Geology and Paleontology 

of Canal Zone and 

Adjoining Parts of Panama

Description of Tertiary Mollusks 

(Gastropods: Eulimidae, Marginellidae to 

Helminthoglyptidae)

GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D
Geology and Paleontology 

of Canal Zone and 

Adjoining Parts of Panama

Description of Tertiary Mollusks 

(Gastropods: Eulimidae, Marginellidae to 

Helminthoglyptidae)

By W. P. WOODRING

GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D

A contribution to the history of 

the Panama land bridge

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1970
UNITED STATES DEPARTMENT OF THE INTERIOR 

WALTER J. HICKEL, Secretary

GEOLOGICAL SURVEY 

William T. Pecora, Director

For sale by the Superintendent of Documents, U.S. Government Printing Office 

Washington, D.C. 20402 - Price 32.25 (paper cover)
CONTENTS

Abstract.._________________________________________

Introduction-______________________________________

Fieldwork and acknowledgments._____________________

Changes in formation and age assignments.____________

Additions to annotated bibliography,_________________

Additions to localities at which fossils were collected_____

New generic and subgeneric names.___________________

Gastropod faunal summaries and age of formations. ____

Eocene series.._________________________________

Gatuncillo formation._______________________

Eocene or Oligocene series.______________________

Marine member of Bohio(?) formation .-_____

Oligocene series__----_-_______________-____-____

Bohio formation.___________________________

Caimito formation._________________________

Caraba formation.__________________________

Panama formation-____-___-________-_-----_

Miocene series________________________________

Culebra formation._________________________

Cucaracha formation..______________________

La Boca formation, including Emperador limestone member.___________________________

Alhajuela formation.._______________________

Gatun formation._____________-_______----_

Chagres sandstone, including Toro limestone 

member_ ________________________________

Holocene series.___________________-____-------_

Informally designated Atlantic and Pacific muck_

Page 

299

299

300

300

301

301

303

303

303

303

304

304

305

305

306

307

308

309

309

311

311

317

317

322

325

325

	Page

Description of Tertiary mollusks continued from chapter C__________________________________________ 326

Gastropods continued from chapter C__________ 326

Family Eulimidae___________________________ 326

Family Marginellidae___-_________-__-_-____ 330

Family Cancellariidae.._____________________ 334

Family Conidae____________________________ 345

Family Turridae___--___-___________________ 360

Family Terebridae__________________________ 403

Family Pyramidellidae____--_-__-____--_____ 414

Family Acteonidae______-__-_-____-_________ 415

Family Ringiculidae________________________ 416

Family Scaphandridae______-____---_______ 417

Family Philinidae_________________________ 420

Family Bullidae---____-_--__---_---_-______ 421

Family Atyidae___________________________ 422

Family Retusidae___---__--------_------___- 423

Family Spiratellidae___-__-__-__-___________- 426

Family Cavolinidae_________________________ 427

Family Ellobiidae_________________________ 429

Family Helminthoglyptidae._________________ 429

Additions and corrections to families covered in

preceding chapters._______________________ 430

References cited____________________________________ 435

Index _________--__-_-_-------__------___------__-- 441

ILLUSTRATIONS

[Plates follow index]

PLATE 48. Middle Eocene mollusk from Gatuncillo formation, late Eocene or early Oligocene mollusks from marine member 

of Bohio (?) formation, late Oligocene mollusks from Bohio and Caimito formations, and early Miocene mollusks 

from La Boca formation.

49. Late Oligocene mollusks from Caimito formation, early Miocene mollusk from La Boca formation, and middle 

Miocene mollusks from Gatun formation.

50. Early Miocene mollusks from La Boca formation and Chipola formation of Florida.

51. Early Miocene mollusk from Emperador limestone member of La Boca formation and middle Miocene mollusks

from Gatun formation.

52-61. Middle Miocene mollusks from Gatun formation.

62, 63. Middle Miocene mollusks from Gatun formation and late Miocene mollusks from Chagres sandstone. 

64, 65. Protoconchs of middle Miocene turrids from Gatun formation.

66. Protoconchs of middle Miocene turrids and terebrids, and pteropod from Gatun formation.

in

GEOLOGY AND PALEONTOLOGY OF CANAL ZONE AND ADJOINING PARTS OF PANAMA

DESCRIPTION OF TERTIARY MOLLUSKS (GASTROPODS: EULIMIDAE, MARGINELLIDAE TO

HELMINTHOGLYPTIDAE)

By W. P. WOODEING

ABSTRACT

The present chapter completes the description of the gastropods in the fossilferous formations, with the exception of some 

20 species in recently acquired collections. It covers 187 species 

and subspecies in 18 families and 13 other unnamed species are 

briefly described or mentioned. This brings the total so far 

covered to 444 and 56, respectively, in the two categories. 

Special attention is devoted to the family Turridae, represented 

by 70 species and subspecies.

Only two of the species were found in the Gatuncillo formation, the middle Eocene part of the formation. A late Eocene or 

early Oligocene age is retained for the marine member of the 

Bohio(?) formation, although Eocene affinities outweigh Oligocene. Perhaps the most striking example of Eocene affinity is 

afforded by Glyptostyla panamensis, which is redescribed and 

reillustrated. It is closely related to an Eocene Nigerian species. 

Averellia stewarti, from the Bohio(?), is the oldest land snail 

from Central America.

The late Oligocene part of the Bohio formation yielded 

Strioterebrum Hstrotum, the first unequivocal Strioterebrum 

from the Oligocene of the Caribbean region. The fossils from 

the Caimito formation, also of late Oligocene age, include 

Vaginella lophota, the first Oligocene American pteropod of the 

genus Vaginella, and the earliest pteropod of the genus Cavolina: C. xenica.

The name Carabra formation is adopted for strata,formerly 

assigned to the Caimito formation. The Panama formation is 

redefined to include strata in the Pacific coastal area formerly 

assigned to the Bohio and Caimito formations. Both the Caraba 

and Panama include marine lenses containing late Oligocene 

larger foraminifera.

Due to the shifting of fossil localities from the Culebra 

formation to the La Boca formation, distribution tables have 

been compiled for all of the gastropods from both of those 

early Miocene formations. Those from the La Boca described in 

the present chapter include Floribella aldrichi, a remarkable 

philinid, and Ellobium aff. E. pellucens, the first American 

fossil representative of the salt-marsh pulmonate genus ElloMum. The Emperador limestone member of the La Boca 

yielded Campanile cf. C. herculeanus. That form and C. herculeanus itself are the last American species of the genus, 

younger than any in Europe. The fine-grained rocks in Madden 

basin, overlying the agglomerate of the pyroclastic-clay member of the Caimito formation of former usage, are now assigned 

to the La Boca.

The early Miocene Alhajuela formation is defined to include 

the youngest strata in Madden basin. It includes a lower member (the calcareous sandstone member of the Caimito formation of former usage) and an upper member (the Alhajuela 

sandstone member of the Caimito formation of former usage).

As in other chapters, the bulk of the fossils 135 species and 

subspecies and two unnamed forms were collected in the 

Gatun formation. They include 18 cancellarids, 16 conids, 50 

turrids, and 13 terebrids. The distribution of the nonendemic 

species of the Gatun indicates that the entire formation is of 

middle Miocene age. The upper part in the western area, west 

of the Canal Zone, formerly was considered to be late Miocene. 

A late Miocene age is adopted for the Chagres sandstone, 

instead of early Pliocene.

According to radiocarbon dates, the informally named Atlantic and Pacific muck, formerly assigned a Pleistocene age, were 

deposited during the post-glacial rise of sea level.

INTRODUCTION

Preparation of chapter D was started in 1962, but 

was interrupted at intervals by other activities. This 

chapter was designed to complete description of the 

gastropods and it does so, with the exception of some 

20 species in recently acquired collections from the 

Gatuncillo, La Boca, and Gatun formations. Six of 

those species of special interest, however, are included 

in the present chapter and additional data, including 

corrections, are presented for three species described in 

preceding chapters.

Chapter D covers 187 species and subspecies and 13 

other unnamed species that are briefly described or 

mentioned. This brings the total so far covered to 444 

and 56, respectively, in the two categories. Two additional chapters are planned to take care of the remaining recently acquired gastropods and also the scaphopods, pelecypods, and cephalopods: an estimated 250 

species.

As shown in the following table, the gastropods so 

far described are unevenly distributed in the marine 

formations.

299
300 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Number 

of

Species of gastropods in marine formations

Formation

Eocene series:

Gatuncillo_ ____________________________________ 33

Eocene or Oligoceiie series:

Marine member of Bohio(?)______________________ 37

Oligocene series:

Bohio_----------______________________________ 29

Caimito--------------__-______________________ 41

Caraba_ _--------______________________________ 2

Miocene series:

Culebra-_-----_---____________________________ 32

La Boca, Gaillard Cut area____________________~_ 78

La Boca, Madden basin.________________________ 6

Alhajuela_ _____________________________________ 5

Gatun_ ________________________________________ 293

Chagres _-----_----____________________________ 38

Though it is improbable that any two formations 

exactly duplicate each other in biofacies, the molluscan 

paleontology would be on a better footing, if the percentage of potential species were as high for all the 

formations as it is for the Gatun. The collections, 

mostly small, from the La Boca formation in Madden 

basin and the overlying Alhajuela formation consist 

chiefly of pelecypods.

Inasmuch as the gastropods make up about twothirds of the molluscan faunas, it is appropriate at this 

time to reappraise the age of the fossiliferous formations. It is appropriate also to correct errors in stratigraphy and areal geology that have come to light since 

publication of chapter A. Other errors are certain to 

turn up as the geology becomes better understood.

FIELDWORK AND ACKNOWLEDGMENTS

Since the project started in 1947 fieldwork was undertaken in the Canal Zone and adjoining parts of 

Panama that year and in 1949, 1954, 1959, 1965, and 

1967. The work in 1965 was specially rewarding so far 

as the La Boca formation is concerned. At that time, 

as a result of the program of widening the Panama 

Canal, excavations on the west side of Las Cascadas 

Reach afforded an opportunity to examine the La Boca, 

including the Emperador limestone member, from the 

base upward through a thickness of 77 meters.

I am specially indebted to R. H. Stewart, geologist 

of the Panama Canal Company, and his assistant Joanne Alien Stewart, Through their official work they 

have uncovered errors in chapter A and through weekend activities they have forwarded the largest collections from the Gatuncillo, La Boca, and Gatun formations. Mr. Stewart also offered advice on subsurface 

geology based on his study of thousands of core holes. 

I have profited from advice from my colleagues A. A. 

Olsson and Druid Wilson, and from members of the 

staff of the Division of Mollusks, of the U.S. National 

Museum.

CHANGES IN FORMATION AND AGE ASSIGNMENTS

The geographic and geologic features mentioned in 

the following summary are shown on plate 1, chapter 

A.

The name Caraba formation, defined on page 307, is 

adopted for strata in the Rio Mandinga area (south of 

Gamboa), on the south side of Madden basin, and 

south of the continental divide east of Madden Highway, formerly assigned to the Caimito formation. Like 

the Caimito formation, it is of Oligocene age.

The Panama formation is redefined on page 308. It 

formerly was considered to be of early Miocene age, 

but now is known to be Oligocene. The strata in the 

Pacific coastal area formerly assigned to the Bohio 

and Caimito formation are reassigned to the Panama.

The agglomerate and tuff in the so-called pyroclasticclay member of the Caimito formation in Madden 

basic (p. 32) are identified by Mr. Stewart as the Las 

Cascadas agglomerate and the overlying fine-grained 

nonvolcanic rocks as the La Boca formation. The finegrained rocks include coralliferous limestone like the 

Emperador limestone member of the La Boca in the 

Gaillard Cut area.

The Alhajuela formation is defined on page 317 to 

include a lower member (the calcareous sandstone 

member of the Caimito formation of page 32) and 

an upper member (the Alhajuela sandstone member) 

of the Caimito formation of page 33).

The upper part of the Gatun formation in the western area is now considered to be of middle Miocene 

age, instead of late Miocene, and the Chagres sandstone 

late Miocene, instead of early Pliocene.

These changes are summarized in the following 

table:

Changes in formation and age assignments

Former usage

Chagres sandstone, early

Pliocene.

Upper part of Gatun formation, western area, late

Miocene. 

Alhajuela sandstone member

of Caimito formation, early

Miocene. 

Calcareous sandstone member

of Caimito formation, early

Miocene. 

Pyroclastic-clay member of

Caimito formation, late

Oligocene. 

Caimito formation, Rio

Mandinga area, late

Oligocene. 

Panama formation, early

Miocene.

Bohio and Caimito formations, Pacific coastal area,

late Oligocene.

Present usage

Chagres sandstone, late 

Miocene.

Upper part of Gatun formation, western area, middle 

Miocene.

Upper member of Alhajuela 

formation, early Miocene.

Lower member of Alhajuela 

formation, early Miocene.

Las Cascadas agglomerate, 

Oligocene(?) and La Boca 

formation, early Miocene.

Caraba formation, late 

Oligocene.

Panama formation, late

Oligocene. 

PanamA formation, late

Oligocene.
GASTROPODS: ETJLIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 301

ADDITIONS TO ANNOTATED BIBLIOGRAPHY

The first three items should have been included in 

former lists.

1862. Blake, C. C., Sharks' teeth at Panama: Geologist, v. 5, 

p. 316.

Three species of shark teeth, including a new species, Lamna eurybathrodon, from Miocene deposits 

at Monkey Hill [upper part of Gatun formation at 

Mount Hope]. According to the author, "it would 

be very injudicious to found a species on one solitary tooth." Nevertheless he did it.

1889. Woodward, A. S., Catalogue of the fossil fishes in the 

British Museum (Natural History), pt. 1, 474 p., 

17 pis.

Blake's species is briefly described on p. 438.

1955. White, E. I., On Lamna eurybathrodon Blake: Annals 

and Mag. Nat. History, 12th ser., v. 8, p. 191-193, 

14 figs.

Blake's species is assigned to Negaprion, redescribed, and recorded also from a Miocene Australian 

locality.

1960. Woodring, W. P., Panama : XX Cong. Geol. Internat, Com. 

Stratigraphique, Lexique Stratigraphique International, v. 5, Amerique Latine, fasc. 2a, Amerique Central, p. 307-357, map.

Alphabetical list of valid, dubious, and nude stratigraphic names.

1962. Tuan, Yi-Fu, A coastal reconnaissance of central Panama : California Geographer, v. 3, p. 79-96, 12 figs. 

Three USGS Cu dates on muck, erroneously considered to be of Pleistocene age on page 50 of chapter A, are cited in a footnote on page 94.

1962. Eames, F. E., and others, Fundamentals of mid-Tertiary 

stratigraphical correlation, 163 p., 17 pis., 20 figs., 

Cambridge Univ. Press.

Gatuncillo, Bohio, and Caimito formations are of 

Aquitanian (early Miocene) age (p. 36-37).

1963. Yokes, E. H., Cenozoic Muricidse of the western Atlantic 

region; pt. 1, Murex sensu stricto: Tulane Studies 

Geology, v. 1, no. 3, p. 93-123, 4 pis. 

Includes species of Murex from Gatun formation.

1964. Yokes, E. H., Additions to the New World Turbinellas: 

Tulane Studies Geology, v. 3, no. 1, p. 95-96. 

Includes discussion of species from Canal Zone.

1964. Jenkins, D. G., Panama and Trinidad Oligocene rocks: 

Jour. Paleontology, v. 38, p. 606.

Foraminifera from Gatuncillo formation at locality 

21 of present report are of early Oligocene age.

1965. Eames, F. E., and others, Dating of some beds in Panama 

and Trinidad: Idem, v. 39, p. 162-163.

Fossil bed at locality 21 is of Aquitanian (early 

Miocene) age.

1965. Whitmore, F. C., Jr., and Stewart, R. H., Miocene mammals and Central American seaways: Science, v. 148, 

p. 180-185, 2 figs.

North American mammals, considered to be of 

middle Miocene age, from Cucaracha formation.

1965. Yokes, E. H., Cenozoic Muricidse of the western Atlantic 

region; pt. 2, Chicoreus sensu stricto and Chicoreus 

(Siratus): Tulane Studies Geology, v. 3, no. 4, p. 

181-208, 4 pis.

Includes a species from Gatun formation.

1966. Woodring, W. P., Estratigrafia Terciaria de la Zona del 

Canal y partes adyacentes de la Repiiblica de Panama (Tertiary stratigraphy of Panama Canal Zone 

and adjoining parts of the Republic of Panama) : 

Inst. Centroamericano In. y Tecnologia Indus., Pub. 

Geol. no. 1, p. 43-45.

Includes correction of formation assignments in 

Pacific coastal area.

1967. Stewart, R. H., The quartz minerals of Panama and the

Canal Zone: Lapidary Jour., v. 21, no. 1, p. 185-190;

no. 2, p. 324-333, illus. 

Includes localities where silicified corals and wood

can be collected. 

1967. Bold, W. A. van den, Miocene Ostracoda from Costa

Rica: Micropaleontology, v. 13, no. 1, p. 75-86, 2 pis.,

1 fig.

Includes a species from Caimito formation and

one from Gatun formation. 

1967. Bold, W. A. van den, Ostracoda of the Gatun formation,

Panama: Idem, v. 13, no. 3, p. 306-318, 2j>ls.

Twenty-eight species. 

1969. Blacut, Gustavo, and Kleinpell, R. M., A stratigraphic

sequence of benthonic smaller foraminifera from the

La Boca formation, Panama Canal Zone: Cushman

Found. Foram. Research Contr., v. 20, pt. 1, p. 1-22,

pis. 1-6, 4 figs.

Foraminifera from a measured section on west

side of Las Cascadas Reach and a nearby core hole. 

1969. Bartlett, A. S., Barghoorn, E. S., and Berger, Rainer,

Fossil maize from Panama: Science, v. 165, p. 389-

390, 3 figs. 

Wild maize pollen and at higher level cultivated

maize in radiocarbon dated muck in Gatun Lake

area.

ADDITIONS TO LOCALITIES AT WHICH FOSSILS 

WERE COLLECTED

No. 

used in 

this 

report

USGS 

Cenozoic 

No.

Description of locality

Gatuncillo formation

23b 

37a

24553 

23648

Upper course of Rio Palenque, 3.4 km in 

direct line west of Nuevo San Juan and 1.3 

km northwest of settlement of Palenque, 

Col6n Province, Panama. Silty mudstone. 

R. H. Stewart, J. L. Alien, and Anselmo 

Mena, 1968. 

Top of Cerro Pelado, 1 km north-northwest of 

Gamboa, Canal Zone, altitude 223 m. 

Leached, soft sandstone. R. H. Stewart, 

1964.
302 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

No. 

used in 

this 

report

112b 

112c

I22c

lOla

lOlb

lOlc 

lOld

lOle

lOlf 

lOlg 

lOlh

lOli

USGS 

Cenozoic 

No.

24502 

24505

23650

23651

23656 

23668

23655

23653 

23654 

23652

23658

Description of locality

Cnlebra formation

Panama Canal, east side of Culebra Reach, 

northwest face of Gold Hill, near Core Hole 

ECB1, Canal Zone. Cliff face of sandstone. 

R. H. Stewart, 1967. 

Panama Canal, east side of Culebra Reach, 

opposite Empire-Culebra slide, about Canal 

station 1730, Canal Zone. M. I. Goldman, 

1912.

Cucaracba formation

northeast slope of Cerro Escobar near 

Borinquin Highway, Canal Zone. Somewhat 

carbonaceous shale (coprolite bed) about 3 m 

above ash flow. J. L. Alien, 1965.

La Boca formation

Panama Canal, west side of Las Cascadas 

Reach, Canal station 1627, Canal Zone. 

Basal echinoid-bearing limestone. R. H. 

Stewart and W. P. Woodring, 1965.

Panama Canal, west side of Las Casoadas 

Reach, Canal station 1631 plus 15 m, Canal

Zone. Basal echinoid-bearing limestone. 

W. P. Woodring, 1965. 

Panama Canal, west side of Las Cascadas 

Reach, Canal station 1622, Canal Zone. Unit 

4 of measured section on p. 312. W. P. 

Woodring, 1965. 

Panama Canal, west side of Las Cascadas 

Reach, Canal station 1622, Canal Zone. 

Unit 5 of measured section on p. 312. W. P.

Woodring, 1965. 

Panama Canal, west side of Las Cascadas 

Reach, Canal station 1628, Canal Zone.

Poorly sorted, somewhat calcareous sandstone about 15 m below Emperador limestone member. W. P. Woodring, 1965. 

Panama Canal, west side of Las Cascadas 

Reach, Canal station 1602, Canal Zone. 

Limestone at top of canal cliff. W. P. 

Woodring, 1965. 

Panama Canal, west side of Las Cascadas 

Reach, Canal station 1601 plus 15 m, Canal 

Zone. Upper limestone in upper part of La 

Boca formation. W. P. Woodring, 1965. 

Panama Canal, west side of Las Cascadas 

Reach, Canal stations 1608 to 1612 plus 23 

meters, near top of canal cliff, Canal Zone.

Somewhat calcareous silty sandstone, about 

25 meters above Emperador limestone 

member. W. P. Woodring, R. H. Stewart

and J. L. Alien, 1965. 

Panama Canal, east side of Las Cascadas 

Reach, opposite measured section at Canal 

station 1622, Canal Zone. Upper part of La

Boca formation, corresponding to unit 12 of 

measured section on p. 312. R. H. Stewart, 

1964.

No. 

Used in 

this 

report

H7a 

Il7b 

H7c

117d

118

138f 

138g

143a

159d 

160b

160c 

160d

I77e

usos

Cenozoic 

No.

23662 

23661 

23660

23657

23659

23663 

23664

24504

24173 

23665

23666 

24503

24174

Description of locality

Emperador limestone member of La Boca formation

Panama Canal, west side of Las Cascadas 

Reach, Canal station 1630, Canal Zone. Two 

beds of coralliferous limestone. R. H. Stewart 

and W. P. Woodring, 1965. 

Panama Canal, west side of Las Cascadas 

Reach, Canal station 1630, Canal Zone. 

Upper noncoralliferous limestone. W. P. 

Woodring, 1965. 

Panama Canal, east side of Las Cascadas

Reach, Canal station 1626, Canal Zone. 

Coralliferous limestone. W. P. Woodring,

1965. 

Panama Canal, west side of Las Cascadas

Reach, Canal station 1622, Canal Zone. 

Unit 7b of measured section on p. 312. 

Coralliferous limestone. W. P. Woodring, 

1965. 

West of Panama Canal, overgrown quarry

about 250 m west of Borinquin Highway, in 

line with Canal station 1870, Empire Reach,

Canal Zone. Same as USGS 6016. Type 

locality of Emperador limestone member. 

Incorrectly plotted on pi. 2, chap. A. R. H. 

Stewart, 1960.

Lower part of Gatun formation

South side of Transisthmian Highway, hillside 

excavation at Colchoneria Yero, about 450 m 

southwest of Cativa, Panama. R. H. Stewart, 

W. P. Woodring, and others, 1965. 

South side of Transisthmian Highway, about 35 

m east of 23663, Panama. R. H. Stewart, 

1965.

Middle part of Gatun formation

Panama Railroad, first cut southeast of Camp 

Totten, Canal Zone. [Near locality 143.] 

M. I. Goldman, 1912. 

Gatun, Canal Zone. [Presumably Gatun Locks 

excavation.] M. I. Goldman and others, 1912. 

South side of Rfo Chagres, 175 m below spillway of Gatun Dam, Canal Zone. Coralliferous conglomerate. W. P. Woodring, 1965. 

Same locality as 23665. Silty sandstone underlying conglomerate. W. P. Woodring, 1965. 

West end of Gatun Dam, Canal Zone. M. I. 

Goldman, 1912.

Upper part of Gatun formation

Mount Hope borrow pits, Canal Zone. M. I. 

Goldman, 1912.
GASTROPODS: EULIMIDAE. MARGINELLIDAE TO HELMINTHOGLYPTIDAE 303

NEW GENERIC AND SUBGENERIC NAMES

The following new generic and subgeneric names are

proposed.

Dolostoma, Turridae, Mangeliinae.

Type: Dolostoma anorhepes Woodring, n. sp., 

Gatun formation, Miocene, p. 398. Gender neuter.

Euglyphostoma, 'subgenus of Glyphostoma, Turridae,

Mangeliinae.

Type: Glyphostoma partefilosa Dall, Gulf of California and west coast of Baja California, p. 401. 

Gender neuter.

Floribetta, Pliilinidae.

Type: Dolabella aldrichi Dall, Miocene, Florida, 

Canal Zone, Cuba, p. 420. Gender feminine.

Lecattia, subgenus of AverelUa, Helminthoglyptidse,

Xanthonycinae.

Type: AverelMa (Lecallia) stewarti Woodring, n. 

sp., marine member of Bohio(?) formation, late 

Eocene or early Oligocene, p. 429. Gender neuter.

Paleocavolina, subgenus of Cavolina, Cavolinidae. 

Type: Cavolina (Paleocavolina) xenica Woodring, 

n. sp., Caimito formation, Oligocene, p. 428. Gender feminine.

Rhiglypliostoma, subgenus of <Glypho8toma, Turridae,

Mangeliinae.

Type: Glyphostoma xeston Gardner, Miocene, 

Florida, p. 401. Gender neuter.

GASTROPOD FAUNAL SUMMARIES AND AGE OF 

FORMATIONS

EOCENE SERIES 

GATUNCILLO FORMATION

As a result of Mr. Stewart's fieldwork and the coredrilling under his supervision the distribution of the 

Gatuncillo formation has been enlarged. Leached fossiliferous sandstone of that formation was found capping Cerro Pelado on the 227-meter peak (223 meters 

on the most recent map) of plate 1, a kilometer northnorthwest of Gamboa (locality 37a). ECB core holes, 

drilled near the northwest foot of Gold Hill a few 

hundred meters northwest of the continental divide on 

the east side of the canal, revealed siltstone and mudstone of the Gatuncillo, containing abundant planktonic foraminifera, underlying the Culebra formation. 

Ruth Todd reports that the poorly preserved foraminifera include Cassigerinella, indicating a latest Eocene 

or early Oligocene age. Another subsurface locality 

(Core Hole CH-5), in the northern part of Gatun 

Lake, is recorded on page 61. These scattered occurrences and the reappearance of late Eocene deposits in 

the Rio Bayano district, east of the area covered by

plate 1, indicate that probably all of central Panama 

was submerged in late Eocene time.

Flat-lying limestone of the Gatuncillo, resting on an 

irregular surface of the basement complex, was found 

along and near a tributary of Rio Agua Sucia a kilometer west of locality 27 on the Transisthmian Highway, in an area shown as basement on plate 1. The 

limestone has a thickness of almost 60 meters, the 

greatest thickness now known for limestone in the 

Gatuncillo. It contains caves and deep, narrow sink 

holes. The stream disappears in the limestone and 

emerges at the contact with basement. The Agua Sucia 

and Rio Gatun faults of plate 1 are misinterpretations.

The Gatuncillo formation has yielded 33 species of 

gastropods so far recorded, including two in the present 

chapter: Persicula cf. P. semen and Conus cf. C. sauridens, both silicified fossils from the Rio Casaya area 

(locality 38). Fifteen of them, however, are not named 

at the specific level. Only four are unequivocally identified, three of which are endemic. About 20 additional 

species are represented in the recently acquired collection from locality 23b, for the most part species of 

genera not found at other localities.

The occurrence elsewhere of the same or related species is as follows:

Gastropods from Gatuncillo formation and occurrence elsewhere of 

same or related species

Species from Gatuncillo formation

Velates perversus (Gmelin), 

subsp.?

Turritella cf. T. carinata Lea.__

Turritella cf. T. samanensis

Olsson. 

Dirocerithium ame Woodring-__

Hannatoma cf. H. emendorferi 

Olsson. i

Calyptraea cf. C. aperta 

(Solander).

Oostrombus aff. 0. chiraensis 

(Olsson).

Terebellum (Terebellum) 

procerum Merian. 2

Terebellum (Seraphs)

belemnitum (Palmer)?. 

Ectinochilus cf. E. gaudichaudi

(d'Orbigny). 

Cypraedia aff. C. subelegans

(Trechmann). 

Pachycrommiumf solenaeum

Woodring.

See footnotes at end of table.

Occurence elsewhere of same or 

related species

V. perversus, early to late

Eocene, principally Tethyan

localities. 

T. carinata, middle Eocene,

southeastern U.S. 

T. samanensis, late Eocene,

Peril. 

D. whitfieldi (Heilprin),

middle Eocene, southeastern U.S. 

H. emendorferi, late Eocene, or

early Oligocene, Peru; late

Eocene, Columbia,

Venezuela. 

C. aperta, Paleocene to

Oligocene, western Europe,

southeastern U.S., Miocene,

Maryland. 

0. chiraensis, late Eocene or

early Oligocene, Peru; 0.

tournoueri (Bayan), middle

Eocene, Italy. 

Middle Eocene, Jamaica,

Haiti, St. Bartholomew;

late Eocene, Trinidad. 

T. belemnitum, late Eocene,

Florida. 

E. gaudichaudi, late Eocene,

Peru. 

C. subelegans, middle Eocene,

Jamaica.
304 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Gastropods from Gatuncillo formation and occurrence elsewhere of 

same or related species Continued

Species from Gatuncillo formation

Galeodea? cf. G. nodosa

(Solander).

Yasila aff. Y. paytensis Olsson_ 

Xancus cf. X. peruvianus

(Olsson).

Ctenilyria ctenista Woodring

Persicula (Gibberula) cf. P.

semen (Lea). 

Conus cf. C. sauridens Conrad-.

Occurrence elsewhere of same or 

related species

G. nodosa, middle to late 

Eocene, western Europe.

Y. paytensis, late Eocene, Peru.

X. peruvianus, middle Eocene 

to late Eocene or early 

Oligocene, Peru.

C. coroni (Morlet), middle to 

to late Eocene, western 

Europe.

P. semen, middle to late 

Eocene, soiitheastern U.S.

C. sauridens, middle Eocene to 

early Oligocene, southeastern U.S.

1 See following paragraph.

2 Described on p. 192 as Terebellum procerumf.

Hannatoma cf. H. emendorferi. was described on 

page 68 as Hannatoma? cf. H. emendorferi. Hundreds 

of specimens of this species from the new locality 23b 

show that it is indeed a Hannatoma, to be described in 

chapter E as a new species. Hannatoma is classified as 

a cerithid, not a thiarid. That it is a brackish genus is 

doubtful.

The Gatuncillo gastropods have Tethyan (Velates, 

Terebellum and its subgenus Seraphs], western European, southeastern United States, West Indian, and 

Peruvian affinities. Those with southeastern United 

States and Peru are preponderant. Velates, Dirocerithium, Hannatoma, Oostrombus, Seraphs, Ectinocliilus, 

CyprcBdia, Pachycrommium, Yasila, and Ctenilyria are 

extinct, and Terebellum survives only in the western 

Pacific Ocean.

Age. No reasons are apparent for changing the age 

assignment in chapter A (p. 22), that is, that the bulk 

of the formation is of late Eocene age, but that it includes middle Eocene. Smaller foraminifera from 

locality 21 have been claimed to be of early Oligocene 

age (Jenkins, 1964), although at locality 23, practically 

along the strike, limestone yielded late Eocene larger 

foraminifera (p. 20). The Gatuncillo may include deposits of early Oligocene age, but the way to determine 

that is to examine a series of samples by trenching or 

augur-holing the upper part of the formation. With 

some effort that can be done. The Gatuncillo has been 

alleged to be of Aquitanian (early Miocene) age, with 

a liberal sprinkling here and there of reworked middle 

and late Eocene fossils (Eames and others, 1962, p. 36). 

As a matter of fact, the basis for that allegation the 

occurrence of Lepidocyclina pustulosa tobleri (Pliolepidina of Eames and others) has been repudiated 

(Eames and others, 1968, p. 302).

EOCENE OB OLIGOCENE SERIES

MARINE MEMBER OF BOHIO(?) FORMATION

The term "marine member of Bohio (?) formation" 

is a poor designation for this unit. Hill's casual designation "Vamos a Vamos [Vamos Vamos] beds" cannot 

be adopted, as it cannot be defined properly. His locality, on Rio Chagres northwest of what is now Barro 

Colorado Island (locality 40), is now under the waters 

of Gatun Lake and the stratigraphic relations were 

uncertain before it was submerged. (See discussion p. 

22-23.) If mapping can be carried out in the peninsula 

ending in Palenquilla Point (locality 41) and the 

peninsula to the west, it may be possible to define properly the strata containing this distinctive fauna.

Species in the families covered in chapter D are 

tabulated below. In that and other distribution tables 

the designation "sp." in the locality columns indicates 

an incomplete or poorly preserved species that may or 

may not be the same as that in the species column, and 

the designation "?sp." indicates that the species is questioned. Symbols for relative frequency are as follows:

Symbols used for relative frequency

Symbol 

R, rare.

Number

of

specimens

1-2

F, few.- _ - -- - ---- - 3-5

C, common----_---__------------------------------ 6-20

A, abundant.- -- >20

Gastropods from marine member of Bohio (?) formation (Eulimidae, 

Turridae to Helminthoglyptidae 

[R, rare; F, few; C, common]

Eulima cf . E. jacksonensis

Niso (Niso) umbilicata (Lea)?___-

Scaphander (Scaphander) cf. S.

Retusa (Cylichnina) aff. R.

Volvulella (Volvulella) aff. V.

Averellia (Lecallia) stewarti

Localities

Vamos 

Vamos

40a

R

R 

F

40d

R 

R

C

R

Palenquilla 

Point

41

F

R

41b

R

C 

R

R

Trinidad 

Island

42

F

?sp. R

If Zemaciesf sp. a is indeed a species of Zemacies, it 

is a representative of a Paleocene to Pliocene New 

Zealand-Australian genus heretofore unknown in 

America. Averellia stewarti, the oldest land snail in 

Central America, is associated with 40 marine species
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 305

at locality 41b. It is the type of the subgenus Lecallia. 

Thirty-seven species are so far recorded from the 

marine member of the Bohio (?) formation a representation comparable to that for the Gatuncillo formation. Half of them are not named at the specific level. 

The seven unequivocally identified species are endemic. 

The affinities of those identified at the specific level and 

of one not so identified are as follows:

Gastropods from marine member of Bohio(?) formation and occurrence elsewhere of related species

Species from marine member of 

Bohio(?) formation

Tricolia calypta Woodring-____ 

Turritella adela Woodring___ __

Turritella cf. T. caleta Olsson L_ 

Architectonica (Stellaxis) aff.

A. alveata (Conrad). 

Architectonica cf. A. fungia

(Conrad). 

Eulima cf. E. jacksonensis

Gregorio.

Niso (Niso) umbilicata (Lea)?._

Neverita (Glossaulax) bolivar ensis tapina Woodring.

Globularia (Ampulella) sp______

Globularia (Ampulella?) nana

Woodring. 

Pachycrommiumf proinum

Woodring.

Morum (Cancellomorum) cf.

M. antiquum (Bayan) 2 . 

Ficus cf. F. mississippiensis

Conrad. 

Typhis (Laevityphis) aff. T.

recurvirostris Conrad. 

Glyptostyla panamensis DalL

Scaphander (Scaphander) cf.

S. jacksonensis Palmer. 

Retusa (Cylichnina) aff. R.

adamsi (Palmer). 

Volvulella (Volvulella) aff. V.

conradiana (Gabb). 

Averellia (Lecallia) stewarti

Woodring, n. sp.

Occurrence elsewhere of related species

T. precursor (Dall), early

Miocene, Florida. 

T. galvesia Olsson, Oligocene,

Peru.

T. caleta, Oligocene, Peru. 

A. alveata, middle to late

Eocene, southeastern U.S. 

A. fungia, middle Eocene,

southeastern U.S. 

E. jacksonensis, middle to

late Eocene, southeastern

U.S. 

N. umbilicata, early to late

Eocene, southeastern U.S. 

N. bolivarensis bolivarensis

Clark, late Eocene,

Colombia. 

G. parisiensis (d'Orbigny),

middle Eocene to early

Oligocene, western Europe. 

G. garzaensis Vokes, middle

Eocene, California. 

P.? gabrielensis (Clark), late

Eocene, Colombia; P.?

jacksonensis (Harris), late

Eocene, southeastern U.S. 

M. antiquum, late Eocene,

Italy. 

F. mississippiensis, Oligocene,

southeastern U.S. 

T. recurvirostris, Oligocene,

southeastern U.S. 

G. striata (Newton), middle

or late Eocene, Nigeria. 

S. jacksonensis, late Eocene,

southeastern U.S. 

R. adamsi, middle Eocene,

southeastern U.S. 

V. conradiana, middle Eocene,

southeastern U.S. 

A. coactiliata (Ferrusac),

Holocene, northern Mexico

to Trinidad.

1 Identified as Turritella olssoni Clark, a late Eocene Colombian species, by Allison 

(in Allison and Adegoke, 1969, p. 1254).

2 Described on page 203 as Morum ("Oniscidia") cf. M. antiquum.

Age. The gastropods from the marine member of 

the Bohio (?) formation have Eocene and Oligocene 

affinities, but the Eocene outweigh the Oligocene. Perhaps the most striking example of Eocene affinity is 

afforded by Glyptostyla panamensis, which is redescribed on page 434 and reillustrated on plate 48, 

figure 23. As a result of reappraisal, Glyptostyla is

monotypic so far as America is concerned. A closely 

related species, however, G. striata (described as a species of Strepsidura] , occurs in the Eocene of Nigeria. 

Newton (1922, p. 109) favored a middle Eocene age 

for the Nigerian deposits, but Eames (1957, p. 30) preferred late Eocene. Of the genera and subgenera in the 

preceding lists, Stellaxis, Globularia, Pachycrommium, 

Glyptostyla, Zemaciesf, and Lecallia are extinct.

If the Eocene trend is continued by the pelecypods, 

a late Eocene age would be justified, presumably latest 

Eocene, younger than any known molluscan fauna in 

the Gatuncillo formation. It is possible that this 

Bohio (?) fauna represents an unusual biof acies in the 

upper part of the Gatuncillo. Stratigraphic relations, 

however, are unknown. A late Eocene age would agree 

with Cole's identification of the larger foraminifera, 

quoted on page 23.

OLIGOCENE SERIES 

BOHIO FORMATION

The only mollusks from the Bohio formation are 

those in collections from the upper part of the formation on Barro Colorado Island. The following species, 

all collected at locality 42d, are covered in the present 

chapter:

Gastropods from upper part of Bohio formation on Barro Colorado 

Island (Marginellidae to Atyidae)

[R, rare; C, common; A, abundant] Locality

Marginella (Eratoidea) aff. M. mollitor Dall___ 

Conus cf. C. sulculus DalL_-__---------------

Conus aff. C. chipolanus Dall_----_-----------

Drilliaf (Neodrilliaf) sp____________________

Strioterebrum listrotum Woodring, n. sp--------

Acteon (Acteon) aff. A. tampae DalL___________

Acteocina cf. A. bullata (Kiener)____--__------

Scaphander (Scaphander) cryus Woodring, n. sp. 

Atys (Roxaniella) rhadina Woodring, n. sp_____

____ R 

	R 

.____ C

.___- R

__-_ R

.____ R

.____ A

__._ C

,___- R

Strioterebrum listrotum is the first unequivocal 

Strioterebrum, from the Oligocene of the Caribbean 

region.

Of the 29 species from the upper part of the Bohio 

formation 10 are not named specifically. Seven of the 

eight unequivocally named species are endemic. The 

occurrence elsewhere and in other Canal Zone formations of the same or related species is tabulated below.

Age. According to the tabulation, the Bohio fossils 

have Oligocene and Miocene affinities. The only species 

that occurs elsewhere (Atys rhadina) is found in the 

Oligocene Mint Spring marl member of the Marianna 

limestone in southeastern United States, which is generally assigned to the middle Oligocene. On the face 

of the table Miocene affinities far outweigh Oligocene. 

In the Caribbean region, however, Oligocene molluscan faunas are meager compared with those of Miocene 

age and for the most part consist of large, robust forms.
306 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Gastropods from upper part of Bohio Formation on Barro Colorado 

Island and occurrence elsewhere and in other Canal Zone formations of same or related species

Species from Bohio formation

Solariella ephnidia Woodring. __

Turritella cf. T. altilira 

Conrad.

Turritella listrota Woodring____

Hemisinus (Longiverena)

oeciscus Woodring. 

Cerithium (Thericium) mimeticum Woodring. 

Orthaulax sp 1 ______________.

Globularia (Globularia) aff. 

G. fischeri (Dall).

Pachycrommium aff. P. guppyi 

(Gabb).

Gonysycon epomis Woodring. 

Gonysycon cf. G. epomis

Woodring. 

Mitrella epacta Woodring. _ _

Marginella (Eratoidea) aff. 

M. mollitor Dall.

Conus cf. C. sulculus Dall____ 

ConUs aff. C. chipolanus DalL.

Strioterebrum listrotum

Woodring, n. sp. 

Acteon (Acteon) aff. A. tampae

Dall. 

Acteocina cf. A. bullata

(Kiener). 

Scaphander (Scaphander) cryus

Woodring, n. sp. 

Atys (Roxaniella) rhadina

Woodring, n. sp.

Occurrence elsewhere and in other

Canal Zone formations of same or

related species

S. altiuscula Guppy, middle

Miocene, Jamaica; S.

depressa Dali, Holocene. 

Subspecies of T. altilira range

from late Oligocene to late

Miocene. 

T. venezuelana Hodson, early

Miocene, Venezuela, Canal

Zone. 

H. atriformis Cooke, late

Oligocene, Antigua.

Possibly related to 0. pugnax 

(Heilprin), late Oligocene, 

Georgia, early Miocene, 

Florida.

G. fischeri, early Miocene, 

Florida. Also Caimito and 

La Boca formations.

P. guppyi, middle Miocene, 

Jamaica; early to middle 

Miocene, Dominican Republic. Also Culebra and 

La Boca formations.

M. acanthodes (Dall), early

Miocene, Florida; M. oxia

Gardner, early Miocene,

Florida. 

M. mollitor, early Miocene,

Florida; M. euancyla

Gardner, early Miocene,

Florida. Also Caimito

formation. 

C. sulculus, early Miocene,

Florida. Also Caimito

formation. 

C. chipolanus, early Miocene,

Florida. Also Caimito,

Culebra, and La Boca

formations. 

S. ischnum Woodring, middle

Miocene, Jamaica. 

A. tampae, early Miocene,

Florida. 

A. bullata, early Miocene to

Holocene. 

Doubtfully also La Boca

formation. 

Middle Oligocene, Mississippi.

1 Described on page 191 as Orthaulax cf. 0. pugnax (Heilprin).

The Suwannee limestone of Florida, of late Oligocene 

age, offers a more suitable biofacies, but its fauna 

shows little affinity with that from the Bohio formation. Orthaulax, Globularia, Pachycrommium, and 

Gonysycon are extinct.

The larger foraminifera from locality 42d (Cole, 

1957) and the stratigraphic position below the Caimito 

formation are decisive for a late Oligocene age.

CAIMITO FORMATION

The Caimito formation includes two distinct biofacies : a shallow-water facies on the Panama Railroad 

north and south of the Bohio Peninsula (localities 56, 

57, 57a) and in the Quebrancha syncline (locality 62); 

and a moderately deep-water facies on Barro Colorado 

Island (localities 54g to 54n) and Pato Horqueto 

Island (locality 55a). Locality 55b is a fossiliferous 

conglomerate interbedded with the moderately deepwater tuffaceous siltstone of locality 55a. The conglomerate evidently is a gravity slide, as its fossils are 

shallow-water forms. It therefore is grouped with' the 

shallow-water facies. The two facies are so different 

that separate tables are presented for them. No species 

is found in both; in fact, only two genera (Architectonica and Conus) occur in both. (The Architectonica 

of the moderately deep-water facies is an unidentified 

species.) Chapter D covers the species in the next two 

tables.

Gastropods from shallow-water facies of Caimito formation (Marginellidae to Turridae)

[R, rare; F, few; C, common]

Localities

55b 56 57

Marginella (Eratoidea) aff. M. mollitor DalL __________ R ___.

Conus sp___________________________________ R _______

Conus cf. C. sulculus Dall_______-_-_-___--______--- C R

Conus aff. C. chipolanus DalL ______________________ R ___.

Gemmula cf. G. arnica Casey________________________ F ___.

Pleurofusia sp------------------------------------ R -__.

Gastropods from moderately deep-water facies of Caimito formation 

(Conidae to Cavolinidae)

[R, rare; C, common]

Conus cf. C.peruvianus Olsson.

Srobinella aff. S. morierei

Paraborsonia aff. P. brassoenVaginella lophota Woodring, n.

Cavolina (Paleocavolina)

Localities

54g

R

R

54h

R

R

R

R

54j

R

C

R

54k

R

R

R

541

R

54m

R

54n

R

R

55a

R 

R

R

The turrids and pteropods in the preceding list are 

noteworthy. Cavolina xenica, the oldest species of the 

genus, is the type of the subgenus Paleocavolina.

Forty-one species of gastropods are recorded from 

the Caimito formation. Half of them are not named 

at the specific level. Five of the seven unequivocally 

identified species are endemic. The next two tables show
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 307

the distribution elsewhere and in other Canal Zone 

formations of the same or related species.

Gastropods from shallow-water fades of Caimito formation and 

occurrence elsewhere and in other Canal Zone formations of the 

same or related species

Species from. Caimito formation

Turritella meroensis Olsson_

Turritella (Bactrospira) 

altilira Conrad, subsp.1

Architectonica (Architectonica)

rhicna Woodring. 

Trochita cf. T. spirata Forbes 2 .

Orthaulax sp 3 __--_________-_.

Cypraea cf. C. chilona Dall____

Globularia (Globularia) aff. 

G. fischeri (Ball).

Ampullinopsis spenceri 

(Cooke).

Semicassis (Echinophoria)

apenes Woodring. 

Semicassis (Echinophoria) sp.

Cymatium (Septa) ogygium

Woodring. 

Ficus cf. F. pilsbryi (B.

Smith).

Marginella (Eratoidea) aff. 

M. mollitor Dall.

Conus cf. C. sulculus Dall. 

Conus aff. C. chipolanus--

Gemmula cf. G. arnica Casey__

Occurrence elsewhere and in other Canal 

Zone formations of same or related 

species

Late Oligocene, Santiago

area, Panama, Ecuador,

Peru. 

Subspecies of T. altilira

range from late Oligocene

to late Miocene. 

A. nobilis Roding, early

Miocene to Holocene. 

T. spirata, possibly middle

Miocene to Holocene. 

Possibly related to 0. pugnax

(Heilprin), late Oligocene,

Georgia, early Miocene,

Florida. 

C. chilona, early Miocene,

Florida. Also Culebra and

La Boca Formations. 

G. fischeri, early Miocene,

Florida. Also Bohio and

La Boca Formations. 

Late Oligocene, Antigua;

possibly Puerto Rico,

Santiago area, Panama,

Ecuador, Peril.

S. intermedia (Brocchi) 

Miocene to Pliocene, 

Italy.

C. nicobaricum (Roding), 

Holocene.

F. pilsbryi, middle Miocene, 

Jamaica, Dominican 

Republic.

M. mollitor, early Miocene, 

Florida; M. euancycla 

Gardner, early Miocene, 

Florida. Also Bohio formation.

C. sulculus, early Miocene, 

Florida. Also Bohio 

formation.

C. chipolanus, early Miocene, 

Florida. Also Bohio, 

Culebra, and La Boca 

formations.

G. arnica, early Oligocene, 

Mississippi.

1 Described on page 104 as Turritella (Torcula) altilira Conrad, subsp.

2 Recorded on page 81 as Trochita cf. T. trochiformis (Born). 

s Described on page 191 as Orthaulax cf. 0. pugnax (Heilprin).

Age. The Caimito gastropods, like those from the 

Bohio formation, have Oligocene and Miocene affinities. Numerically Miocene outweighs Oligocene. Nevertheless Oligocene affinities are more pronounced for 

the Caimito than for the underlying Bohio and the 

percentage of extinct genera is almost twice as high 

for the Caimito. The two unequivocally identified species that are not endemic (Turritella meroensis and 

Ampullinopsis spenceri) occur elsewhere in deposits of

Gastropods from moderately deepand occurrence elsewhere and 

of related species

 water fades of Caimito formation 

in other Canal Zone formations

Species from Caimito formation

Conus cf. C. peruvianus 

Olsson. 

Scobinella aff. S. morierei 

(Cossmann). 

Paraborsonia aff. P. brassoensis 

(Mansfield).

Vaginella lophota Woodring, 

n. sp.

Woodring, n. sp.

Occurrence elsewhere and in other Canal 

Zone formations of related species

C. peruvianus, late Eocene, 

Peril. 

S. morierei, early to late 

Miocene. 1 

P. brassoensis, middle Miocene, 

Trinidad. 

S. listrotum Woodring, Bohio

formation. 

V. depressa Daudin, Miocene, 

western Europe ; V. chipolana 

Dall, early Miocene, 

Florida.

i See page 373 for localities.

Oligocene age. Though Ampullinopsis (Megatylotus 

of much European literature) has an age range of late 

Eocene to early Miocene, it is especially characteristic 

of Oligocene throughout the Tethyan region, in the 

Rupelian (or Stampian) of western Europe, in southeastern United States, and in the Tertiary Caribbean 

province. The following genera and subgeiiera are extinct : Orthaulax, Globularia, Ampullinopsis, Echinophoria, "Gemmula" Zemaciesf, Scobinella, Paraborsonia, and Paleocavolina.

The beautifully preserved planktonic and benthonic 

foraminifera in the moderately deep-water facies of 

the Caimito on Barro Colorado Island were identified 

by Bolli and assigned by him to the Globorotalia kugleri zone (Bolli, in Woodring, 1958, p. 22-23, 27). The 

smaller foraminifera, larger foraminifera (p. 29-30; 

Cole, 1957), and inollusks indicate a late Oligocene age.

As part of the sweeping allegation that no marine 

Oligocene is known in America, except in the Tampico 

area of Mexico and Cuba, the Bohio and Caimito formations have been alleged to be of Aquitanian (early 

Miocene) age (Eames and others, 1962, p. 36-37). The 

sweeping allegation has been tacitly repudiated 

(Eames and others, 1968, p. 292-295).

CARABA FORMATION

On the recommendation of Mr. Stewart the name 

Caraba formation is adopted for strata formerly assigned to the Caimito formation. The name, in the 

form Caraba facies of the Caimito formation, was 

proposed by Jones (1950, p. 901). The thickest wellexposed section so far found is in the type region. It is 

located south of the Gamboa Reach of the Panama 

Canal, along a tributary of Rio Mandinga, about 4 

kilometers southwest of Gamboa and about 750 meters
308 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

east of Rio Caraba. The section exposed along the 

stream, as recorded by Mr. Stewart and later by Woodring, is as follows :

Section of Caraba formation along stream in type region

Approximate 

thickness 

(meters)

8. Andesitic lava_ __________________________________ 15

7. Agglomerate. _-__ ________________________________ 60

6. Sandy siltstone_ _________________________________ 8

5. Hard, dense, buff limestone; Clypeaster f ragments _ _ _ _ 3

4. Sandy siltstone and calcareous, pebbly sandstone;

Clypeaster concavus and few mollusks (locality 60) __ IS 

3. Hard, dense, buff limestone; Clypeaster f ragments ____ 8

2. Sandy siltstone and poorly sorted, silty sandstone; 

many Heterostegina israelskyi, also Lepidocyclina 

asterodisca (locality 59)__~ __--__ _________________ 18

1. Conglomerate and coarse-grained, poorly sorted 

conglomeratic sandstone; boulders have maximum 

length of 60 cm, but cobbles having length of 3 to 6 

cm more common than boulders; dacite prophyry 

conspicuous among clasts____-_-_----_---_----__- 70

Approximate thickness of section.________ 200

These strata, characterized by the exceptional thickness of conglomerate and conglomeratic sandstone, presumably interfinger with the Caimito formation. The 

total thickness of the Caraba is unknown, even in the 

type region, as the base and top have not been recognized.

Where the Caraba formation reappears northeast 

of the Canal, it consists almost wholly of agglomerate, 

in which blocks and slabs of decite porphyry generally 

predominate, and agglomeratic tuff. North of Pedro 

Miguel the agglomerate is shown on the geologic map 

(pi. 1) as part of the Pedro Miguel agglomerate, and 

in the area straddling the part of Madden Highway 

south of the Transisthmian (or Boyd-Roosevelt) Highway overpass as part of the Caimito formation. Exposures may be seen at the falls on the east side of Madden Highway at the monument site four kilometers 

south of the overpass and on abandoned Army roads 

east of the highway. Lenses of dense marine limestone, 

containing calcareous algse and larger foraminifera, 

have been found at locality 97, off Madden Highway, 

and six kilometers north-northwest of Pedro Miguel.

The fossils so far recorded from the Caraba formation and the occurrence of the same or related species 

elsewhere and in other Canal Zone formations are tabulated below.

Age. The fossils other than larger foraminifera 

indicate a late Oligocene or early Miocene age. According to the larger foraminifera, however, the age is late

Fossils from Caraba formation and occurrence elsewhere and in 

other Canal Zone formations of same or related species

Species from Caraba formation

Larger foraminifera:

Nummulites panamensis

Cushman, near locality 97.1 

Heterostegina israelskyi Gravell

and Hanna, locality 59.2

Lepidocyclina asterodisca 

Nuttall, locality 59.

Lepidocyclina sp., near

locality 97. 

Coral:

Goniopora cf. G. cascadensis

Vaughan, locality 61.3

Gastropods:

Pachycrommiumf aff.

P.? trinitatensis (Mansfield),

locality 60. 

Ficus sp., group of F. ventricosa

(Sowerby), locality 60.

Echinoid:

Clypeaster concavus Cotteau, 

locality 60.4

Occurrence elsewhere and in other 

Canal Zone formations of same or 

related species

Late Oligocene, Trinidad.

Also Caimito formation. 

Late Oligocene, Texas,

Florida, Mexico. Also

Caimito formation. 

Early Oligocene, Mexico;

late Oligocene, Texas,

Trinidad.

G. cascadensis, La Boca 

formation; late Oligocene, 

Antigua; early Miocene, 

Auguilla.

P.? trinitatensis, early 

Miocene, Trinidad. Also 

La Boca formation.

Species of F. ventricosa 

group range from late 

Oligocene to Holocene. 

Possibly also Caimito 

formation.

Late Oligocene, Antigua; 

early Miocene, Anguilla.

1 Across Madden Highway from locality 97; identification by K. N. Sachs, Jr.

2 See p. 30; identifications by W. S. Cole, 

s See p. 30; identification by J. W. Wells. 

4 See p. 31; identification by C. W. Cooke.

Oligocene: the equivalent of part, or all, of the Caimito formation.

PANAMA FORMATION

The Panama formation is redefined to include strata 

formerly assigned not only to the Panama itself, but 

also to the Bohio and Caimito formations, and the 

Pedro Miguel agglomerate. As redefined the Panama 

consists chiefly of agglomerate and tuff, extending from 

the Miraflores Lake area to Panama City, and also 

northeastward across the continental divide and eastward in the Pacific coastal area to and beyond the limit 

of plate 1. The formation also includes tuffaceous sandstone, tuffaceous siltstone, lenses of stream deposits, 

and lenses of marine limestone.

Mr. Stewart showed many outcrops of agglomerate, 

including many new exposures resulting from highway 

construction and suburban development northeast of 

the main part of Panama City and along the Transisthmian Highway. The agglomerate consists of angular to subrounded blocks, mostly andesitic, generally 

widely scattered in a matrix of fine-grained tuff. The
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 309

proportion of angular and subrounded blocks and the 

amount of tuff interbedded with, or overlying, agglomerate changes from place to place.

Stream deposits, made up of crudely bedded, tuffaceous sandstone containing scattered rounded, subrounded, and subangular boulders and cobbles, formerly assigned to the Bohio formation, are shown on 

plate 6. In the Pacific coastal area eastward from Kio 

Abajo tuffaceous sandstone and fine-grained tuff make 

up a progressively larger part of the formation and 

in the Tocumen Airport area farther east only finegrained tuff was seen in excavations during construction.

The thickest lens of limestone was observed at locality 44, about 150 meters north of the Transisthmian 

Highway, where almost cylindrical stacks of algal limestone rise above the general surface to a maximum 

height of 6 meters. The lowest exposed part of the 

limestone contains scattered pebbles and sand grains.

Pectinids were found in algal limestone at localities 

44 and 45, but no other mollusks. Larger foraminifera 

from algal limestone and their distribution elsewhere 

and in other Canal Zone formations are as follows:

Larger foraminifera from Panamd formation and their occurrence 

elsewhere and in other Canal Zone formations

[Identifications by W. S. Cole]

Species from Panama formation

Heterostegina antillea 

Cushman, localities 43, 1 

45, 1 95.2

Lepidocyclina giraudi R. 

Douville,3 localities 43, 95.

Lepidocyclina waylandvaughani 

Cole, locality 45.

Lepidocyclina yurnagunensis 

Vaughan, 4 localities 43, 45, 

95.

Lepidocyclina vaughani

Cushman, localities 45, 95.

Lepidocyclina favosa Cushman, 

localities 43, 45.

Lepidocyclina gigas Cushman,

locality 43. 

Miogypsina antillea Cushman,

locality 95.

Occurrence elsewhere and in other 

Canal Zone formations

Late Oligocene, Antigua, 

Trinidad. Also Bohio and 

Caimito formations.

Widespread in Oligocene, 

especially late Oligocene, of 

Caribbean region. Also 

Bohio, Caimito, and La 

Boca formations.

Late Oligocene, Antigua, 

Trinidad, Mexico. Also 

Bohio and Caimito formations.

Widespread in late Oligocene 

of Caribbean region. Also 

Caimito formation.

Late Oligocene, Antigua. Also 

Bohio and Caimito formations.

Widespread in late Oligocene 

of Caribbean region. Also 

Bohio formation.

Widespread in late Oligocene 

of Caribbean region.

Late Oligocene, Trinidad; 

early Miocene, Anguilla. 

Also Bohio, Caimito, 

Culebra, and La Boca 

formations.

1 See p. 27.

2 See p. 33.

3 Recorded on p. 27 and 33 as Lepidocyclina parvula Cushman.

4 Includes Lepidocyclina yurnagunensis morganopsis Vaughan of p. 27 and 33.

Age. The larger foraminifera in the preceding list 

are typical for a late Oligocene age in the Caribbean 

region. All except two occur also in the Caimito formation. Two range upward into the La Boca formation. 

Like the Caraba formation, the Panama formation is 

inferred to be the equivalent of part, or all, of the 

Caimito. The early Miocene age formerly assigned to 

the Panama was based on supposed stratigraphic relations to the La Boca, wThich have turned out to be 

erroneous.

Between Madden Highway and the Chiva Chiva 

road (the road extending from Miraflores Lake northeastward to the Transisthmian Highway) the Caraba 

and Panama formations presumably interfinger. Much 

fieldwork, however, remains to be done in that heavily 

forested and other areas before the stratigraphic relations of the agglomerates of those formations and of 

the Las Cascadas agglomerate are properly known.

MIOCENE SERIES 

CULEBRA FORMATION

As restricted on page 244, the Culebra formation is 

of limited extent along and near the Culebra Reach of 

the Canal, where it underlies the Cucaracha formation. 

Not only are fossil localities 98 to 101 to be transferred 

from the Culebra to the La Boca formation, as noted 

on page 244, but also localities 113 to 116a.

The Culebra was assumed to rest on the Las Cascadas agglomerate, but when the first core holes recently penetrated its base, it was found unexpectedly 

to rest directly on the Gatuncillo formation. The 

Culebra represents the early stage of a marine transgression culminating in the moderately deep-water 

siltstone in the upper part of the La Boca formation. 

In the Culebra Reach area the transgression was interrupted by deposition of a northwestward thinning 

wedge of nonmarine tuff, later altered to bentonitic 

clay, of the Cucaracha formation. The Culebra itself 

wedges out northwestward, as along the Empire and 

Las Cascadas reaches the La Boca overlies the Las 

Cascadas agglomerate; that is, as the marine transgression ~ continued, it extended farther northward. '

During La Boca time doubtless the entire central 

Panama area was submerged.

The present chapter covers five species from the 

Culebra formation, but none is specially noteworthy. 

On account of the transfer of fossil localities, the table 

on page 310 includes all of the Culebra gastropods: 32 

species, about half of which are not named at the 

specific level.
310 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Gastropods from Culebra formation (Neritidae to Terebridae) and occurrence in La Boca formation

[R, rare; F, few; C, common; A, abundant]

Neritina (Vittaf) sp__ __ _ _ _______

Turritella (Bactrospira?) amaras Woodring J _ _ _

Turritella venezuelana Hodson _ _

Crucibulum sp__ _ _ _ __ _

Polinices? sp ____ ___ __

Sin um sp ________ ___

Pachycrommium? cf. P. guppyi (Gabb)

Ficus carbasea micronematica (Brown and Pilsbry)___ 

Murex (Siratus) cf. M. polynematicus Brown and 

Pilsbry 2 __ ____________ _ _ ___

l\^f>t/tt In CIT*

Cymatophos? cf. C. veatchi (Olsson) _ _

Antillophosf (Antillophos?) cf. A candei gatunensis 

(Toula), ___ ___ ___ _ _ ________ __ ___ _____

1?it Q-^'OT ti Q$ cY"i

Mitraf (Cancilla?) sp.3 _ _ _

Persicula (Rabicea venezuelana amydra Woodring,

RtvmlpYphvitYn QY~i

Localities

102

R

R

F

103

R

104a

R

R

104b

___-

R

R

R

106

R

R

R

R 

R

R

R

107

A 

R

108b

R

R

108C

F

F

C

R 

R

R

110

R 

A 

F 

C 

R

R 

R

R

R

R

F

R

HOa

R

R,

R

R

Ilia

C

c

F

R

R

lllb

F

R 

R

R

112

C 

F

?R

R

R

R 

R

R

112a

F 

C

R

R

La 

Boca formation

X 

X 

X

X

X

X

X 

X

X 

X

X

X 

X

1 Described on page 101 as Turritella (Torcula?) amaras.

2 Described on page 215 as Murex (Murexf) cf. M. polynematicus.

3 Recorded on page 283 as Mitra (Tiara) sp.

4 Described on page 286 as Xancus cf. X. rex Pilsbry and Johnson.
GASTROPODS: ETJLIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 311

Though most of the species listed in the table lived 

in marine waters of normal salinity, Neretina, Littorina aff. L. angulifera, and Terebralia dentilabris? 

indicate brackish water, and Potamides suprasulcatus 

tolerated water of low salinity. These species are most 

abundant in the transition zone between the Culebra 

and the nonmarine Cucaracha formation (localities 

108c, 110).

The following table shows the occurrence elsewhere 

and in other Canal Zone formations of the same or related species:

Gastropods from Culebra formation and occurrence elsewhere and 

in other Canal Zone formations of same or related species

Species from Culebra formation

Littorina aff. L. angulifera.

(Lamarck). 

Turritella (Bactrospiraf)

amaras Woodring.1

Turritella venezuelana Hodson__

Potamides suprasulcatus 

(Gabb).

Terebralia dentilabris (Gabb)?_

Calyptraea cf. C. centralis 

(Conrad).

Cypraea cf. C. chilona Dall____

Pachycrommium? cf. P. guppyi 

(Gabb).

Semicassisf (Tylocassisf) cf.

S. aldrichi (Dall). 

Ficus carbasea micronematica

(Brown and Pilsbry). 

Murex (Siratus) cf. M. polynematicus Brown and

Pilsbry.2 

Cymatophosf cf. C. veatchi

(Olsson).

Antillophosf (Antillophosf) cf. 

A. candei gatunensis (Toula).

Xancus cf. -X". validus 

(Sowerby) .3

Persicula (Rabicea) venezuelanaamydra Woodring, n. subsp.

Conus aff. C. chipolanus Dall.__

Occurrence elsewhere and in other

Canal Zone formations of same or

related species

L. angulifera, late Miocene

to Holocene. 

T. caparonis Maury, early

Miocene, Trinidad. Also

La Boca formation. 

Early Miocene, Venezuela.

Also La Boca formation. 

Widespread in Caribbean

region, late Oligocene to

middle Miocene. Also La

Boca formation. 

T. dentilabris, Miocene,

Dominican Republic. 

C. centralis, early Miocene to

Holocene. Also La Boca

formation. 

C. chilona, early Miocene,

Florida. Also La Boca

formation. 

P. guppyi, middle Miocene,

Jamaica; early to middle

Miocene, Dominican

Republic. Also Bohio and

La Boca formations. 

S. aldrichi, early Miocene,

Florida. 

Early Miocene, Peru. Also

La Boca formation. 

M. polynematicus, Gatun

formation. Also La Boca

formation. 

C. veatchi, Gatun formation;

middle Miocene, Costa

Rica. Also La Boca formation. 

A. candei gatunensis, Gatun

formation. Also La Boca

formation. 

X. validus, early to Middle

Miocene, Dominican

Republic. Also La Boca

formation. 

P. venezuelana venezuelana,

early Miocene, Venezuela.

Also La Boca formation. 

C. chipolanus, early Miocene,

Florida. Also Caimito and

La Boca formations.

1 Described on p. 101 as Turritella (Torcula?) amaras.

2 Described on p. 215 as Murex (Murex?) cf. M. polynematicus. 

s Described on p. 286 as Xancus cf. X. rex Pilsbry and Johnson.

368-278 O - 70 - 2

Age. Though the Culebra fauna is small, it has an 

unmistakable Miocene stamp the earliest of the Miocene faunas in the Canal Zone. Moreover, the mollusks 

indicate the earliest part of the Miocene. All except 

three of the species in the preceding table occur also 

in the La Boca formation, an indication of the close 

age association of the two formations. Orthaulax and 

Pachycrommium are extinct, but both genera are 

doubtfully identified.

CUCARACHA FORMATION

Fresh-water snails of the genus Hemisinus (H. aff. 

H. oeciscus) are the only gastropods found in the 

Cucaracha formation.

Age. The Cucaracha formation is bracketed by formations containing early Miocene marine fossils. 

Therefore the Cucaracha itself is of early Miocene age 

in terms of the marine succession. North American 

land mammals recently found in the Cucaracha are 

considered to be of middle Miocene age (Whitmore 

and Stewart, 1965, p. 182).

LA BOCA FORMATION, INCLUDING EMPERADOR LIMESTONE 

MEMBER

Gaillard Out area. Isolated outcrops of fine-grained 

rocks of the La Boca formation have been accessible 

along the Canal, and also outcrops of sandstone some 

distance from the Canal, as at the abandoned quarry 

south of Summit Gardens (locality 128). No outcrop 

localities showing a considerable thickness, however, 

were available until the excavation involved in the program of widening the Canal reached the Las Cascadas 

Reach in 1964-67. (The Las Cascadas Reach is the second reach southeast of Gamboa. Localities 99a, 99g, 100, 

101, and 120 are plotted on the reach on plate 2.) The La 

Boca was exposed on the west side of the reach in a strip 

about 125 meters wide and a little more than a kilometer 

long from the major fault at Canal station 1598, which 

brings the underlying Las Cascadas agglomerate up 

above the level of Gaillard Cut, southeastward to station 1633. Many minor faults are apparent in the strip. 

The following section was exposed at station 1630, near 

the southeast end of the excavated strip:

Section of lower part of La Boca formation on west side of Las

Cascadas Reach at Canal Station 1630

Approximate 

thickness

6. Emperador limestone member:

e. Noncoralliferous limestone; Aequipecten canalis, 

Amusium, Spondylus, Clypeaster concavus 

(locality Il7b)___-- -- - - 1- 5

d. Poorly exposed sandy siltstone______-_____---- 3

c. Coralliferous limestone; corals, small Spondylus-- 1. 5 

b. Sandy siltstone------------------------------ 1-5
312 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Section of lower part of La Boca formation on west side of Las 

Cascadas Reach at Canal Station 1630 Continued

Approximate 

thickness 

(meters)

6. Emperador limestone member Continued

a. Coralliferous limestone; corals, Turritella amaras

and other mollusks (locality H7a)___________ 1. 5

5. Greenish silty clay_______________________________ 1. 5

4. Carbonaceous shale_______________________________ . 3

3. Siltstone, clay, silty sandstone, poorly sorted somewhat calcareous sandstone, and conglomeratic 

sandstone at foot of steep slope; mollusks in somewhat calcareous sandstone 30 meters to northwest 

(locality 101e)___--_--_________________________ 10. 5

2. Siltstone and clay, poorly exposed on bench_________ 12

1. Lens of disintegrating echinoid-bearing limestone just 

beyond station 1632; mollusks and echinoids 

(Echinolampas cf. E. lycopersicus, Clypeaster concavus, identifications by P. M. Kier; locality 101b)_ 3 

Las Cascadas agglomerate. _____

Approximate thickness of section.______________ 36. 3

At station 1627 another lens of basal echinoid-bearing limestone is represented by huge masses of hard, 

scabrous limestone. Mollusks and echinoids that weathered out of the hard rock were collected there (locality 

lOla). The echinoids, identified by P. M. Kier, are as 

follows: Echinolampas cf. E. lycopersicus and Agassizia clevei.

The Emperador limestone member was exposed on a 

bench at station 1626 (locality Il7c). Corals, especially 

Acropora and Porites, and a few mollusks are strewn 

on the bench. The mollusks include five specimens of a 

big cerithid, Campanile cf. C. herculeanus.

Some 60 species of mollusks were collected from 

somewhat calcareous, silty sandstone on a bench extending from station 1608 to 23 meters beyond station 

1612 (locality lOlh). This fossiliferous bed, characterized by the abundance of a small Aequipecten and 

a small Dimya, was not seen elsewhere. It is about 25 

meters above the Emperador limestone member. The 

collection contains also many well-preserved specimens 

of a small, heavily pillared Lepidocyclina, identified by 

K. N. Sachs, Jr., as L. giraudi.

The following section was measured at station 1622. 

The base is actually at the west end of a bench formed 

by the Las Cascadas agglomerate, 75 meters west of the 

Canal cliff at station 1622, and the top at the top of 

unweathered rock near the top of the steep slope at 

station 1621.

The samples listed in the following section are foraminiferal samples, which have been studied by Blacut 

and Kleinpell (1969). The section is shown in graphic 

form in their publication.

Section of La Boca formation on west side of Las Cascadas Reach 

at Canal station 1622

Thickness 

(meters)

16. Thin-bedded, dark gray, sandy siltstone; sample 

 410-10; overlain by weathered rock and residual 

clay._-_----_----___--___-_----_-__-_-_______ 5. 1

15. Thin-bedded, brownish, sandy siltstone- _ __________ 4.8

14. Silty sandstone and very sandy siltstone. __________ . 7

13. White vitric tuff, lenses out farther northwest than

unit ll____-_-_--___--_______________________ . 9

12. Silty sandstone and very sandy siltstone; crumbly

Lepidocyclina and Acila- _______________________ 17. 1

11. White vitric tuff, lenses out to northwest_________ 1.3

10. Siltstone and sandy siltstone; sample 410-4, 2.8 m 

above base; 410-5, 4m; 410-6, 4.8 m; 410-7, 6 m; 

410-8, 7.4 m; 410-9, 8.8 m___________________ 9. 2

9. Claystone and mudstone, lower half dark gray, upper

half chocolate brown__________________________ 1. 8

8. Soft mudstone and silty mudstone; leached foraminifera; includes two beds of calcareous silty sandstone, 

30 to 60 cm thick_________-___-__-___________. 24. 6

7. Emperador limestone member:

d. Thin-bedded limestone, mat of broken branching corals_----_-_--_-_----_-_----_----__- 2. 1

c. Thin-bedded, calcareous siltstone-____________ . 2

b. Thin-bedded limestone, mat of broken branching

corals (locality H7d)____________________ . 6

a. Thin-bedded, calcareous, sandy siltstone_-____- 1. 5

6. Sandstone, mudstone, and carbonaceous shale___-_-_ . 4 

5. Mudstone and sandy siltstone; many Bittium scotti 

and molds of other small mollusks (locality lOld); 

sample 410-1, 1.1 m above base_______________ 3. 3

4. Sandy siltstone; mollusks, for most part fragments: 

Cypraea, Anadara, Trachycardium, Tagelus (locality 

101c)_-----------_------_----_-------_------- . 5

3. Carbonaceous shale____-_-_-_----_----_-_--_----_ .6

2. Mudstone, black calcareous nodules; ostracodes, fragmentary mollusks___-______-__________--_-__-_ 1. 2

1. Bentonitic clay_________-_-___--_-_--___----_--- 1. 3

Las Cascadas agglomerate.

Thickness of section___________________________ 77. 2

Half a kilometer farther northwest a lens of dense 

limestone crops out in the upper part of exposed strata, 

as shown in the following section:

Section of La Boca formation on west side of Las Cascadas Reach

from Canal station 1605 to 1601

Approximate 

thickness 

(meters)

8. Silty sandstone and sandy siltstone; scattered foraminifera and mollusks_____________---__-_-_----- 15

7. Lens of white vitric tuff___________-_-------------- 3

6. Silty sandstone and sandy siltstone---------------- 6

5. Lens of dense limestone; Pachycrommiumf cf. P.? 

trinitatensis, Aequipecten canalis, Amusium, echinoids 

(Echinolampas semiorbis, Clypeaster concavus, identified by P. M. Kier; locality 101g)__-_--_--------- 4. 5

4. Sandy siltstone and silty sandstone, few thin calcareous

beds--.-------------------------------------- 20
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 313

Section of La Boca formation on west side of Las Cascadas Reach 

from Canal station 1605 to 1601 Continued

Approximate

thickness

(meters)

3. Poorly exposed sandy and silty beds________________ 7.5

2. Emperador limestone member:

e. Lenticular limestone, many Amusium.- _________ 6

d. Alternating siltstone and limestone in beds 30 to

60 cm thick_______________________________ 2

c. Limestone__________________________________ 3

b. Siltstone__--------____________________..__ .3

a. Coralliferous limestone near level of Canal__-__- 3

1. Carbonaceous clay and tuffaceous siltstone^_________ . 5

Approximate thickness of section.______________ 70. 8

Immediately south of the major fault at station 1598 

the base of the La Boca, overlying the Las Cascadas 

agglomerate, consists of variegated clay overlain by 

fine-grained vitric tuff.

Core hole LBW 149, located 250 meters west of the 

top of the measured section at station 1622, penetrated 

strata higher than the measured section. On the assumption that a 2.3-meter bed of vitric tuff penetrated 

in the core hole corresponds to unit 13 of the measured 

section, the combined outcrop and subsurface thickness of the La Boca is 137 meters, and the top of the 

formation is not represented.

Madden ~basin. As mentioned on page 300, the agglomerate on the south side of Madden basin, exposed 

on the Transisthmian Highway immediately north of 

the intersection with Madden Highway, is identified 

by Mr. Stewart as the Las Cascadas agglomerate and 

the overlying fine-grained rocks as the La Boca formation. Both were formerly grouped as the pyroclastic-clay member of the Caimito formation (p. 32). The 

fine-grained rocks are poorly exposed or unexposed. 

Molds of foraminifera, including Siplwgenerina, were 

observed in calcareous mudstone on a tributary of Rio 

Chilibre (Quebrada Ancha, not shown on plate 1) 

about 500 meters east-northeast of the Transisthmian 

Highway bridge across Rio Chilibre. Other streams, 

however, in that and adjoining areas have not been 

traversed.

The lenses of limestone, on which localities 71, 72, 

and 73 are plotted on plate 1 evidently represent outcrops of different parts of a limestone unit, now identified as the Emperador limestone member of the La 

Boca. In unpublished reports it has been designated 

informally as the Chilibre limestone.

Gastropods, GaiUard Out area. Seventeen species 

from the La Boca formation in the Gaillard Cut area 

are described or recorded in the present chapter. Floribella aldrichi is the most noteworthy of these species. 

It is considered to be a remarkable philinid, by far the 

largest fossil or living species. Ellobiium aff. E. pellucens is the first American fossil representative of the 

salt-marsh pulmonate genus Ellobium. Terebra dicker es is the earliest known species of the subgenus 

Oreoterebra.

The La Boca fauna has been greatly enlarged by the 

collections from localities lOlh and 116a (some 60 and 

some 90 species of mollusks respectively). As for the 

Culebra formation, all the gastropods from the La 

Boca formation of the Gaillard Cut area are listed in 

the table on pages 314-315. The table lists 78 species, half 

of which are not named at the specific level. Three of 

those not so named, chiefly from localities lOlh and 

116a, are to be described in chapter E. No mollusks 

have been recovered in Madden basin, except from the 

Emperador limestone member. (See table, p. 314.)

The occurrence elsewhere and in other Canal Zone 

formations of the same or related species is as follows:

Gastropods from La Boca formation of Gaillard Cut area and 

occurrence elsewhere and in other Canal Zone formations of 

same or related species

Species from La Boca formation

Turritella cf. T. collazica

Maury. 

Turritella (Bactrospiraf)

amaras Woodring. 1

Turritella (Bactrospira) altilira 

Conrad, subsp.2

Turritella subgrundifera Dall 3 _ 

Turritella venezuelana Hodson_

Turritella cf. T. berjadinensis

cocoditana Hodson. 

Hemisinus (Longiverend) aff.

H. oeciscus Woodring. 

Potamides suprasulcatus

(Gabb).

Bittium scotti Brown and

Pilsbry. 

Architectonica (Architectonica)

nobilis Roding, subsp.4 

Calyptraea cf. C. centralis

(Conrad).

Orthaulax gabbi, Dall________

Cypraea cf. C. chilona Dall___

Globularia (Globularia) aff. 

G. fischeri (Dall).

Pachycrommiumf cf. P.?

trinitatensis (Mansfield). 

Pachycrommiumf cf. P.

guppyi (Gabb).

Morum (Cancellomorum) cf.

M. chipolanum Maury.5 

Ficus carbasea micronematica

(Brown and Pilsbry).

Occurrence elsewhere and in other

Canal Zone formations of same or

related species

T. collazica, late Oligocene,

Puerto Rico. 

T. caparonis Maury, early

Miocene, Trinidad. Also

Culebra formation. 

Subspecies of T. altilira range

from late Oligocene to late

Miocene.

Early Miocene, Florida. 

Early Miocene, Venezuela.

Also Culebra formation. 

T. berjadinensis cocoditana,

middle Miocene, Venezuela. 

H. oeciscus, Bohio formation.

Widespread in Caribbean 

region, late Oligoeene to 

middle Miocene. Also 

Culebra formation.

B. permutabile, early Miocene, 

Florida.

A. nobilis, early Miocene to 

Holocene.

C. centralis, early Miocene to 

Holocene. Also Culebra 

formation.

Early Miocene, Florida.

C. chilona, early Miocene, 

Florida. Also Culebra, and 

Caimito formations.

G. fischeri, early Miocene, 

Florida. Also Caimito and 

Bohio formations.

P.? trinitatensis, early 

Miocene, Trinidad.

P. guppyi, middle Miocene, 

Jamaica; early to middle 

Miocene, Dominican Republic. Also Culebra and 

Bohio formations.

M. chipolanum, early 

Miocene, Florida.

Early Miocene, Peru. Also 

Culebra formation.

Continued on page 316, with, footnotes.
314 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Gastropods from La Boea formation of Gaillard

[R, rare; F, few;

ATprvfn «n

Potamides suprasulcatus (Gabb) _ _

Campanile? sp_ _ _ __ _ _____

S*t'n /i/'yn cn

7V/Y/7v»7/^r P «:n

Northiaf cf. N. northiae (Gray)_ _ ___ _ _

f)l'J1}011 ft C!T"i

PT^/OT iiwi ( 7l//V_**T/') QTft'i'Ytft\ tiff P fi<nft1ft/*h 00 ( flcnT-T- &\*\

P Tt 1 ft 1 1 *ffl ( /l/i iFYfltffflYfl 1 «!Ti

~P0ir<i'ift ii 1 ft ( 7? ft h vf^p n^\ ?) PW f> yn P! frn ft ftvniirivft A^rirkH 1*1 no* n QII Vi^JTi

Crassismra (Crassisvirelld) SD _ _ _ _

Localities

98

R

99a

?sp.R

R

R

R

99b

R

A

R

C

R

C

R

A

R

C 

R

R 

?F

?F

R 

?R

99c

R

F

R

F 

F

R

____

F 

R

99d

R

?R

C 

?sp.R

R 

R 

F

R

R

R

R

99e

R

 

 

----

99f

R

R

R

99g

?R

R 

F 

sp.R

R

F

99h

F

R

R

100

C.

R

R

R

F

C

?C

?C

lOOa

A

sp.R

R

C

See footnotes at end of table.
Cut area (Acmaeidae to 

O, common; A, abundant]

GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE

Ellobiidae)

315

Localities Continued

lOOb

R

R

R

101

A

R

?F 

?F

lOla

F

lOlb

R ._

R

?R

lOlc lOld lOle lOlg

R

lOlh

A 

R 

C

R

R

A

R

R

R 

R

C 

F

R

R 

A 

C 

R 

A

C 

?C

R

R

R 

C 

R

lOli

C 

F

C

C 

R

R 

R

114

R

F

R

R,

R 

R

R 

R 

R,

R,

R 

R 

R

F 

F

R

115

 

----

R

R 

R

115a

?F

F

R

F 

sp.R

R 

F

R

R

R

R

F 

R

115b

R

C

R

F

R

R

C

116

R

----

R

F

R

----

----

----

116a

R 

R 

C 

C 

R 

R 

R 

R 

F

C

R 

R 

F 

F

R

F 

C 

A 

F

C

F 

R 

R 

R 

R

R

R

F 

F 

F 

R

R

A

R 

C

R

119

sp.R

R

119a

R

R

120

R

R 

sp.R

R

R

125

R

 

130

 

R
316 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Gastropods from La Boca formation of Gaillard

Clathrodrillia sp__ _ _ _ _ ___ __

Terebra (Oreoterebra) dicker es Woodring, n. sp

Strioterebrum cf. S. clethra (Maury) _

Acteocina sp___ ____

Atysf sp _ _

Retusaf (Cylichninaf) sp ___ ____ __

Floribella aldrichi (Dall)

Ellobium aff. E. pellucens (Menke) _

Localities

98 99a 99b

R

99c 99d 996 99f 99g 99h 100 lOOa

F

1 Described on page 101 as Tunitella (Torcula?) amaras.

2 Recorded 011 page 103 as Turritella (Torcula) altilira, subsp.

3 Described on page 105 as Tunitella cf. T. subgrundifera.

4 Recorded on page 81 as Trochita cf. T. trochiformis (Born).

5 Described on page 203 as Morum ("Oniscidia") cf. M. chipo anum.

6 Described on page 215 as Murex (Murex?) cf. M. polynematicus.

7 Described on page 286 as Xancus cf. X. rex Pilsbry and Johnson.

Gastropods from La Boca formation of Gaillard Cut area and 

occurrence elsewhere and in other Canal Zone formations of 

same or related species. Continued from p. 313

Species from La Boca formation

Murex (Siratus) cf. M.

polynematicus Brown and

Pilsbry.6 

Strombina cf. S. quirosana

Hodson. 

Cymatophos? cf. C. veatchi

(Olsson).

Antillophosf (Antillophosf) cf.

A. candei gatunensis

(Toula). 

Northiaf cf. N. northiae

(Gray). 

Nassarius (Uzitaf) praeambiguus (Brown and Pilsbry). 

Xancus cf. X. validus

(Sowerby) .7

Prunum (Microspira) aff. P.

apalachee (Gardner). 

Persicula (Rabicea) venezuelana

amydra, Woodring, n. subsp.

Conus cf. C. planiceps Heilprin. 

Conus aff. C. chipolanus DalL-.

Gemmula cf. G. vaningeni

(Brown and Pilsbry). 

Terebra (Oreoterebra) dicheres

Woodring, n. sp. 

Strioterebrum cf. S. clethra

(Maury). 

Scaphander (Scaphander) cryus

Woodring?, n. sp. 

Floribella aldrichi (Dall)______.

Ellobium aff. E. pellucens

(Menke).

Occurrence elsewhere and in other

Canal Zone formations of same or

related species

M. polynematicus, Gatun 

formation. Also Culebra 

formation.

S. quirosana, early Miocene, 

Venezuela.

C. veatchi, Gatun formation; 

middle Miocene, Costa 

Rica. Also Culebra formation.

A. candei gatunensis, Gatun 

formation. Also Culebra 

formation.

N. northiae, Holocene.

Also Gatun formation.

X. validus, early Miocene,

Dominican Republic. Also

Culebra formation. 

P. apalachee, early Miocene,

Florida. 

P. venezuelana venezuelana,

early Miocene, Venezuela.

Also Culebra formation. 

C. planiceps, early Miocene,

Florida. 

C. chipolanus, early Miocene.

Also Culebra and Caimito

formations.

G. vaningeni, Gatun formation. 

T. odopoia Gardner, early to

middle Miocene, Florida. 

S. clethra, early Miocene,

Brazil. 

Perhaps same species as S.

cryus, Bohio formation. 

Early Miocene, Florida, Cuba. 

E. pellucens, Holocene.

1 Described on p. 101 as Tunitella (Torcula?) amaras.

2 Recorded on p. 104 as Tunitella (Torcula) altilira Conrad, subsp.

3 Described on p. 105 as Tunitella cf. T. subgrundifera.

4 Described on p. 165 as Architectonica (Architectonica) cf. A. nobilis Roding.

5 Described on p. 203 as Morum ("Oniscidia") cf. M. chipolanum.

6 Described on p. 215 as Murex (Murex?) cf. M. polynematicus.

7 Described on p. 286 as Xancus cf. X. rex.

Gastropods, Emperador limestone member. The 

Emperador limestone member yielded the gastropods 

tabulated below. Aside from species identified only at 

the generic level, Campanile cf. C. herculeanus is the 

only one not in the preceding table. The Emperador 

species and C. herculeanus, from the early Miocene 

Anguilla formation of Anguilla, are the last American 

species of the genus, younger than any in Europe.

Gastropods from Emperador limestone member of La Boca 

formation

Turritella cf. T.

Turritella (Bactrospiraf) amaras

Turritella (Bactrospira) altilira

Campanile cf. C. 

herculeanus (Cooke) _

Cypraea cf. C. chilona 

Dall-. -------------

Ficus cf. F. carbasea 

micronematica 

(Brown and 

Pilsbry)_ __ _______

Localities

Gaillard cut area

117a

R

117c

F

118

R

R 

R

R,

121

R

123

R

129a

R,

Madden 

basin

71

R

R,

R 

R

R,

73

R

R

Age. The La Boca fossils have marked early Miocene affinities. They suggest correlation with the Chipola formation of Florida. Nevertheless, the La Boca 

is considered to be older than the Chipola, chiefly on 

the grounds that Lepidocyclma and Nemocardium sur-
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE

Cut area (Acmaeidas to Ellobiidae) Continued

317

Localities C ontinued

lOOb 101 lOla lOlb lOlc lOld

C

lOle lOlg lOlh lOli

R

114 115 115a

R

- R

R

1151)

R,

116 11 6a

sp.F

R 

R

R

119 119a 120 125 130

vived in La Boca time. To be sure Lepidocyclina is unknown in southeastern United States later than Oligocene. Aside from the survival of Netnocardium in the 

Tampa limestone (Mansfield, 1937, p. 258; recorded as 

Protocardid) the La Boca fossils show little resemblance to those of the Tampa, although they are 

thought to be of the same age. Orthaulax, Globularia, 

Pachycrommiumf, and Flonbella are extinct.

The smaller benthonic foraminifera from the La 

Boca indicate correlation with the early Miocene Saucesian stage of California (Blacut and Kleinpell, 1969, 

p. 5). In addition to Lepidocyclina giraudi from locality lOlh, a collection from the east side of Las Cascadas Reach (locality lOli), at a horizon corresponding to unit 12 of the measured section on the west side 

of the reach, contains many well preserved specimens 

identified by K. N. Sachs, Jr., as Lepidocyclina tniraflorensis. Those two species are the last representatives 

of the genus in the Canal Zone.

The large, high-domed echinoid, Echinolampas semiorbis, is conspicuous in the Emperador limestone member in the Gaillard Cut and Madden basin areas. It 

occurs in the Anguilla formation of Anguilla, which 

is correlated with the La Boca.

ALHAJUELA FORMATION

The Alhajuela formation is defined to include the 

calcareous sandstone member and Alhajuela sandstone 

member of the Caimito formation in Madden basin, as 

used in chapter A (p. 32-33). It is the youngest formation in Madden basin, which is the type region. The 

base is tentatively selected as the base of conglomerate 

formerly exposed on the Transisthmian Highway at 

locality 74, 1.5 kilometers south of Eio Chilibrillo 

bridge.

The formation is divided into a lower member and 

an upper member. The lower member consists of the 

former calcareous sandstone member of the Caimito 

formation and includes the lens of sandy limestone 

formerly designated the Chilibrillo limestone member 

of the Caimito (p. 32). The upper member is the 

former Alhajuela sandstone member of the Caimito. 

The thickness of the formation is estimated to be 115 

to 145 meters.

Gastropods are rare in the Alhajuela formation and 

with few exceptions are represented by molds. The following species are recorded in earlier chapters:

Gastropods from Alhajuela formation

Tunitella (Bactrospira)

Tunitella gatunensis

Ficus carbasea carbasea 

(Guppy) . -------------

Localities

Lower member

77

cf. R

sp. R

79

R

80

cf. R

82

R

82a

'R

Upper member

85

R

85a

?R

88

?

R

R

89

C

92

?

i Described on p. 102 as Turrtietta (Torcula) aMira attilira.

Age. The upper member contains the typical form 

of Turntella altilira and the typical form of Ficus carbasea the earliest known occurrence of those forms, 

which are found in the Gatun formation. That the age 

of the entire Alhajuela formation is late early Miocene, rather than middle Miocene, is indicated by the 

pelecypods.

GATUN FORMATION

The present chapter covers 135 species and subspecies 

from the Gatun formation and two unnamed ones are
318 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Gastropods from Gatun formation (Eulimidae, Marginellidae to

[R, rare; F, few;

Clathrodrillia saavedrai Woodrine. n. sn.. --.... ___ .-- _ ___ .. ____ - ___ .--

Localities

Lower part

S

R

F

A 

F

R

F

R

R

R

C3 

%

-

C

R

C

R

0

F

R

R

R

R

r*-

co

R

F

R

R

R

R

R

F

C3 

?!

-

F

R 

R

R

R

F

R

8

-

R

F

R,

F

R

F

R

R

R

R

R

C

C3 oo

CO

F

F

R

'R

R

F

R 

R

R

R,

R,

R

R

F

R

R

R

,Q

s

-

R

R

R

R

R

£

.R 

R,

C

R,

?R

K

_F

A

R,

O

R

F

A

A

A

R

C 

R

0

F

0

R

R

0

0

0

R

R

.A

.A

R

C

C

 0

oo

R

R

R,

F 

F

R,

A

C

A

F

R

R,

R

R,

R,

R,

C

C

F

F

1

C 

R

--

R

F

F

C

R,

0

F

F

R

R

8

-

-

R

F

C

C

C 

0

R,

R,

R

R

F

F

R

R

tf

-

-

-

--

§

 

---

---

Middle part

Eastern area

,Q

8

-

R,

R,

R

0

--

R

F

R

F

*

F

R

R

R

0

C

F

R,

R,

R

R,

0

F

C

R

R

F

R

F

 o

S

R

--

--

R,

R

1

R

m

R

R

R

F

R,

F

C

R,

R,

R,

-

C

R

R,

R

R,

R

R

IP

R

-

o

R

R,

--

C^

R,

-

R,

R,

^i

--

R,

S

A

-

R,

K,

R

F

R,

R,

r~ *

 

?F

& 

5

R

R,

R

O

A

0

?R

 'R

R,

0

0

R

F

0

R

t>-

Tt<

-

r- *

 

C

?R

R

R

R

R

R
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 319

Cavolinidae and a few species of other families) 

C, common; A, abundant]

Localities Continued

Middle part Continued

Eastern area Continued

M

M

R

C

A

...

C

F

C

F

F

C

B

M

R

R

F

--

R

R

R

R

R

R

"R

Ji

--

-

--

--

US

R

--

R

R

F

F

F

C

F

=3

--

R

---

--

--

03 =2

R

--

R

?R

--

-

-x

R

--

...

R

--

--

--

>o 

>o

R

R

C

R

R

R

R

C 

C

F

C

C 

R

R

C

C

C

C

3

R

R

F 

F 

R

R

F

R

R

F

R

R

JD

s

R

3

R

C 

C

F

C

R

C

F

F

F

8

--

c

R

--

R 

R

R

C

R

C

R

C

C

<% £

R

F

... I

?R 1

R <

... ]

. R

R

SR

---

,

t?R

*

R ...

R

R

C ..

OS 10

R

F 

R

R

R

-

3 

>o

-

-

R

5

o

R

F

R

T3

OS>o

R

R

R

R

7

R

R,

R

R

R

r

r

c

R

r

F

is.2 U"S

JPSJ

R

R

R

Western area

o 

so

?R

g

R

-

s

R

-

o

R

R

H 1

03 

SO

R

^J

F

A

R

R

,0

so

R

R

F

-

o

SO

R

R

R

C

R

R

R

T3 

to

R

F

R

-o

--

R

SrO

...

3

-

O t>-

m

?R

F

...

cS

R

Upper part

Eastern area

3 t>-

R

...

?R

R

R

N

F

R

R

R

R

«

R

F 

R

R

F

R

03 

«

"

-

R

R

-*

-

-

-O

R,

R

A

R

R

R 

R

F

R

A

R

C

R

C

F

so

F

R

R

F

R

C

R

R

sg

R

R

R

R

-

R

t>- t>-

R

!R

C

R

R

R

03

R

R

--

,Q

p,

F

R

A

R

R

R

R

o

C

R

R

R

--

T3 t>- 

J>j

R

F

?R

...

<a

--

00j>

?R

R

«

R

Other col-lections

R

....

R

Western area

g

...

§

H,

R

K,

R

R,

R

R

--

03

2

R

R

R

R

R

--

R

oo

00

R

C

R

to

00

sp.R

?R

F

R

R

R

~~~R
320 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Gastropods from Gatun formation (Eulimidae, Marginellidae to

Clathrodrillia aff. C. lelandi (Olsson)

Microdrillia trina Mansfield

Kurtziella (Cryoturris) sp

Nannodiella rintriada (Mansfield)

Nannodiella cf. N. melanitica (Ball)

Localities

Lower part

£

R

F

F

0

-R

03 

%

R

F

r

A

F

r~

03

 R.

0

R

R

?R

F

F

R

R

03 

5

R

R

R

§8

R

F

F

R

F 

F

R

F

C 

C

f!

A

F

o

F

03 

8

?R

0

R

F

R

R

R

R

C 

C

r,

c

F

F

f!

A

R

r

,0 

88

R

R

F

o 

§8

F

R

C 

F

R

R

R,

R

C

C

R

R

C 

C

A 

A

A

R 

R

r

R

F

R

A 

R

r

R

S

F

?R

F 

R

R

R

R 

R

R

R

R

F 

A

A

R

R

C

«

R

R

m

F 

?R

c

R

R

R

R

F

R

i

R

R

R

f!

A

C

R

bo §8

R

C

F

R

R

02

03

?R

Middle part

Eastern area

p 

§

R

R

C

R

r,

R

f!

R

F

R

F

R

R

A

F

R

F

r,

r,

o

3

R

C 

F

F

F

R

o

R

R

R

F

F

R

F

r,

F

F

f!

A

R

R

F

F

 d 

§

R

c 

§

R

F

F

1

R

r,

R

1

R

r

R

R

0

3

R

 *

R

(M * 3

CD

3

R

f!

F

R

F

R

R

F

R

F

0

R

R

t3

,0 

r~ 

3

F

A

A

A

R

A

A

A

r

A

R,

A

R

F

R

R

n

A

A

A

n

A

F

0

3

R

r~-*i

F

F

C

R

R

C

F
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO

Cavolinidae and a few species af other families) Continued

HELMINTHOGLYPTIDAE 321

Localities Continued

Middle part Continued

Eastern area Continued

u> 

-*

F 

A

F

A

C 

C

F

R

F

R

C

R

A 

C

F

R

R

Tjl

F

C

C

F 

F

R 

R

-

R

F

R

R

 *

F

?R

R

>n

R

R

C

s?

---

03

s

R

R

R

R

-

R

s

-

18

R

R,

R

"R,

F

F

 R.

R

F

R

R

F

03 tf3 >O

R

F

f:

s

R

F

R

pp R

0

R

8

R

p

F

R

R

R

F

R

>o

R

o

R

^

R

R

F

S ib

R

F

03 Oi >o

Oi

>o

g

5

F

F

R

R,

R

R

R,

R,

F

R

R

F

R

R

R

R

Other col-lections

R

R

Western area

o

CO

m

S § to

R

R

R

03

to

r,

.0

0

R

R

R

F

R

R

F

R

R

R

?R

T3

?R

i

R

R

03

to

?R

S

o!>-

m

S

R

C

R

R

F

Upper part

Eastern area

S

R,

R,

tR,

R

R

CO

R

F

£

R

R,

R

R,

R,

 *

R

R

>o

R

R

R

R

A 

A

R

to

R

R

A

03 to

R

R

t~ t~

R

R

R

R

03

R

F

X! t^r^

R

R

C

C 

C

R

R

o

F

R

R

C

R

T3t^

R

R

R

5

hC

00r~

R

col- 3therlections

C

Western area

o t^

R

?R

00

R

R

03 

C^ 00

R,

»

00

R

?R

>o

00

R

R

F
322 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

mentioned. Two other species, described in earlier chapters, are redescribed on the basis of better material in 

recently acquired collections.

The Gatun fossils embrace 18 cancellarids (including a species of the subgenus Charcolleria and of the 

genus Aphera), 16 conids, 50 turrids (including two 

species of the new mangeline genus Dolostoma, and a 

species each of Euglypliostoma and Rhiglyphostoma, 

new subgenera of the mangeline genus Glypliostoma) , 

and 13 terebrids (including five large species and subspecies of Terebra, assigned to the subgenera Oreoterebra, Paraterebra, and Panaterebra).

The distribution of the species and subspecies in the 

formation is shown in the table on pages 318-321.

The Gatun gastropod fauna now totals 293 species 

and subspecies (plus eight others not formally described). To that total should be added 5 that remain 

to be described and 10 undescribed pyramidellids. For 

the purpose of tabulation the arbitrarily separated 

eastern and western areas of the middle part (p. 46) 

are consolidated, but the geographically isolated eastern and western areas of the upper part are retained. 

The number of species and subspecies in the three parts 

of the formation, and the largest collections are tabulated below.

Gastropods from Gatun formation

Part

Lower __ _ __

Middle__--__________.______

Upper, western _ ___ _

Number of 

species and 

subspecies

145 

181 

82 

53

Largest collection

Number of 

species and 

subspecies

125 

104 

35 

25

Locality

138c 

139c 

177b 

185

Ninety-five of the 230 unequivocally identified species 

and subspecies are endemic. The occurrence elsewhere 

of the other 135 is shown in the table on pages 323-324.

Age. The distribution elsewhere supports a middle 

Miocene age for the Gatun formation, even for the 

upper part in the western area, which was formerly 

considered to be late Miocene. The closest similarity is 

with unrecorded middle Miocene faunas from Darien 

Province and Chiriqui Province, both on the Pacific 

side of Panama, and also with the middle Miocene of 

southeastern Costa Rica.

For some unknown reason the small fauna of 53 

species from the upper part in the western area is 

unique in biofacies, although 30 of the species occur 

elsewhere in the Gatun.

Diarecallus, Apiocypraea, Strombinophos, Strombinella, Subterynotus, Panamurex, Pilsbry ty phis ^ Galophos, Rhipophos, Nicema, Nanarius, Psttarius, Charcolleria, Scobinella, and Dolostoma are extinct.

Taxa that formerly lived in the western Atlantic 

part of Tertiary Caribbean province, but now are extinct there and survive in eastern Pacific waters, have 

been designated paciphiles (Woodring, 1966, p. 426). 

The following paciphile genera and subgenera are 

represented in the gastropod fauna of the Gatun formation :

Paciphile genera and subgenera in gastropod fauna of Gatun 

formation

Cymatophos

Oliva (Strephonella)

Mitra (Cancilla) 2

Cancellaria (Pyruclia)

Cancellaria (Euclia)

Cancellaria (Narona)

Aphera

Pleurofusia (Cruziturricula}

Agladrillia (Agladrillia}

Glyphostoma (Euglyphostoma)

Terebra (Panaterebra}

Rictaxis

Teinostoma (Aepystoma)

Solariorbis (Hapalorbis}

Heliacus (Astronacus)

Rhinoclavis (Ochetoclava}

Trochita

Neverita (Glossaulax)

Neverita (Hypterita}

Malea

Thais (Vascula)

Strombina (Sincola)

Solenosteira l

Metula

1 Described on p. 256 as Hanetia.

2 Described on p. 283 as Tiara.

A few paciphiles, listed below, are at the specific 

level:

Paciphile species in gastropod fauna of Gatun formation

Trochita spirata Forbes? l

Neverita (Glossaulax) reclusiana xena Woodring 

Neverita (Hypterita} helicoides Gray 

Distorsio (Rhysema) decussata gatunensis Toula 

Cochlespira (Ancistrosyrinx) cedonulli (Reeve) 

Volvulella (Volvulella) cylindrica parallela (Pilsbry and Johnson) 

i Described on p. 81 as Trochita trochiformis (Born).

Still fewer species survive on both sides of the Panama 

land bridge, as follows:

Gastropod species of Gatun formation that survive on both sides of 

Panama land bridge

Architectonica (Architectonica} nobilis Roding

Colubraria obscura (Reeve)

Murex (Murex) recurvirostris Broderip

CHAGRES SANDSTONE, INCLUDING TORO LIMESTONE 

MEMBER

Locality 208 yielded the following gastropods described or mentioned in the present chapter. All are 

rare; that is, represented by one or two specimens.

Gastropods from Chagres sandstone at locality 208 

(CanceUaridce to Terebridce)

Cancellaria (Cancellaria) aff. C. epistomifera dariena Toula

Cancellaria (Cancellaria) sp.

Cancellaria (Pyruclia) diadela Woodring, n. sp.?

Trigonostoma n. sp.

Conus imitator Brown and Pilsbry?

Conus tortuosostriatus Toula

Pleuroliria (Polystira) ecuadoriana (Olsson)
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE

Occurrence elsewhere of gastropods from Gatun formation

323

Cydostremiscus (Ponocyctus) pentagonus (Gabb) __ .

Tunttella (Bactrospira) attilira praecellens Pilsbry

Arcltitectonica (ArcMttctonica) nobtiis nobilis Roding--

Siphocypraea (Muracypraea) henekeni (Sowerby) 8...

Murez (Mmex) recurvirostris recurvirostris Broderip.-

Paziella (Panamurex) gatunensis (Brown and 

Pilsbry).... .......................................

AnacMs (Costoanachis) mira fugax Brown and 

Pilsbry

Strombina (Strombina) lessepsiana Brown and 

Pilsbrv

Antillophos (AntUlophos) candei gatunensis (Toula)--

Fasciolaria gorgasiana Brown and Pilsbry, subsp -- 

Oliva (StrephoneJla) colpotus Woodring, n. narne 11 -.

Mitra (Concilia) dariensis Brown and Pilsbry H._._.

Prumtm (Microspira) gatunense (Brown and Pilsbry) 

Cancellaria (Cancellaria) epistomifera dariena Toula. 

Cancellaria (Cancellaria) epistomifera lipara WoodCancellaria (Pyruclia) cibarcola cibarcola Anderson.-

Aphera islacolonis (Maury)... -... _ -...-.-_.-. ...

Gatun formation

Lower part

X 

X 

X

X 

X 

X 

X

xxxxx

X

xxxx

X 

X

X 

X

X 

X

X 

X

X

X 

X 

X

X

X

X 

X 

X

X

X

X

X 

X

X

X

X 

X 

X

X 

X

X 

X 

X 

X

Middle part

XXX XXXXXX 

X 

X

X 

X 

X

X

X 

X

X 

X

X 

X

X 

X

X 

X

X 

X 

X

X

XXXXXXX X 

X 

X 

X

X

X 

X

XXXXXXXXX

X 

X

X 

X 

X

Upper part,eastern 

area

X 

X

X

X

X

X 

X

X

X

X

X

X

X

X 

X 

X

X

X 

X 

X

X 

X

X

X 

X

X 

X

Upper western part, 

area

X

X

X

X

X

X

X

X

X

X

X

X

Early 

Miocene

Chipola formation, 

Florida

------

0

A 

0

X

X

X

X

X

Middle Miocene

Southeastern Costa 

Rica

X 

X

X 

X

X

X

f

X

X

X

X

Cercado formation, 

Dominican Eepublic

X

X

X 

X

X 

X

X

X

X

X

X

X 

X

X

X

X

X

X

formation, Gurabo 

Dominican Bepublic

X

X

X 

X

X 

X

X

X

X

X

X

X 

X

X

X

X

Bow formation, den 

Jamaica

..

X

X

X 

X

X

X

X

X

X

X

X

Shoal B,iver 

Florida formation, 

X

X

Other 2

X

X

X

X 

X

X

X 

X

X

X

X

X

X 

X

X 

X

X

X

X

X

X

X

X

X

X

X

X 

X

X

X 

X 

X

X

X

X

X 

X

X

X

X

X

X

X

Late 

Miocene

formation, Limon 

Bica Costa 

X

X

X

X

X

?

X

X

X

X

X

Other 3

X

X

X

X

X

X

X

X

X

Early 

Pliocene

Caloosahatchee 

Florida formation, 

X

X

X

X

X

Other *

X

X

X

X

X

X

Late Pliocene

formation, Mofn 

Rica Costa 

X

X

X

Holocene

X

X

X 

X

X

X

X

X

See footnotes at end of table.
324 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Occurrence elsewhere of gastropods from Gatun formation Continued

Conus recognitus Guppy _ ........................

Conus musaensis Olsson ............................

Conus spurius Gmelin. .............................

Conus bravoi Spieker .. ... - . ------- .

Conus molis Brown and Pilsbry ---------

Conus symmetricus Sowerby. ------- . --------- . .

Conus imitator imitator Brown and Pilsbry . . . . . . .

Conus multiliratus multiliratus Bose .---.-... .. -

Conus burckhardti burckhardti Bose .-..--.-... -

Conus burckhardti harrisi Olsson. .._, . .............

Conus tortuosostriatus Toula.--.-- .... --.....--....

Oemmula vaningeni (Brown and Pilsbry) -----------

Oemmula machapoorensis (Maury) .... .... --------

Pleuroliria (Polystira) tenagos (Gardner) _ ----------

Pleurofusia (Cruziturricula) fusinus fusinus (Brown

Cochlespira (Ancistrosyrinx) cedonulli (Reeve). ------

Scobinella morierei (Cossmann). ....................

Drittia (Neodrillia) riogurabonis eurysoma Gardner. .. 

Crassispira (Hindsiclava) consols conscrs (Sowerby) . 

Clathrodrillia gatunensis (Toula) ..--....----..---...

Agladrillia (Agladrillia) acarii ectypha Woodring, n.

Eudathvrella (Miraclathurella) eucharis Woodring,

Terebra (Oreoterebra) subsulcifera subsulcifera Brown 

and Pilsbry. .-_______-_-.--.___ __ ..............

Terebra (Panaterebra) cucurrupiensis Oinomikado.-

Strioterebrum gausapatum (Brown and Pilsbry).-- ..

Acteon (Acteon) punctostriatus (C. B. Adams)...---

Ringicula (Ringiculella) semistriata d'Orbigny,... .. 

Acteocina bullata (Kierier). ..---... . ..... .

Cylichnella atacata stibara Woodring, n. subsp. .....

Retusa (Cylichnina) quercinensis biforis Pilsbry and 

Johnson __ ........ ----- .

Volvulella (Volvulella) micratracta Brown and

Volvulella (Volvulella) cylindrica parallela (Pilsbry

Gatun formation

part Lower 

X 

X 

X 

X

X 

X

X

X

X 

X

X 

X 

X 

X

X 

X 

X 

X

X 

X 

X

X 

X

X 

X 

X 

X 

X

X 

X 

X 

X

X 

X 

X 

X

X 

X

X 

X

Middle part

X 

X 

X 

X

X 

X 

X 

X 

X 

X

X

X 

X

X 

X 

X

X 

X 

X 

X 

X 

X

X 

X 

X

X

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X 

X

X 

X 

X

X

X 

X

X

eastern Upper part,

area

X

"x~~

X 

X 

X

X 

X

X

X 

X

X 

X

X

X

X

X

X 

X

X

X 

X

X 

X

X

X

Upper western part, 

area

X 

X 

X

X

X

X

X

X

X

X

Early 

Miocene

formation, Chipola 

Florida

X

Is

0

X

X

X

X

X

X

Middle Miocene

Southeastern Costa 

Rica

X

X

X

X

X

X

X

X

Cercado formation, 

Dominican Republic

X

X

X

X 

X

X

X

X

X

formation, Gurabo 

Dominican Republic

X

X

X 

X

X 

X

X

X

X

X

X

X

X

formation, Bowden 

Jamaica

X

X

X

X 

X

X

X

X

X

X 

X

X

X

River Shoal 

Florida formation, 

X

1o

X

X 

X

X 

X 

X

X 

X

X

X

X

X

X

X

X

X

X

X 

X

X

X

X

X

X

Late 

Miocene

Linon formation, 

Rica Costa 

X

X 

X

X 

X 

X

X

X

X

X

X

X

Is

o

X

X

X

X

X

Early 

Pliocene

Caloosahatchee 

Florida formation, 

X

X

| O

X

X

X

X

Late Pliocene

formation, Moin 

Rica Costa 

X

Holocene

X

X

X

X 

X

X

1 Early Miocene part of Uscari shale, southeastern Costa Rica; Baitoa formation, 

Dominican Republic; Thomonde formation, Haiti; Aymamon limestone, Puerto 

Rico; deposits of early Miocene age in Sinu area, Colombia; Las Perdices shale, 

Colombia; outcrop La Rosa formation, Venezuela; Subibaja formation, Ecuador; 

lower part of Zorritos formation, Peru.

2 Presumably Cercado or Gurabo formation, Dominican Republic (Gabb's stratigraphically unallocated species); Tubara formation, Colombia; subsurface La Rosa 

formation, Venezuela; middle Miocene part of Brasso formation, Trinidad; San Jose 

calcareous silt member of Manzanilla formation, Trinidad; Agueguexquite formation, 

Mexico; middle Miocene deposits in Chiriqui Province and Darien province, Panama; 

middle Miocene deposits in valley of Rio San Juan, southwestern Colombia; Angostura, Picaderos, Progreso, and Daule formations, Ecuador; Upper part of Zorritos 

formation, and Cardalitos and Montera formations, Peru.

3 Deposits of late Miocene age, Colombia; Punta Gavilan formation, Venezuela; 

Savaneta glauconitic sandstone member and Melajo clay member of Springvale formation, Trinidad; deposits of late Miocene age (Spencer's Coatzacoalcos formation), 

Tehuantepec area, Mexico; deposits of late Miocene age, including Olsson's Pinecrest

beds and Mansfield's Cancellaria zone, Florida; Duplin marl, North Carolina; 

Esmeraldas formation, Ecuador.

* Deposits of early Pliocene age, Bocas Islaid, Panama; Mare and Playa Grande 

formations, Venezuela; deposits of early Pliocene age in Cumana area, Venezuela; 

Matura formation, Trinidad; Charco Azul formation, southeastern Costa Rica and 

southwestern Panama.

5 Assigned to subgenus Torcula on p. 102 and 105, respectively.

e Described on p. 161 as Petaloconchus afl. P. floridanus.

7 Described on p. 81 as Trochita trochiformis.

8 Described on p. 194 as Cypraea (Muracypraea) henekeni.

9 Assigned to genus Hanetia on p. 256 and 257, respectively.

10 Described on p. 272 as Leptarius leptus.

n Described on p. 278 as Oliva (Strephonella) plicata.

13 Assigned to subgenus Tiara on p. 283 and 284, respectively.

13 Described on p. 286 as Xancus validus falconensis.

M Described on p. 287 as Valuta alfaroi eurytera.
GASTROPODS: EULIMIDAE, MARGENTELLIDAE TO HELMINTHOGLYPTIDAE 325

Pleurofusia cf. P. fenimorei (Bartsch)

Scobinella ecuadoriana Olsson

Carinodrillia (Carinodrillia) sp.

Darbya? (Buridrilllal) sp.

Grassispira (Hindsiclava) pyrgoma Woodring, n. sp.

Clathrodrilliaf sp.

Striotereftrum sp.

The gastropod fauna of the Chagres sandstone 

proper is meager, like that of the early Tertiary faunas 

under consideration: 30 species, seven of which are not 

named at the specific level. Seven of the unequivocally 

identified species are endemic and four others evidently 

are. The occurrence elsewhere and in the Gatun formation of the same or related species is tabulated below.

Gastropods from Chagres sandstone and occurrence elsewhere and in 

Gatun formation of same or related species

Species from Chagres sandstone

Calliostoma (Calliostoma)

metalium Woodring. 

Architectonica (Architectonica)

nobilis karsteni Rutsch.

Epitonium ("Depressiscala") 

eucteanum Woodring.

Sthenorytis toroensis euthynta 

Woodring.

Scalina weigandi (Bose) _____

Bathygalea (Miogalea) hadra

Woodring and Olsson. 

Stigmaulax guppiana (Toula)_.

Distorsio (Rhysema) decussata

gatunensis Toula. 

Ficus carbasea carbasea

(Guppy). 

Solenosteira dalli Brown and

Pilsbry, subsp.2

Cymatophos? acolus Woodring. 

Amarophos bothrus Woodring..

Latirus (Polygona) anapetes

Woodring. 

Valuta eurytera Woodring 3 __.

Cancellaria (Cancellaria) aff.

C. epistomifera dariena

Toula. 

Cancellaria (Pyruclia) diadela

Woodring, n. sp.? 

Conus imitator Brown and

Pilsbry? 

Conus tortuosostriatus Toula _.

Pleuroliria (Polystira) 

ecuadoriana (Olsson).

Pleurofusia cf. P. fenimorei 

(Bartsch).

Scobinella ecuadoriana Olsson.

Crassispira (Hindsiclava) 

pyrgoma Woodring, n. sp.

Occurrence elsewhere and in Gatun 

formation of same or related species

C. aurora Dall, Holocene.

Middle Miocene, Ecuador;

late Miocene, Venezuela,

Panama, Mexico. 

E. scipio (Dall). Holocene.

S. toroensis toroensis (Dall), 

Toro limestone member.

Late Miocene, Tehuantepec 

area, Mexico.

B. andersoni (Abbott), 1 early 

Miocene, Colombia.

Middle to late Miocene. Also 

Gatun formation.

Middle to late(?) Miocene. 

Also Gatun formation.

Early to late Miocene. Also 

Gatun formation.

Subspecies of S. dalli range 

from middle to late Miocene.

A. dentalis (Olsson), early 

Miocene, Costa Rica.

L. taurus Olsson, late Miocene, Panama.

V. alfaroi Dall, middle Miocene, Costa Rica. Also 

Gatun formation.

C. epistomifera dariena, middle to late Miocene.

Possibly C. diadela, Gatun

formation. 

Possibly C. imitator, middle to

late Miocene. 

Middle to late Miocene. Also

Gatun formation.* 

Late Miocene, Panama,

Ecuador. 

P. fenimorei, Holocene.

Late Miocene, Ecuador. 

C. consors (Sowerby), early 

Miocene to late Pliocene.

1 Recorded on p. 198 as Bathygalea dalli.

2 Assigned to genus Hanetia on p. 258.

3 Described on p. 287 as Valuta alfaroi eyrytera.

Though the Chagres formerly was considered to be 

of early Pliocene age, a late Miocene age is justified. 

Three species occur elsewhere exclusively in formations 

of that age and the age range of four others includes 

late Miocene. Five and possibly two others occur in the 

Gatun formation. None is exclusively Pliocene and the 

age range of none, except a related species, includes 

Pliocene. So far as known, however, three species are 

closely related only to living species. Miogalea, Amarophos, and ScobineUa are extinct.

Two of the Chagres turrids are found in the Esmeraldas formation of northern Ecuador, although the 

outcrop areas are 800 kilometers apart. Olsson (1964, 

p. 12) designated a late Neogene age for the Esmeraldas, but late Miocene is preferable. Perhaps in suggesting a correlation between the Chagres and Esmeraldas a comparable moderately deep-water f acies rather 

than age is being correlated. Nevertheless both formations overlie middle Miocene deposits. In three widely 

separated areas the Tehuantepec region of Mexico, 

northern Panama and northern Ecuador moderately 

deep-water formations seem to be of late Miocene age.

Only eight gastropods have been recovered from the 

Toro limestone member, the shallow-water basal part 

of the Chagres. Six, represented by molds, are aragonite-shelled, one of which is identified as Turritella 

altilira. The other two are calcite-shelled epitonids: 

Sthenorytis pemobilis? (a living species) and S. toroensis toroensis, which may occur in Pliocene deposits 

on Montserrate Island, in the Gulf of California.

HOLOCENE SERIES 

INFORMALLY DESIGNATED ATLANTIC AND PACIFIC MUCK

The informally designated Atlantic and Pacific 

muck is described on page 50. The following radiocarbon dates on black organic muck and wood were 

measured in the ITSGS laboratory: 

Radiocarbon dates on muck and wood from Atlantic and Pacific muck

Lab. 

No.

W-958

W-959

W-960

Location of sample

Core Hole BBR 53, west side of Pacific 

entrance to Panama Canal, 9 kilometers west-northwest of Point Farfan; depth 10 meters, 4.2 meters below 

top of muck, which is 9.9 meters thick. 

Black organic muck.

Core Hole BBR 128, east side of Pacific 

entrance to Panama Canal, 0.5 kilometer east of terminus of Thatcher 

Ferry; depth 22 meters, 9 meters below 

top of muck, which is 10.5 meters 

thick. Black organic muck.

Core Hole Mindi 2, Mindi Dairy Farm, 

6.4 kilometers south of Col6n; depth 

10.6 meters, 1.5 meters below top of 

muck, which is 3 meters thick. Wood.

Age (years 

before present)

6, 720 ±300

7, 680 ±300

7, 240 ±3 00
326 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

The muck formerly was thought to be of Pleistocene 

age, but according to the radiocarbon dates it was deposited during the postglacial rise of sea level.

Dall (1912) and Brown and Pilsbry (1913a) described a few species of mollusks from marine layers 

in the muck near Mount Hope on the Caribbean side 

of the Canal. Three of them are now known to be living off Payardi Island, 8 kilometers northeast of 

Colon: Minioliva myrmecoon (Dall), Stri-oterebrum 

spei (Brown and Pilsbry), and Adrana perprotracta 

(Dall). As pointed out by Altena (1968), A. newcombi 

(Angas) is an earlier name for Dall's species.

Mr. Stewart called my attention to a constructionperiod map based on some 200 shallow drill holes along 

and near the alinement of Gatun Dam. Contours drawn 

on the top of bedrock show a channel, filled with muck, 

west of the present channel of Rio Chagres, sloping 

northwestward to a depth of 300 feet (91.4 meters). 

Major (later Major General) W. L. Sibert, the engineer in charge of the construction of Gatun Dam, Spill1 

way, and Locks described the bedrock surface as follows: "The rock surface under the Gatun Locks was 

very uneven, being composed of hills and valleys; in 

the valleys the softest of mud was found, corroborating the statements of the geologists that a subsidence 

of more than 300 feet occurred at one time at Gatun" 

(Sibert, 1916, p. 392-393).

Other radiocarbon dates for muck in the Gatun Lake 

area, ranging in age from about 11,300 to 1,275 years 

before the present, have been published in a recent 

account that describes the core occurrence of wild maize 

pollen and at a higher level cultivated maize (Bartlett, Barghoorn, and Berger, 1969).

DESCRIPTION OF TERTIARY MOLLUSKS 

CONTINUED FROM CHAPTER C

GASTROPODS CONTINUED FROM CHAPTER C

Family EULIMIBAE 

The family Eulimidae was omitted in its proper place.

Genus Enlima Risso

Bisso, Histoire naturelle des principales productions de I'Burope 

meridionale, v. 4, p. 123, 1826.

Type (logotype, Herrmannsen, Indicis generum malacozoorum, 

v. 1, p. 431, 1847) : Turbo subulatus Donovan (=Strombiformis glaber da Costa), living, eastern Atlantic Ocean.

Eulima cf. E. jacksonensis Gregorio 

Plate 48, figure 8

Very small, slender, 9-whorled. Protoconch acute, tip 

slightly bulging. Post-protoconch whorls practically 

flat-sided. Suture indistinct. Aperture elongate, narrow, ovate.

Height (incomplete) 3.1 mm (estimated restored 

height 4 mm), diameter 1.1 mm (figured specimen).

The marine member of the Bohio (?) formation on 

Trinidad Island yielded five small specimens of 

Eulima, They resemble E. jacksonensis Gregorio 

(Palmer in Harris and Palmer, 1946-47, p. 224, pi. 26, 

fig. 16, 1947; middle and late Eocene, southeastern 

U. S.), but are about half as large, even when specimens having the same number of whorls are compared.

Occurrence: Marine member of Bohio (?) formation 

(late Eocene or early Oligocene), locality 42.

Eulima nobilis Guppy 

Plate 49, figure 1

Eulima (Liostraca) nobilis Guppy, in Guppy and Dall, U.S. 

Natl. Mus. Proc., v. 19, p. 315, pi. 30, fig. 9, 1896 (Miocene, Jamaica).

Melanella (Eulima) cercadica Maury, Bull. Am. Paleontology, 

v. 5, no. 29, pi. 25, fig. 1, 1917 (Miocene, Dominican Republic).

Strombiformis prcelubrica Pilsbry and Johnson, Acad. Nat. Sci. 

Phila. Proc., v. 69, p. 183, 1917 (Miocene, Dominican 

Republic). Pilsbry, Idem, v. 73, p. 395, pi. 35, fig. 10). 

1922 (Miocene, Dominican Republic).

Strombiformis ischna Gardner, U.S. Geol. Surrey Prof. Paper 

142, p. 573, pi. 55, fig. 17 (misprinted 16), 1947 (Miocene, 

Fla.).

Small, slender, 11- or 12-whorled. Protoconch acute, 

tip slightly bulging. Post-protoconch whorls practically flat-sided. Suture indistinct. Aperture elongate, narrow, ovate.

Height 5.9 mm, diameter 1.7 mm (figured specimen).

Type: Lectotype, herewith designated, USNM 

107071.

Type locality: Bowden, Jamaica, Bowden formation.

Eulima nobilis, represented by 35 specimens, is fairly 

widely distributed in the lower and middle parts of 

the Gatun formation, but is nowhere abundant. The 

Gatun shells are smaller than those from Jamaica, the 

Dominican Republic, and Costa Rica, which reach a 

height of 9.5, 10, and 8.5 millimeters, respectively.

The type lot of Guppy's misnamed species consists 

of four specimens. The lectotype is the largest and was 

illustrated. Maury's synonomy and discussion indicate 

that she intended Melanetta cercadica to be a substitute 

name for Gabb's Eulima acicularis, a homonym, although she did not have access to Gabb's type. Nevertheless, her expression "n. sp.", whether intentional or 

not, is the basis for accepting her name as the name 

for the species she illustrated. Two weeks after her 

name appeared, a properly proposed substitute name 

for Gabb's species was published. It is not the same as 

Maury's. (See the second following species.)
GASTROPODS: EULIMIDAE, MARGESTELLIDAE TO HELMINTHOGLYPTIDAE 327

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138, 

138a, 138b, 138c, 138d. Middle part, eastern area, localities 139b, 139c, 147b, 147h, 153a, 155 (identification 

doubtful), 159d. Cercado and Gurabo (USGS 8735) 

formations (middle Miocene), Dominican Kepublic. 

Bowden formation (middle Miocene), Jamaica. Middle 

Miocene deposits, Costa Kica (USGS 5882f and other 

localities). Shoal Kiver formation (middle Miocene), 

Florida.

Eulima acuta Sowerby

Plate 51, figure 13

Eulima acuta Sowerby, Zool. Soc. London Proc., p. 8, 1834

(living, eastern Pacific Ocean). 

Leiostraca acuta (Sowerby), Sowerby, Thesaurus conchy liorum,

p. 803, pi. 170, fig. 11 (misprinted 25 in text), 1854

(living, eastern Pacific Ocean). 

Strombiformis acuta (Sowerby), Bartsch, U.S. Natl. Mus. Proc.,

v. 53, p. 347, pi. 47, fig. 2 (enlargement of Sowerby's

illustration), 1917 (Sowerby's record). 

Eulima scotti Maury, Bull. Am. Paleontology, v. 4, no. 21, p. 30,

pi. 7, fig. 21, 1910 (Miocene, Florida). 

Strombiformis scotti (Maury), Gardner, U.S. Geol. Survey Prof.

Paper 142, p. 573, pi. 55, fig. 16 (misprinted 17), 1947

(Miocene, Florida). 

Strombiformis ischnon Pilsbry and Johnson, Acad. Nat. Sci.

Philadelphia Proc., v. 69, p. 183, 1917 (Miocene, Dominican Republic). Pilsbry, Idem, v. 73, p. 395, pi. 35, fig. 8,

1922 (Miocene, Dominican Republic).

Small, very slender, 12-whorled. Protoconch acute, 

tip slightly bulging. Post-protoconch whorls practically flat-sided. Suture indistinct. Aperture elongate, narrow, ovate.

Height 7.3 mm, diameter 1.2 mm (figured specimen).

Type: Presumably in British Museum (Natural 

History).

Type locality: Living, Montijo Bay, Pacific Coast 

of Panama.

A small, very slender Eulima from the middle part 

of the Gatun formation is identified as the species 

ranging from North Carolina to the West Indies, for 

which the name E. acuta, has been used, for example 

by Ball and Simpson (1901, p. 413). Though no eastern Pacific specimens are available, that name is 

adopted. In any event, the fossil and living species is 

not E. auricincta (Abbott) (1958, p. 106, fig. 5; Grand 

Cayman Island), which is less slender.

The type of E. ischnon and specimens from the Cercado formation at USGS locality 8525 are smaller and 

more slender than the Gatun fossil, but others from 

both the Cercado (USGS 8521) and Gurabo (USGS 

8734) formations closely resemble that fossil.

Occurrence: Middle part of Gatun formation (middle Miocene), eastern area locality 159d. Chipola formation (early Miocene), Florida. Cercado and Gurabo

formations (middle Miocene), Dominican Kepublic. 

Middle Miocene deposits, Costa Kica (USGS 5882 m). 

Deposits of late Miocene age, Florida. Caloosahatchee 

formation (Pliocene), Florida. Living, North Carolina 

to West Indies; Pacific coast of Panama.

Eulima sarissiformis (Pilsbry and Johnson) 

Plate 49, figure 2

Eulima acicularis Gabb, Am. Philos. Soc. Trans., n. ser., v. 15, 

p. 227, 1873 (Miocene, Dominican Republic). Not EuUma 

acicularis A. Adams, 1861.

Strombiformis sarissiformis Pilsbry and Johnson, Acad. Nat. 

Sci. Phila. Proc., v. 69, p. 183, 1917 (Miocene, Dominican 

Republic) ; new name for Eulima acicularis Gabb. Pilsbry, Idem, v. 73, p. 394, pi. 35, fig. 9, 1922 (Miocene, 

Dominican Republic).

Relatively large, slender. Protoconch and early postprotoconch whorls missing. Remaining post-protoconch 

whorls practically flat-sided. Suture distinct or moderately distinct. Aperture elongate, narrow ovate.

Height (incomplete) 11.5 mm (estimated restored 

height 14 mm), diameter 2.7 mm (figured specimen).

Type: Acad. Nat. Sci. Phila. 3010.

Type locality: Dominican Republic, Miocene.

This relatively large species is represented by five 

specimens from the lower part of the Gatun formation. The protoconch and early post-protoconch whorls 

are missing on all. Comparable, but smaller, species 

are living in both western Atlantic and eastern Pacific 

waters: Eulima rectiuscula (Dall) (1890-1903, p. 160, 

1890) and E. towensendi (Bartsch) (1917, p. 340, pi. 

46, fig. 4), respectively.

An incomplete, partly corroded shell from the upper 

part of the Gatun formation in the western area is 

listed as Eulima sp.

Occurrence: Lower part of Gatun formation (middle 

Miocene), localities 138, 138c, 138d. Miocene (presumably Cercado or Gurabo formation), Dominican Republic.

Genus Balcis Leach

Leach, Annals and Mag. Nat. History, v. 20, p. 271, 1847. 

Type (monotype): Balcis montagui Leach (= Strombiformis 

albus da Costa), eastern Atlantic Ocean.

The western Atlantic species of the genus in the 

collections of the U.S. National Museum are not satisfactorily identified and the eastern Pacific species are 

overnamed.

Submenus Balcis s.s.

Balcis (Balcis) jacululum (Maury) 

Plate 51, figures 1, 2

Melanella (Eulima) jacululum Maury, Bull. Am. Paleontology, 

v. 5, no. 29, p. 143, pi. 25, fig. 3, 1917 (Miocene, Dominican Republic).

368-278 O - 70 - 3
328 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Eulima robusta Gabb, Am. Philos. Soc. Trans., n. ser., v. 15, 

p. 227, 1873 (Miocene, Dominican Republic). Not Eulima 

robusta A. Adams, 1861.

Mclanella astuta Pilsbry and Johnson, Acad. Nat. Sci. Philadelphia Proc., v. 69, p. 182, 1917 (Miocene, Dominican 

Republic) ; new name for Eulima robusta Gabb. Pilsbry, 

Idem, v. 73, p. 394, pi. 35, fig. 7, 1922 (Miocene, Dominican Republic).

Small, slender, about 8-wliorled. Protoconch missing. 

Post-protoconch whorls practically flat-sided. Suture 

fairly distinct. Body whorl rounded. Penult whorl 

practically flat or faintly angulated just above bodywhorl suture. Aperture short, ovate.

Height (not quite complete) 3.3 mm, diameter 1.1 

mm (figured specimen).

Type: Cornell University.

Type locality: Bluff 3, Cercado de Mao, Rio Mao, 

Dominican Republic, Cercado formation.

Six small, slender fossils in five lower and middle 

Gatun collections are identified as Balds jacululum. 

They are considerably smaller than topotypes of that 

species, which reach a height of 5.5 mm. Maury emphasized the "overhanging whorls," that is, the faint 

angulation of the penult whorl just above the bodywhorl suture. That feature, however, is not consistently 

shown by topotypes or Gatun shells. The trivial name 

evidently is a noun in apposition.

Three additional specimens that have a wider apical 

angle are doubtfully identified (localities 138c and 

139c). They may represent B. maoica (Maury, 1917, 

p. 142, pi. 25, fig. 2), should that be a valid species.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 137, 

138, 138c (identification doubtful). Middle part, 

eastern area, localities 139b, 139c (identification doubtful), 147b, 159d. Cercado formation (middle Miocene), 

Dominican Republic.

Balcis (Balcis) lipara Woodring, n. sp. 

Plate 51, figure 14

Of medium size, slender, about 12-whorled. Outline 

of spire faintly concave. Protoconch acute. First two 

or three and last two post-protoconch whorls slightly 

bulging, remainder practically flat-sided. Suture 

distinct. Aperture short, ovate.

Height (not quite complete) 8.3 mm, diameter 2.5 

mm (type).

Type: USNM 646076.

Type locality 138 (USGS 16909, North and south 

sides of Transisthmian Highway, 1.6 kilometers northeast of Canal Zone boundary, Panama), lower part of 

Gatun formation.

Balcis lipara is represented by 13 specimens from 

the lower and middle parts of the Gatun formation.

It is characterized by its faintly concave spire and 

slightly bulging last two whorls. Those features distinguish it from B. jamaicensis (C. B. Adams) (Clench 

and Turner, 1950, p. 296, pi. 36, fig. 5; living, Jamaica) 

and a similar species in USNM collections from localities between Massachusetts and Florida. This Atlantic 

coast species seems to be the same as that named B. 

bartschi (Gardner and Aldrich) (1919, p. 37, pi. 2, fig. 

1) for late Miocene fossils and B. calkinsi (Olsson and 

Harbison) (1953, p. 332, pi. 59, fig. 5) for Pliocene 

fossils.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138, 

138a, 138c. Middle part, eastern area, localities 139c, 

147b, 147g.

Balcis (Balcis) aulaca Woodring, n. sp. 

Plate 51, figure 12

Of medium size, slender, about 12-whorled. Protoconch and earliest post-protoconch whorls missing. 

Suture deeply impressed. Early whorls practically flatsided; remaining whorls slightly bulging. Base of all 

except first few spire whorls beveled adjoining suture. 

Aperture short, ovate; outer lip broken back. Columellar lip thin.

Height (not quite complete) 7.8 mm, diameter (not 

quite complete) 2.2 mm (type).

Type: USNM 646077.

Type locality: 138 (see preceding species), lower 

part of Gatun formation.

Though the outer lip of the type the only specimen is defective, Balcis aulaca is a distinctive species. 

In suture and beveling adjoining the suture it is similar to the next species, but the shell is smaller and 

more slender, and the whorl height is greater.

Occurrence: Lower part of Gatun formation (middle 

Miocene), locality 138.

Balcis (Balcis) cetia Woodring, n. sp. 

Plate 49, figure 8

Very large, slender, about 18-whorled, whorl height 

low. Protoconch missing. Post-protoconch whorls 

slightly bulging. Suture deeply impressed. Base of all 

except first few spire whorls beveled adjoining suture. 

Aperture short, ovate. Columellar lip thick.

Height (not quite complete) 28.3 mm, diameter 8.7 

mm (type).

Type: USNM 646078.

Type locality: 138f (USGS 23663, South side of 

Transisthmian Highway, hillside excavation at Colchoneria Yero, about 450 meters southwest of Cativa, 

Panama), lower part of Gatun formation.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 329

Fourteen specimens of this large Niso-like species were 

found in the lower and middle parts of the Gatun formation. It is a representative of a closely knit group of 

Miocene species that are characterized by their large 

size, low whorl height, and deeply impressed suture. 

This group of four species left no known descendants. 

The earliest and smallest (height 20 mm) of these 

species occurs in the Thomonde formation of Haiti 

(USGS 9946), the last and largest in the Savaneta 

glauconitic sandstone member and Melajo clay member 

of the Springvale formation of Trinidad, both of late 

Miocene age: Balds egregia (Guppy) (Jung, 1969, p. 

465, pi. 47, figs. 1, 2; height 50 mm). The fourth species, B. maUsta (Gardner) (1926-47, p. 574, pi. 72, 

figs. 6-8, 1947), occurs in the middle Miocene of 

Florida. Like B. cetia, it is of intermediate size (estimated height 23 mm). It is, however, more slender 

than the Gatun species and the base of its spire whorls 

is not bevelled.

B. dalli (Bartsch) (1917, p. 302, pi. 35, fig. 5; Gulf 

of California) is a large species (height 20 mm), but 

its outline, suture, and whorl height are like those of 

smaller species.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138c, 

138d, 138e, 138f. Middle part, eastern area, localities 

139b, 139c.

Genus Niso Risso

Risso, Histoire naturelle des principales productions de I'Europe

meridionale, v. 4, p. 218, 1826. 

Type (monotype) : Niso eburnea Risso, Pliocene, France and

Italy.

Submenus Niso s.s.

Niso (Niso) umbilicata (Lea)? 

Plate 48, figure 7

Of medium size, moderately inflated, about 10- 

whorled. Protoconch missing. Spire whorls low, 

practically flat-sided to slightly bulging. Body whorl 

rounded to faintly angulated. Umbilicus partly exposed, apparently narrow. Peristome missing.

Height (incomplete 11.2 mm (estimated restored 

height 15 mm), diameter 5.5 mm.

The marine member of the Bohio(?) formation at 

the submerged Vamos Vamos locality and Palenquilla 

Point yielded eight specimens doubtfully identified as 

Niso umbilicata. The identification is doubtful only 

because these fossils are defective. So far as observable 

features go they closely resemble that species, which 

ranges through the Eocene in southeastern United 

States (Palmer, 1937, p. 66, pi. 6, figs. 22-25; Palmer 

in Harris and Palmer, 1946-47, p. 225, pi. 26, fig. 17, 

1947). No comparable Oligocene species is known in 

that area.

Occurrence: Marine member of Bohio (?) formation 

(late Eocene or early Oligocene), localities 40a, 40d, 

41, 41b.

Niso (Niso) mesata Woodring, n. sp.

Plate 49, figure 5, plate 51, figure 3

Large,, moderately inflated, about 15-whorled. Protoconch acute. Post-protoconch whorls low, first few 

slightly bulging, later whorls practically flat-sided, 

last few slightly bulging. Intermediate whorls weakly 

angulated at base adjoining suture, angulation fainter 

or absent on last few whorls, including corresponding 

part of body whorl. Occasional microscopic axial 

threads on some specimens. Umbilicus moderately wide, 

bordering ridge subdued. Basal lip missing.

Height (incomplete) 14 mm (estimated restored 

height 18 mm), diameter (incomplete) 6.4 mm (type). 

Height (not quite complete) 21.1 mm, diameter (not 

quite complete) 8 mm (paratype).

Type: USNM 646079; paratype USNM 646080.

Type locality: 138c (USGS 21956, About 100 meters 

north of Transisthmian Highway and about 75 meters 

west of road to refinery site on Payardi Island, Panama ; immediately east of Cativa and 100 meters north 

of locality 138), lower part of Gatun formation.

Locality of paratype: 142 (Stanford Univ. 2698, 

Northeast of Fort Gulick, latitude 9° 20' N., longitude 

79° 52' W., plus 1,010 feet [310 meters], middle part 

of Gatun formation.

Though 61 specimens of Niso mesata are in collections from the three parts of the Gatun formation, 80 

percent are immature, including all from six localities. 

The early and intermediate whorls of the paratype 

the largest specimen are worn. The type shows the 

angulation of intermediate whorls.

Several similar species, differing chiefly in apical 

angle, are found in the Miocene of the Caribbean 

region. N. grandis Gabb (Pilsbry, 1922, p. 394, pi. 34, 

fig. 17; Dominican Republic) and its probable 

synonym, N. striatula Bb'se (in Bose and Toula, 1910, 

p. 227, pi. 12, fig. 7; Tehuantepec area) have the widest 

apical angle, and a species from the Cercado farmation 

of the Dominican Kepublic (USGS 8534) the narrowest. (The type of N. grandis is a fragment of four 

whorls. A 10-whorled specimen, lacking a few early 

whorls (height 23.8 mm) has been found in the Gurabo 

formation at USGS locality 8519.) The apical angle 

of N. mesata is of intermediate width; wider, however, than that of a smaller species from the Bowden 

formation of Jamaica and the Limon formation of the 

Bocas del Toro area.

N. mesata is more closely related to the Panamic N. 

excolpa Bartsch (1917, p. 348, pi. 48, fig. 4) than to
330 GEOLOGY AND PALEONTOLOGY OP CANAL ZONE

any western Atlantic species. The whorl height of the 

Panamic species is greater, and its late spire whorls 

and body whorl are angulated.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 137, 138a, 138c, 138d. Middle part, eastern area, 

localities 142, 147b, 147g, 147h, 159d; western area, 

locality 161. Upper part, eastern area, locality 175.

Family MARGINELLIDAE

Unlike most of the families of gastropods under consideration, the family Marginellidse has a richer representation in the Carribbean Sea than in eastern Pacific 

waters.

In addition to the species described or mentioned, 

unidentified marginellids, or probable marginellids, 

are in collections from the Caimito and Culebra formations.

Genus Marginalia Lamarck

Lamarck, >Soc. Histoire Nat. Paris Me'm., p. 70, 1799.

Type (monotype) : Valuta glabella Linne, living, west Africa.

Subgenus Eratoidea Weinkauff

Weinkauff, Systematisches Conchylien-Cabinet von Martini und

Chemnitz, v. 5, pt. 4, p. 140, 1879. 

Type (logotype, Cossmann, Essais paleoconchologie comparee,

pt. 3, p. 87, 1899) : Marginella margarita Kiener, living,

West Indies.

Classification of the numerous fossil and living marginellids still is unsatisfactory. Eratoidea seems to be 

better than Serrata (Jousseaume, 1875, p. 167, 230-232; 

type (tautotype) : Marginella serrate Gaskoin, living, 

Mauritius) for high-spired Caribbean species, although 

Serrata was used in 1928 (Woodring, 1928, p. 239). 

Should the objective be smaller groups, the recently 

proposed name Ebumospira (Olsson and Harbison, 

1953, p. 201; type (orthotype) : Marginella ebumeola 

Conrad, Miocene, Virginia, and North Carolina) is 

available.

Marginella (Eratoidea) aff. M. mollitor Ball

Plate 48, figures 15, 16

Small, slender or moderately inflated. Spire high, 

about 2/5 of height of shell. Later half of body whorl 

slightly shouldered. Outer lip greatly thickened, 

sharply limited. Inner edge of outer lip coarsely denticulate. Space between posteriormost denticle and insertion of lip relatively wide. Columella bearing four 

folds.

Estimated restored height 10.5 mm, diameter 5.7 mm 

(figured specimen). Estimated restored height 11 mm, 

diameter 6 mm (largest specimen).

Three damaged specimens are referred to this 

species: one from the upper part of the Bohio formation on Barro Colorado Island and two from the 

Caimito formation of the Gatun Lake area. The Bohio 

specimen, the best of the lot, is illustrated. One of the 

Caimito specimens is mature. It is a little larger and a 

little more inflated than the Bohio specimen, and has 

also a wider outer lip.

The greatly thickened outer lip is the most distinctive feature of this unnamed species. Otherwise it is 

similar to early Miocene species from Florida: Marginella mollitor Dall (1915, p. 52, pi. 12, fig. 1; Tampa 

formation) and M. euancycla Gardner (1926-47, p. 388, 

pi. 46, fig. 20, 1938; Chipola formation). M. hematita 

Kiener (1834-41, p. 11, pi. 7, fig. 31, 1834) is the most 

similar of the living Caribbean species. Aside from its 

thinner outer lip, it tends to have a lower spire and a 

more distinctly shouldered body whorl than the Panama fossils. No comparable living Panamic species has 

been recognized.

Two poorly preserved small specimens reveal the 

presence of a species of Eratoidea in the La Boca formation (locality 99b). It has a lower spire than M. aff. 

M. mollitor.

Occurrence: Upper part of Bohio formation (late 

Oligocene), locality 42d. Caimito formation, Gatun 

Lake area (late Oligocene), locality 56.

Genus Volvarina Hinds

Hinds, Zool. Soc. London, Proc., pt. 12, p. 75, 1844. 

Type (logotype, Redfield, Am. Jour. Conchology, v. 6, p. 221, 

1870) : Marginella nitida Hinds, living, locality unknown.

Hinds' statement that Marginella avena Valenciennes 

is a typical species of Volvarina is not a type designation, although it has been accepted doubtfully as such 

(Woodring, 1928, p. 236). Marginella nitida was based 

on a specimen (or specimens) of unknown habitat in 

the Cuming collection. Sowerby's (1847, p. 389, pi. 76, 

fig. 131) illustration of that species presumably represents the type. Tomlin (1917, p. 284) thought that M. 

nitida is the Mediterranean species M. mitrella Risso. 

In any event, according to Sowerby's illustration, Volvarina is properly used for a small group of slender 

species that have an exposed spire, slightly or moderately thickened, nondenticulate or faintly denticulate 

outer lip, and four columellar folds. Hyalina (Schumacher, 1817, p. 234; type (monotype) : H. pellucida 

Schumacher, living, locality unknown) is also in use 

for that group of species. Without type material or an 

illustration of the type, however, Hyalina is considered 

a nomen dubium. Despite statements to the contrary 

(Tomlin, 1917, p. 288; Woodring, 1928, p. 237), Schuinacher's species was based on a specimen. Though he 

cited an illustration in Martini's Conchylien-Cabinet, 

he went on to remark that Martini's illustration is
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 331

smaller than his shell, but is much better depicted than 

by an illustration in a publication by Schroter.

Volvarina leander (Brown and Pilsbry) 

Plate 51, figures 4-6

Marginella leander Brown and Pilsbry, Acad. Nat Sci. Phila.

Proc., v. 63, p. 347, pi. 24, fig. 13, 1911 (Miocene, Canal

Zone). Olsson, Bull. Am. Paleontology, v. 9, no. 39, p.

98, pi. 6, fig. 22, 1922 (Miocene, Canal Zone). 

Marginella aff. nitida Hinds, Toula, K. k. Geol. Reichsanstalt

Jahrb., v. 61, p. 504, pi. 31, fig. 18, 1911 (Miocene, Canal

Zone).

Of medium size, slender. Spire low, thinly glazed. 

Outer lip somewhat thickened, sharply bordered. Inner 

edge of outer lip smooth or faintly denticulate. Aperture narrow. Columellar lip bearing four slender folds. 

Parietal callus moderately thick.

Height 8 mm, diameter 3.7 mm (small figured specimen). Height 9 mm, diameter 4.4 mm (figured specimen of intermediate size), height 11.4 mm, diameter 

5.2 mm (large figured specimen).

Type: Acad. Nat. Sci. Phila. 1708.

Type locality: Gatun Locks excavation, Canal Zone, 

middle part of Gatun formation.

This marginellid is widespread in the middle and 

upper parts of the Gatun formation, including the 

upper part in the western area, but was not found in 

the lower part. It is most abundant in old collections 

from the middle part along the Panama Railroad 

south and southeast of Gatun; for example, some 150 

specimens were collected at locality 147b. None from 

that locality and nearby has a height greater than 

about 8 millimeters. One of these small fossils is shown 

on plate 51, figure 6. A specimen of average size has a 

height of 9 millimeters (pi. 51, fig. 5). The largest 

(pi. 51, fig. 4) is in a collection from the upper part of 

the Gatun near Mount Hope. With the exception of a 

relatively inflated shell in the same collection (height 

8.2 mm, diameter 4.2 mm), the range of variation in 

degree of inflation is very slight. Toula's Marginella 

aff. M. nitida is a typical example of Volvarina leander.

V. collina (Olsson) (1922, p. 97, pi. 7, figs. 26, 27), 

a middle Miocene Costa Rican species, is a little larger, 

a little more slender, and has a slightly higher, more 

thickly glazed spire. The living Caribbean species V. 

avena (Valenciennes) (in Kiener, 1834-41, p. 17, pi. 6, 

fig. 24, 1834) is a little larger than F. collina, has a 

higher spire, and lacks the sharply bordered outer lip 

of F. collina and F. leander. V. avena has been recognized in deposits of late Miocene age in Costa Rica 

(Olsson, 1922, p. 97, pi. 7, figs. 21, 28). F. tceniolata 

Morch (Coan and Roth, 1966, p. 287, pi. 50, figs. 48-55), 

which ranges from southern California to Costa Rica?,

has the outline of F. leander, but its outer lip is not 

sharply bordered.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

localities 139b, 139e (identification doubtful), 146, 

147b, 147f, 147g, 147h, 151, 153a (identification doubtful), 155 (identification doubtful), 155b, 155c, 157; 

western area, locality 161a. Upper part, eastern area, 

localities 173, 177, I77b, 177c, 177d, 178 (identification 

doubtful); western area, localities 183, 185 (identification doubtful).

Genus Prunum Herrmannsen

Herrmannsen, Indicis generum malacozoorum, v. 2, supplement, 

p. 113, 1852.

Type (monotype and tautotype) : Valuta prunum Gmelin, living, West Indies.

At the present time it is appropriate to use Prunum 

at the generic level for a large group of maginellids, 

the largest group of fossil species in the Caribbean 

region. The species assigned to Prunum in the unrestricted sense are of medium to large size, more or 

less inflated, and low-spired. Their aperture is narrow 

or wide; the outer lip is sharply bordered and is smooth 

or denticulate. The parietal wall and the ventral face 

of the spire are free of callus or covered with callus. 

The columellar lip bears four folds.

The subgenus Prunum s.s. is a small group. The 

species have a wide aperture and nondenticulate outer 

lip. The spire and parietal wall have the usual glaze 

of marginellids, but are not callused. The thickened 

outer lip tapers off abruptly toward the posterior end. 

The type species has a distinct notch, formed by the 

thinned continuation of the lip.

Subgenus Microspira Conrad

Conrad, Am. Jour. Conchology, v. 4, p. 66, 1868. 

Type (monotype) : Volutella (Microspira) ovifortms Conrad, 

Miocene, Virginia.

The type specimen of the type species was illustrated 

by Gardner (1926-47, p. 397, pi. 48, figs. 11, 12, 1938). 

It has a very low spire, denticulate outer lip, and thin 

parietal callus. Egouena (Jousseaume, 1875, p. 167, 

192-214; type (tautotype); Egouena egouen Jousseaume = Marginella amygdala Kiener, living, west Africa) is available for species that have a nondenticulate 

outer lip and callus completely encircling the aperture. 

Leptegouana (Woodring, 1928, p. 237; type (orthotype) : Voluta guttata Dillwyn, living, West Indies) 

was proposed for species characterized by no callus 

on the spire except in line with the weakly denticulate 

outer lip. Both names, however, can be suppressed 

without any great loss.
332 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Prunum (Microspira) aff. P. apalachee (Gardner) 

Plate 48, figure 25

Relatively small, moderately inflated, spire relatively 

high. Outer lip moderately thickened, relatively narrow, its inner edge smooth. Parietal callus thin.

Height 10.2 mm, diameter 5.9 mm (figured specimen) .

The La Boca formation at locality 116a near Paraiso 

yielded a small specimen of Prunum. Though it is not 

well preserved, it evidently represents a small species 

allied to P. apalachee (Gardner) (1926-47, p. 395, pi. 

47, fig. 5, 1938), of the early Miocene Chipola formation of Florida. The spire of the La Boca species is, a 

little lower and its outer lip is narrower.

P. aff. P. apalacliee is doubtfully recognized at locality 99b.

Occurrence: La Boca formation (early Miocene), 

localities 99b (identification doubtful), 116a.

Prunum (Microspira) gatunense (Brown and Pilsbry) 

Plate 51, figures 10, 11

Marginella gatunensis Brown and Pilsbry, Acad. Nat. Sci.

Phila. Proc., v. 63, p. 347, pi. 24, fig. 10, 1911 (Miocene,

Canal Zone). 

Marginella coniformis Sowerby, Brown and Pilsbry, Idem, p.

348, pi. 24, fig. 12, 1911 (Miocene, Canal Zone). 

Marginella (Bullata) mindiensis Cossmann, Jour. Conchyliologie, v. 61, p. 61, pi. 5, figs. 13-15, 1913 (Miocene, Canal

Zone).

Marginella mindiensis Cossmann, Olsson, Bull. Am. Paleontology, v. 9, no. 39, p. 96, pi. 6, figs. 16, 23, 1922 (Miocene,

Canal Zone). 

Marginella latissima, pilslryl Olsson, Idem, p. 96, pi. 10, fig. 2,

not fig. 1, 1922 (Miocene, Canal Zone, Costa Rica).

Of medium size, moderately to strongly inflated, spire 

extending well above posterior end of outer lip. Aperture very narrow. Outer lip strongly thickened, its 

inner edge weakly denticulate. Callus extending from 

outer lip over much, or exceptionally all, of ventral 

face of spire, thickly covering parietal wall and thinning somewhat on columellar lip. Outer edge of callus 

generally fading out on parietal wall and columellar 

lip, exceptionally sharply defined.

Height 17.8 mm, diameter 12 mm (larger figured 

specimen).

Type: Acad. Nat. Sci. Phila. 1706.

Type locality: Gatun Locks excavation, Canal Zone, 

middle part of Gatun formation.

Unfortunately the type of Marginella gatunensis is 

immature (height 11.4 mm), whereas the type of J/. 

mindiensis is mature.

Though Prunum gatunense is of modest size, it is 

the largest marginellid in the Gatun formation. It

occurs in the lower, middle, and upper parts, but is 

represented by a total of only 23 specimens. The outline is variable. The type of P. mindiense and Olsson's 

illustrated specimens referred to that species are 

moderately inflated, like the immature type of P. 

gatunense and the immature shell shown on plate 51, 

figure 10. Brown and Pilsbry's P. coniformis is mature 

and inflated. Olsson's illustrated P. latissima pilsbryi 

from the middle part of the Gatun in the western area 

is strongly inflated, but not as strongly inflated as the 

illustrated specimen from Rio Banana, Costa Rica, 

wT hich is herewith designated the lectotype. Other 

specimens from the western area are not so strongly 

inflated.

According to Crossmann's illustrations, the type of 

P. mindiense has exceptionally thick and sharply 

limited callus. A moderately inflated specimen (height 

20.3 mm, diameter 12 mm) from locality 157 has such 

callus. Not only is it thick and sharply limited, it 

extends to the tip of the spire and covers the entire 

apertural face of the spire, as on Cossmann's type.

Prunum coniforme (Sowerby) (Pflug, 1961, p. 55, 

pi. 15, figs. 2-6, 8, 9: Miocene, Dominican Kepublic, 

Jamaica) is larger, less inflated, less heavily callused, 

and has a lower spire. P. precursor (Dall) (1890-1903, 

p. 47, pi. 5, fig. 4, 1890; Pliocene, Florida) also has 

a lower spire. P. latissimum (Dall) (in Guppy and 

Dall, 1896, p. 308, pi. 29, fig. 11; Miocene, Dominican 

Kepublic; Pliocene, Costa Kica) is smaller, more inflated, and its aperture is dilated near the base of the 

outer lip. P. storerium (Couthouy) (1837, p. 440, pi. 

9, figs. 1, 2), now living along the Caribbean coast of 

Panama, is less inflated and its outer lip is smooth. 

Coan and Koth's (1966, p. 280, pi. 48, figs. 7, 8) illustrations of an unnamed shell dredged off Costa Rica indicate that a species similar to P. gatunense is living in 

eastern Pacific waters. As they noted, the inner edge of 

the outer lip of Brown and Pilsbry's Marginella coniformis (that is, P. gatunense} is straighter than that 

of their species.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 138c, 138d. Middle part, eastern area, localities 

139c, 139d, 157, 160d; western area, localities 161, 161b. 

Upper part, eastern area, localities 171, 175 (immature). Middle Miocene, Costa Eica. Middle Miocene, 

Chiriqui area, Panama (USGS 7955).

Genus Persicula Schumacher

Schumacher, Essai d'un nouveau systeme des habitations des 

vers testaces, p. 235, 1817.

Type (monotype and tautotype) : Persicula varlaUlls Schumacher (=Voluta persicula Linne), living, west Africa.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 333

Subgenus Gibberula Swainson

Swainson, A treatise on malacology, p. 323, 1840.

Type (monotype) : Gibberula sonata Swainson (= Volvaria

oryza Lamarck), living, west Africa. 

Gibberula is low-spired, whereas Persicula s.s. has 

no spire; in fact, the type has a slight apical depression.

Persicula (Gibberula) cf. P. semen (Lea) 

Plate 48, figure 3

Small, inflated or moderately slender, spire low. 

Aperture very narrow. Outer lip thickened, but not 

sharply bordered. Inner edge of outer lip finely denticulate. Siphonal canal notched. Columellar lip bearing five or six folds.

Height 6.5 mm, diameter 4 mm (figured inflated 

specimen). Restored height about 6.Y mm, diameter 3.6 

mm (moderately slender specimen).

Two silicified small marginellids from the Gatuncillo 

formation are similar to Persicula semen, a middle and 

late Eocene species of southeastern United States 

(Palmer, 193Y, p. 422, pi. 6Y, figs. 13, 14, 16-19). They 

have a slightly higher spire and narrower aperture 

than P. semen. One of the fossils from the Canal Zone 

is inflated and the other is fairly slender. The preservation of both is poor, as the granular silica is too coarse 

for satisfactory preservation of such small fossils.

Occurrence: Gatuncillo formation (middle Eocene), 

locality 38.

Sugenus Rabicea Gray

Gray, Guide to the systematic distribution of Mollusca in the

British Museum, pt. 1, p. 37, 1857. 

Type (monotype) : Persicula interrupta (Lamarck) (Marginella

interrupta Lamarck=FoZ«£a interruptolineata Megerle),

living, west Africa and Caribbean Sea.

The heavy, more or less bifid second (from anterior 

end) fold on the columellar lip, callus spread over the 

entire parietal wall, and absence of an apical depression distinguish Rabicea from Persicula s.s.

Persicula (Rabicea) venezuelana amydra Woodring, n. subsp. 

Plate 48, figure 26

Small, strongly inflated, apex slightly below level of 

posterior end of outer lip. Aperture very narrow. Outer 

lip wide, weakly bordered by low ledge. Inner edge of 

outer lip finely denticulate. Siphonal canal notched. 

Five folds visible on columellar lip; second (from 

anterior end) widest, heaviest, indistinctly bifid, extending out on body whorl beyond lip. Parietal callus 

moderately thick. Corroded shells showing 20 to 22 

narrow, closely spaced spiral bands, representing color 

bands that have lost their color, or the spaces between 

color bands.

Height Y.6 mm, diameter 4.9 mm (type).

Type: USNM 645860.

Type locality: 116a (USGS 20956, east bank of 

Panama Canal at Canal station 18YO near Paraiso, 

Canal Zone; YOO feet (213 meters) southeast of locality 

116), La Boca formation.

This marginellid is identified in three collections 

from the La Boca formation and one from the Culebra. 

It is abundant only at the type locality, which yielded 

30 specimens. Most of these fossils from all four localities are more or less corroded and show the spiral 

bands mentioned in the description. Faint color bands 

and corresponding false sculpture of corroded shells 

are not unusual in fossil species of Persicula. They 

have been observed on shells of the Eocene species P. 

semen (Lea) (Palmer, 193Y, p. 423) and on shells of P. 

venezuelana venezuelana (F. Hodson, in Hodson, Hodson, and Harris, 192Y, p. YY).

P. venezuelana amydra agrees with P. venezuelana 

venezuelana (F. Hodson) (in Hodson, Hodson, and 

Harris, 192Y, p. YY, pi. 40, figs. 13, 14) in essential 

features, but is less inflated and a little smaller. The 

nominate subspecies occurs in strata of early Miocene 

age in Venezuela. A larger middle and late Miocene 

subspecies in Venezuela has been named P. venezuelana 

lavelana (F. Hodson) (in Hodson, Hodson, and Harris, 

192Y, p. Y8, pi. 40, figs. 3, 10, 11; Eutsch, 1934, p. 91, 

pi. 6, figs. 9-11, 12).

P. venezuelana is more inflated and has a less distinctly bordered outer lip than P. interruptolineata 

(Megerle) (Kiener, 1834-41, p. 25, pi. 5, fig. 21, 1834; 

as Marginella interrupta Lamarck), the type of 

Rabicea. Though those two species are similar, P. cercad.ensis (Maury) (191Y, p. Y3, pi. 11, fig. Y), a small, 

slender middle Miocene species from the Dominican 

Republic, perhaps more closely resembles the type 

species. Despite its small size (height Y mm), P. cercademis has a wide, sharply bordered outer lip, like 

the type species. P. interruptolineata is living in both 

west African and Caribbean waters. A comparable, if 

not identical, species, P. imbrwata (Hinds) (Coan and 

Both, 1966, p. 283, pi. 48, fig. 18, pi. 49, figs. 19-33), 

is found in the Panamic region.

Occurrence: Culebra formation (early Miocene), 

locality 106. La Boca formation (early Miocene), localities 114, 115b, 116a.

Persicula (Rabicea) couviana stenygra Woodring, n. subsp. 

Plate 51, figure 7

Of medium size, strongly inflated, apex about at level 

of posterior end of outer lip. Aperture moderately narrow. Outer lip narrow, weakly bordered by a low ledge. 

Denticles on inner edge of outer lip fairly strong. 

Siphonal canal notched. Columellar lip bearing seven
334 GEOLOGY AND PALEONTOLOGY OP CANAL ZONE

folds; second (from anterior end) widest, heaviest, 

weakly bifid, extending out on body whorl; fifth to 

seventh progressively fainter in that order. Parietal 

callus thick. Faint spiral color bands visible on later 

half of body whorl under glaze of enamel.

Height 10 mm, diameter 6.8 mm (type).

Type :USNM 645711.

Type locality: 138c (USGS 21958, about 100 meters 

north of Transisthmian Highway and about 75 meters 

west of road to refinery site on Payardi Island, Panama ; immediately east of Cativa and 100 meters north 

of locality 138), lower part of Gatun formation.

The type, found in the lower part of the Gatun formation, is the only mature specimen of this marginellid. Four minute immature shells, one of which is 

associated with the type, presumably are to be assigned 

to it. Nevertheless they are doubtfully identified, as 

a growth series is unavailable. Two of the immature 

specimens, both from the middle part of the Gatun, 

are corroded and show about 12 spiral bands.

Persicula couviana stenygra is less inflated and a 

little smaller than P. couviana couviana (Maury) 

(1925, p. 202, pi. 34, fig. 11), and has a narrower outer 

lip. The outer lip of both subspecies is weakly bordered. 

The nominate subspecies occurs in the late Miocene 

Savaneta glauconitic sandstone member of the Springvale formation of Trinidad. The outer lip of the type 

of P, couviana couviana, and also of the type of its 

junior synonym P. propeobesa (Mansfield) (1925, p. 

41, pi. 6, fig. 10) is broken back beyond the thickened 

margin.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138a 

(immature, identification doubtful), 138c. Middle part, 

western area, locality 170 (immature, identification 

doubtful).

Family CANCELLARIIDAE

Cancellarids present a great array of form and 

sculpture. Classification of such diverse species is difficult and so far unsatisfactory. Ample precedent is 

available for both conservative treatment and more 

narrowly restricted genera and subgenera.

The cancellarid fauna of the Panamic region is much 

richer than that of the Caribbean region. As in many 

other families, the roots of the present Panamic cancellarid fauna lie in the enlarged Miocene Caribbean 

province, which embraced western Atlantic and eastern 

Pacific waters. Perplicaria clarki M. Smith (1947, p. 

55, pi. 2, fig. 9; Olsson and Bergeron, 1967) and the 

loosely coiled Trigonostoma milleri Burch (1949) are 

perhaps the most remarkable Recent cancellarids. Both 

are Panamic species and both have Miocene predecessors in the present Caribbean region or in Florida. The

fossil species of Perplicaria have been reviewed by 

Wilson (1948). His upper Miocene collections from 

peninsular Florida include a loosely coiled Trigonostoma allied to T. milleri, recently described as T. 

hoerlei (Olsson, 1967, p. 24, pi. 8, figs. 6, 6a).

The Gatun formation contains 18 species and subspecies of cancellarids in nine genera and subgenera. 

Four of the genera and subgenera (Pyruclia, Euclia, 

Narona, and Aphera) are no longer living in the 

present Caribbean region, but survive in the Panamic 

region. Unidentified cancellarids, or doubtful cancellarids, are represented by poorly preserved material in 

the marine member of the Bohio(?) formation, the 

upper part of the Bohio formation of Barro Colorado 

Island, the Caimito formation of the Gatun Lake area, 

and the La Boca formation.

Genus Cancellaria Lamarck

Lamarck, Soc. Histoire Nat. Paris Mem., p. 71, 1799. 

Type (monotype): Valuta reticulata Linne, living, Cape Hatteras to Caribbean Sea.

Subgenus Cancellaria s. s.

Cancellaria (Cancellaria) anomoia Woodring, n. sp. 

Plate 52, figures 1, 2

Small, moderately inflated, spire distinctly turreted. 

Protoconch relatively large, naticoid, 3^4-whorled, end 

marked by abrupt appearance of sculpture. First 

sculptured whorl bearing 11 axial ribs and seven relatively wide, closely spaced, flat spiral threads. Sculpture of late whorls consisting of axial ribs, basically 

narrow or of moderate width (18 to 21 on body whorl), 

and closely spaced, flat spiral threads or narrow cords 

(seven on penult). On shoulder of body whorl and 

near base a secondary spiral thread in some of the 

spaces between primary threads. A widened varix-like 

rib or two present on body whorl; present or absent 

on preceding two whorls. Outer lip broken back; ribs 

and growth lines showing a shallow notch near base.

Height 19.1 mm, diameter 11.7 mm (type). Height 

(not quite complete) 19.8 mm, diameter 13.6 mm 

(largest specimen).

Type: USNM 645712.

Type locality: 138c (USGS 21956, about 100 meters 

north of Transisthmian Highway and about 75 meters 

west of road to refinery site on Payardi Island, Panama ; immediately east of Cativa and 100 meters north 

of locality 138), lower part of Gatun formation.

The type of Cancellaria anomoia was found in the 

lower part of the Gatun formation at locality 138c. 

The same locality yielded a minute shell, consisting of 

the protoconch and 2% sculptured whorls, and three 

imperfect shells, one of which is a little larger than the
GASTROPODS: ETJLIMIDAE, MARGINELLIDAE TO HELMESTTHOGLYPTIDAE 335

type. That the type is immature is indicated by its 

size and also by the narrow space between the edge of 

the columellar lip and the siphonal fasciole.

The sculpture, consisting of closely, and essentially 

uniformly, spaced flat spiral threads (or narrow cords) 

and axial ribs, which are basically narrow or of moderate width, gives this species a characteristic appearance. The spiral sculpture suggests that of C. rowelli 

Dall, which is discussed under the next heading. Both 

spiral and axial sculpture of C. anomoia, however, 

continue with undiminished strength beyond the stage 

when they are subdued on C. rowelli. C. anomoia is 

more similar to C. defuniak Gardner (1926-47, p. 365, 

pi. 44, figs. 1, 2, 1938), a middle Miocene species from 

Florida, but is not as slender and has less sharply 

chiseled spiral sculpture.

Occurrence: Lower part of Gatun formation (middle 

Miocene), locality 138c.

Cancellaria (Cancellaria) aff. C. macneili Mansfield 

Plate 52, figures 3, 4

Small, moderately inflated, spire distinctly turreted. 

Protoconch worn, naticoid, 21^-whorled, end marked 

by abrupt appearance of sculpture. First sculptured 

whorl somewhat worn, bearing 10 axial ribs, a spiral 

thread (later two) on upper part of whorl, and three 

low, flat spiral threads on main part of whorl. Early 

half of penult whorl sculptured with 15 narrow axial 

ribs, a doubled spiral thread adjoining suture, another 

spiral thread on upper part of whorl, and four closely 

spaced, low, flat spiral threads (or narrow cords) on 

main part of whorl. On later part of penult and on 

body whorl axial ribs greatly subdued. At same stage 

spiral sculpture disappears, except in narrow band in 

sutural area and in wider band adjoining siphonal 

fasciole. Sculptured spire whorls, except first, bearing 

a widened varixlike rib.

Height 19 mm, diameter 12 mm (figured specimen).

The lower part of the Gatun formation yielded a 

small weakly sculptured Cancellaria and a fragment of 

a somewhat larger specimen, also weakly sculptured. 

Both of these fossils doubtless are immature. The outline and strong suppression of sculpture suggest relationship to C. macneili (Mansfield, 1937a, p. 609, pi. 

85, figs. 1, 4; middle Miocene, Florida), if not that 

species. The early sculpture of C. Icevescens Guppy 

(Woodring, 1928, p. 220, pi. 12, figs. 7, 8; middle Miocene, Jamaica), is reticulate and stronger than that of 

G. macneili, whereas C. rowelli Dall (in Guppy and 

Dall, 1896, p. 307, pi. 29, fig. 1; middle Miocene, Dominican Kepublic) is more slender and its sculpture is 

not as strongly suppressed on late whorls.

Occurrence: Lower part of Gatun formation (middle 

Miocene), localities 136, 137.

Cancellaria (Cancellaria) tapeina Woodring, n. sp. 

Plate 51, figures 8, 9

Of medium size, moderately inflated, spire distinctly 

turreted. Protoconch missing, early sculptured whorls 

worn. Earliest preserved sculpture consisting of axial 

ribs and eight low, flat spiral threads. On later part of 

penult whorl and on body whorl axial ribs more closely 

spaced and subdued; spiral threads (or narrow cords) 

subdued, and an intervening secondary spiral thread 

appearing. Penult whorl bearing 29 axial ribs and 12 

primary spiral threads. Slightly widened ribs, which 

appear on last three whorls, take the place of varices. 

Outer lip broken back; ribs and growth lines showing 

a shallow notch near base.

Height (almost complete) 34.7 mm, diameter 20.5 

mm (type).

Type :USNM 645714.

Type locality: 182a (USGS 8488, Caribbean coast 

east of San Miguel [Rio Miguel], station 25 plus 400 

feet (120 meters), Panama), upper part of Gatun 

formation.

Cancellaria tapema is based on a somewhat worn 

specimen, found in the upper part of the Gatun formation in the western area. Though the weak sculpture 

suggests C. rowelli, this cancellarid evidently is not a 

form of that species. The low, flat spiral threads of C. 

tapeina are separated by wider spaces, in which a 

secondary thread appears on the last whorl and a half. 

The body-whorl axial ribs of the Gatun fossil also are 

less subdued.

C. anomoia, C. aff. C. macneili and C. tapeina are 

more similar to C. reticulata than the other cancellarids 

of the Canal Zone and adjoining parts of Panama. 

Nevertheless the relationship is not close.

Occurrence: Upper part of Gatun formation, western area (middle Miocene), locality 182a.

Cancellaria (Cancellaria) epistomifera dariena Toula

Plate 52, figures 11-18

Cancellaria dariena Toula, K. k. Geol. Reichsanstalt Jahrb., v. 

58, p. 703, pi. 28, fig. 2, 1909 (Miocene, Canal Zone). 

Brown and Pilsbry, Acad. Nat. Sci. Philadelphia Proc., 

v. 63, p. 345, pi. 24, figs. 3, 4, 1911 (Miocene, Canal Zone). 

Pilsbry and Brown, Idem, v. 69, p. 32 (list), 1917 (Miocene, Colombia). Olsson, Bull. Am. Paleontology, v. 9, 

no. 39, p. 80, pi. 6, fig. 8, 1922 (Miocene, Canal Zone, 

Panama, Costa Rica). Weisbord, Idem, v. 14, no. 54, 

p' 50, pi. 6, fig. 8, 1929 (Miocene, Colombia). Anderson, 

California Acad. Sci. Proc., 4th ser., v. 18, no. 4, p. 115, 

1929 (Miocene, Colombia). Jung, Bull. Am. Paleontology, 

v. 49, no. 223, p. 549, pi. 75, fig. 4, 1965 (Miocene, Canal 

Zone).
336 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Cancellaria dariena n. sp., var., Toula, K. k. Geol. Reichsanstalt 

Jahrb., v. 58, p. 704, pi. 28, fig. 1, 1909 (Miocene, Canal 

Zone).

Cancellaria dariena trachyostraca Brown and Pilsbry, Acad. 

Nat. Sci. Philadelphia Proc., v. 63, p. 345, pi. 24, figs. 1, 2, 

1911 (Miocene, Canal Zone).

Cancellaria darienensis Toula, Cossmann, Jour. Conchyliologie, 

v. 61, p. 51, pi. 4, figs. 9, 10, 1913 (Miocene, Canal Zone).

Cancellaria epistomifera Guppy, Cossmann, Idem, p. 53, pi. 4, 

figs. 5, 6, 1913 (Miocene, Canal Zone).

Uxia mioccenica Cossmann, Idem, p. 54, pi. 4, figs. 11, 12, 1913 

(Miocene, Canal Zone).

Cancellaria (Cancellaria) dariena Toula, Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst, p. 118, 1964 (Miocene, Ecuador).

fCancellaria (Cancellaria) dariena Toula, Oinomikado, Geol. 

Soc. Japan Jour., v. 46, p. 623, pi. 29, fig. 19, 1939 (Miocene, Colombia).

Not Cancellaria? cf. C. dariena n. sp., Toula, K. k. Geol. Reichsanstalt Jahrb., v. 58, pi. 25, fig. 13, 1909 (Miocene, 

Canal Zone) ;= AntillopJios candei gatunensis (Toula).

Of medium size, inflated to moderately slender, spire 

slightly to strongly turreted. Protoconch naticoid, 3- 

whorled, end marked by gradual appearance of sculpture. First sculptured whorl bearing four or five spiral 

threads and generally 10 or 11 (exceptionally 14) 

widely spaced axial ribs. Terminal varix of varying 

strength present or absent on mature shells. Body 

whorl of mature shells generally bearing one or two 

earlier varices (exceptionally none or as many as 

three). One or two varices exceptionally present on 

penult whorl or even on antepenult. Penult whorl 

of mature shells sculptured with four or five spiral 

cords (exceptionally six) and 23 to 25 axial ribs (exceptionally as many as 28). Spiral cord adjoining 

suture narrower and closer to adjoining cord than 

others. Width of spiral cords and spacing of axial ribs 

variable. Narrow cords and widely spaced ribs most 

common. A secondary spiral thread generally present in 

at least some of spaces between spiral cords, especially 

on and above periphery. Basal part of outer lip flaring 

and bearing a shallow notch. Siphonal f asciole strongly 

inflated. Parietal callus moderately thick.

Height 34 mm, diameter (including strong terminal 

varix) 21.5 mm (figured inflated specimen). Height 31 

mm, diameter (including weak terminal varix) 16.5 

mm (figured slender specimen).

Type material: Lectotype (herewith designated), 

Tech. Hochschule, Vienna.

Type locality: Presumably Gatun Locks site, Canal 

Zone, middle part of Gatun formation.

Cancellaria epistomifera dariena is the most widely 

distributed cancellarid in the Gatun formation and is 

locally abundant, especially in the lower part of the 

formation at locality 138c, where about 100 specimens 

were collected. Its distribution in the middle and

upper Miocene deposits of the western part of the 

present Caribbean province is fairly wide. Though it 

was not found in the upper part of the Gatun in the 

western area, two incomplete worn shells, listed as C. 

aff. C. epistomifera dariena, were found in the Chagres 

sandstone.

A pronounced range of variation, in outline and 

sculpture, is shown by the fossils referred to C. epistomifera dariena. The greatest range in outline is 

afforded by the samples from the lower part of the 

Gatun. An inflated specimen (pi. 52, figs. 17, 18), a 

slender specimen (pi. 52, figs. 15, 16), and a distinctly 

turreted specimen (pi. 52, figs. 13, 14), were selected 

from the large collection just mentioned. That collection includes 13 distinctly turreted shells. A strongly 

turreted specimen (pi. 52, figs. 11, 12), in a Stanford 

University collection from the lower part of the Gatun, 

is associated with six slightly turreted shells. No other 

shell is so strongly turreted. It is so different from the 

ordinary run of C. epistomifera dariena that it suggests a different species and is perhaps to be treated as 

such. It represents, however, only a step from the 

distinctly turreted form. One distinctly turreted shell 

was found in the middle part of the formation and 

none in the upper part. The illustrated slender specimen (pi. 52, figs. 15, 16) is exceptionally slender. The 

lectotype (the specimen shown on Toula's plate 28, 

figure 2) is moderately inflated and slightly turreted. 

It has a height of 27 mm and a diameter of 15.2 mm. 

Its flat siphonal fascicle and slender columellar folds 

indicate that it is not quite mature.

The range of variation in spacing of axial ribs, in 

width of spiral cords, and in strength of secondary 

spiral threads (or their absence) is too great to show 

with a reasonable number of illustrations. Some of the 

range is shown on plate 52 for specimens from the 

lower part of the Gatun. Toula's and Brown and Pilsbry's illustrations serve the same purpose for the middle part, and Cossmann's for the upper part in the eastern area. Five of some 250 shells in the collections at 

hand lack secondary spiral threads. At the other extreme, one (locality 138c) has exceptionally strong 

secondary threads.

C. epistomifera epistomifera Guppy occurs in the 

Cercado and Gurabo formations of the Dominican 

Republic. A lectotype, presumably from the Gurabo 

formation, was selected and illustrated by Pflug (1961, 

p. 52, pi. 14, figs. 1, 2, 9). Its protoconch is not preserved. Maury (1917, p. 63) pointed out that two types 

of protoconch and first post-protoconch whorl are 

represented in her collections: (1) protoconch small, 

axial ribs appearing at beginning of first post-protocoiich whorl; (2) protoconch large, axial ribs appear-
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 337

ing after first third of first post-protoconch whorl. 

According to collections in the U.S. National Museum, 

her first type is shown by specimens from the Cercado 

formation and her second type by specimens from the 

Gurabo formation. 0. epistomifera dariena shows the 

first type, although some variation in the size of the 

protoconch is apparent. The contrast in the two types 

of protoconch is shown in illustrations of a specimen 

from the Gurabo formation and one from the Gatun 

formation (Jung, 1965, pi. 75, figs. 3, 4, respectively).

The chief difference between C. epistomifera dariena 

and the nominate subspecies lies in the greater range 

of variation, in outline and sculpture, of the Canal 

Zone subspecies.

The subgenus Cancellaria s.s. is an appropriate assignment for C. epistomifera, although that species 

formerly (Woodring, 1928, p. 221) was referred to the 

subgenus later named SivetieUa by Wenz (1943, p. 

1356; type (orthotype: Cancellaris similis Sowerby, living, west Africa).

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 136, 136a, 137, 138, 138a, 138b, 138c, 138d, 138e, 

138f. Middle part, eastern area, localities 139c, 139e, 

139f, 140, 147 (identification doubtful), 147f (identification doubtful), 154 (identification doubtful), 155, 

155a, 155b, 155c, 157, 159, 159b, 159d; western area, 

localities 161, 161a, 161c, 170 (identification doubtful). 

Upper part, eastern area, localities 172, 175, 176, 176a, 

177 (identification doubtful), 177b, 178. Middle Miocene, Costa Rica, Colombia, Darien area (USGS 8477), 

Chiriqui area (USGS 7955). Limon formation (Late 

Miocene), Bocas del Toro area, Panama. Miocene, 

Ecuador. Chagres sandstone (late Miocene), locality 

208 (C. aff. C. epistomifera dariena).

Cancellaria (Cancellaria) epistomifera lipara Woodring, 

n. subsp.

Plate 52, figures 7, 8

Smaller than Cancellaria epistomifera dariena, inflated, spire somewhat turreted. Axial ribs closely 

spaced, 25 to 33 on penult whorl. Secondary spiral 

threads absent. Terminal varix and a nearby earlier 

varix subdued, or both absent. Notch on basal part of 

outer lip very shallow or absent.

Height 28 mm, diameter 17.5 mm (type). Estimated 

restored height 30 mm, diameter 18 mm (largest specimen).

Type :USNM 645719.

Type locality: 183 (USGS 8487, Caribbean coast 

east of Eio San Miguel [Eio Miguel], station 4 plus 40 

feet (12 meters), Panama), upper part of Gatun formation.

Cancellaria epistomifera lipara is represented by 17 

specimens from the type locality and one from locality 

182a. Both localities are in the upper part of the Gatun 

formation in the western area. In marked contrast to 

the diversity shown by C. epistomifera dariena, these 

18 cancellarids are fairly uniform. The combination of 

relatively small size, closely spaced axial ribs, absence 

of secondary spiral threads, subdued varices (or their 

absence), and very shallow notch (or its absence) on 

the outer lip distinguish them from G. epistomifera 

dariena and C. epistomifera epistoinifera. Despite their 

small size, the largest specimens of C. epistomifera 

lipara, like the small form of 0. epistomifera epistomifera in the Gurabo formation of the Dominican Republic, have a swollen siphonal fasciole and heavy 

columellar folds, indicating that they are mature.

The Limon formation, of late Miocene age, on Cayo 

de Toro, in the Bocas del Toro area of northwestern 

Panama (USGS 8326), yielded one specimen of C. 

epistomifera lipara (height 31 mm), associated with 

four of C. epistomifera dariena.

The axial ribs of an immature shell (height 16.3 

mm) in the large collection from the lower part of the 

Gatun formation at locality 183c are even more closely 

spaced than those of C. epistomifera lipara, and so are 

the spiral cords. This fossil is doubtfully identified as 

an aberrant form of C. epistomifera dariena.

Occurrence: Upper part of Gatun formation, western area (middle Miocene), localities 182a, 183. Limon 

formation (late Miocene), Bocas del Toro area, 

Panama.

Cancellaria (Cancellaria) apimela Woodring, n. sp. 

Plate 52, figures 5, 6

Small, slender, spire distinctly turreted. Protoconch 

missing, early sculptured whorls corroded. Earliest 

preserved sculpture consisting of axial ribs and eight 

low, flat spiral cords. Axial ribs strong and widely 

spaced on last two whorls. Penult whorl bearing 12 or 

13 axial ribs and nine low, flat spiral cords, separated 

by narrow grooves. Spiral sculpture subdued on main 

part of body whorl. One or two axial ribs on last two 

whorls wider and lower than others.

Height 22.6 mm, diameter 13 mm (type).

Type:USNM 645720.

Type locality: 182a (USGS 8488, Caribbean coast 

east of San Miguel [Rio Miguel] station 25 plus 400' 

feet (120 meters), Panama, upper part of Gatun 

formation.

This distinctive species is based on two specimens 

from the upper part of the Gatun formation in the 

western area. Part of both shells the part exposed 

when they were collected is corroded.
338 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Though the low flat spirals, which are separated by 

narrow grooves and are subdued on the body whorl, 

recall those of Cancellaria rowelli, the combination of 

such spiral sculpture with strong, widely spaced axial 

ribs and slender outline is unusual.

Occurrence: Upper part of Gatun formation, western 

area (middle Miocene), localities 182,182a.

Cancellaria (Cancellaria) species

An incomplete and badly damaged specimen of Cancellaria was found in the Chagres sandstone at locality 

208. It is small and slender, like C. apimela. Also like 

that species, it has strong, widely spaced axial ribs. 

Unlike that species, however, the spiral cords are 

widely spaced and are strong even on the body whorl. 

The penult whorl is sculptured with 10 axial ribs and 

six spiral cords.

Occurrence: Chagres sandstone (late Miocene), 

locality 208.

Submenus Pyrudia Olsson

Olsson, Bull. Am. Paleontology, v. 19, no. 68, p. 160, 1932. 

Type (orthotype): Cancellaria solida Sowerby, living, Gulf of 

California to Peru.

The earliest species of Pyrudia are of middle Miocene age and the survivors live in the eastern Pacific 

Ocean. Miocene representatives are found along or near 

the south border of the present Caribbean Sea in 

Panama, Colombia, Venezuela, and Trinidad and also 

in the Darien area, Ecuador and Peru. The latest species in the present Caribbean region occur in deposits 

of late Miocene age.

Cancellaria (Pyruclia) cibarcola cibarcola Anderson 

Plate 52, figures 9, 10; plate 53, figures 8, 10-12

Cancellaria cibarcola Anderson, Calif. Acad. Sci. Proc., 4th ser., 

v. 18, no. 4, p. 116, pi. 14, figs. 1-3, 1929 (Miocene, Colombia). Barrios, Colombia Servico Geol. Nac. Bol. Geol., 

v. 6, nos 1-3 (Informe 1082), p. 290, pi. 11, fig. 7, 1960 

(Miocene, Colombia).

Not Cancellaria cibarcola Anderson, Rutsch, Naturforsch. Gesell. Basel Verb., v. 54, p. 163, pi. 9, figs. 6a, 6b, 1942 

(Miocene, Trinidad) ', C. auriculaperta Vokes.

Of medium size to large, slightly to moderately 

shouldered. Spire of varying height, slightly to distinctly turreted, strongly inflated. Protoconch naticoid, 

2i/^-whorled. Early sculpture consisting of closely 

spaced axial ribs and low spiral threads. Sculpture 

gradually disappearing on third to fifth whorl later 

than protoconch, except faint spiral threads in sutural 

area and weak spiral threads on basal part of body 

whorl. Ribbed whorls later than first generally bearing 

a rib (exceptionally two) wider than others. Growth 

lines showing shallow notch on lower part of outer lip.

Height (practically complete) 49.2 mm, diameter 

(incomplete) 30 mm (exceptionally large, high-spired, 

figured specimen). Height (practically complete) 35.7 

mm, diameter (incomplete) 25 mm (moderately large, 

relatively low-spired, figured specimen).

Type: Calif. Acad. Sci. 4643.

Type locality: Between Chorrera and Cibarco, Dept. 

of Atlantico, Colombia, Tubara formation.

Cancellaria cibarcola cibarcola is fairly widespread 

in the lower part of the Gatun formation and is rare 

in the middle part. It is abundant at locality 138c, 

where 40 specimens were collected. The collection from 

that locality is unique in including 11 high-spired 

shells and exceptionally large shells. The largest, which 

is also high-spired, is illustrated (pi. 53, figs. 10, 12). 

The largest of 22 specimens in eight other collections 

is a little larger than that shown on plate 53, figures 8, 

11, and all are relatively low-spired. The type and 

other available shells from Columbia also are relatively 

low-spired. On at least some of the Colombian fossils 

the space between the edge of the columellar lip and 

the siphonal fascicle is a little wider than on Gatun 

specimens. These Colombian fossils have also a slightly 

more abruptly inflated siphonal fascicle and a slightly 

channeled suture. Specimens from the Darien area 

closely resemble those from the Gatun formation.

C. pycta Olsson (1964, p. 122, pi. 21, figs. 3, 3a) is 

considered to be a middle Miocene Ecuadoran subspecies of C. cibarcola, distinguished by the persistence of sculpture to a slightly later stage.

A late Miocene species from Trinidad, 0. auriculaperta Vokes (1938, p. 22, figs. 19, 20), is the most similar described species in the present Caribbean region. 

C. cibarcola is less distinctly turreted, and has an unchanneled or less distinctly channeled suture, and narrower axial ribs on early whorls. Rutsch, however, 

thought that C. auriculaperta is C. cibarcola. In general features C. cibarcola is similar to C. solida (Keen, 

1958, p. 441, fig. 704), the type of Pyruclia, but is 

more distinctly shouldered and has a higher spire.

Occurrence: Lower and possibly middle parts of 

Gatun formation (middle Miocene). Lower part, localities 136a, 137, 137a, 138, 138a, 138b, 138c, 138d, 138e, 

138f. Middle part, eastern area, locality 159d; western 

area, locality 161c (identification doubtful). Middle 

Miocene, Colombia, Darien area (USGS 8433, 8476, 

8493), Panama.

Cancellaria (Pyruclia) diadela Woodring, n. sp. 

Plate 53, figures 7, 9

Moderately large, pyriform, broadly and strongly 

shouldered, strongly inflated, low-spired. Suture deeply 

channeled on late whorls. Early sculpture not well pre-
GASTROPODS: EULIMIDAE, MARGESTELLIDAE TO HELMINTHOGLYPTIDAE 339

served, but consisting of axial ribs and spiral threads. 

Sculpture of late whorls reduced to faint spiral threads 

in sutural area and weak spiral threads on basal part 

of body whorl. Basal part of outer lip bearing a shallow notch.

Height (practically complete) 39.5 mm, diameter 33 

mm (type).

Type :USNM 645724.

Type locality: 182 (USGS 8408, Caribbean coast 

east of San Miguel [Rio Miguel], station 25 plus 600 

feet 150 meters, Panama), upper part of Gatun formation.

The broadly and strongly shouldered, pyriform outline, and low spire of Cancellaria diadela distinguish 

it from other species of the subgenus Pyruclia. C. scheilei Anderson (1929, p. 115, pi. 10, figs. 1-4), from the 

middle and upper (?) Miocene of Columbia, is the most 

similar described species, but is not as strongly shouldered or as pyriform.

Unfortunately, like another species of Cancellaria 

from the upper part of the Gatun formation in the 

western area (C. tapeina), C. diadela is based on only 

one specimen. A fragmentary shell from the Chagres 

sandstone is doubtfully referred to it.

Occurrence: Upper part of Gatun formation, western area (middle Miocene), locality 182. Chagres sandstone (late Miocene), locality 208 (identification doubtful).

Subgenus Euclia H. and A. Adams

H. and A. Adams, The genera of Recent Mollusca, v. 1, p. 277,

1854. 

Type (logotype, Cossmann, Essais de paleoconchologie comparee, pt. 3, p. 10, 1899): Cancellaria cassidiformis Sowerby, living, Gulf of California to Peru.

Euclia, like Pyruclia, occurs in the Miocene deposits 

of the present Caribbean region, but survives in eastern 

Pacific waters. The Miocene distribution has the same 

pattern as that of Pyruclia: Panama, Colombia, Venezuela, Trinidad, Darien area, Ecuador, and Peni. The 

survival pattern in the present Caribbean region also is 

like that of Pyruclia. The latest species there are of late 

Miocene age.

Cancellaria (Euclia) codazzii Anderson 

Plate 54, figures 3, 4, 7, 8, 11, 12

Cancellaria codazsii Anderson, Calif. Acad. Sci. Proc., 4th ser., 

v. 18, no. 4, p. 116, pi. 14, figs. 4-7, March 29, 1929 (Miocene, Colombia). Barrios, Colombia Servicio Geol. Nac., 

Bol. Geol., v. 6, nos. 1-3 (Informe 1082), p. 291, pi. 11, 

fig. 5, 1960 (Miocene, Colombia).

Cancellaria karsteni Anderson, Calif. Acad. Sci. Proc., 4th ser., 

v. 18, no. 4, p. 114, pi. 10, figs. 7-9, 1929 (Miocene, 

Colombia).

Cancellaria hettneri Anderson, Idem, p. 114, pi. 10, figs. 5, 6, 

1929 (Miocene, Colombia).

Cancellaria (Euclia) maldonadoi Olsson, Neogene molluskg 

from northwestern Ecuador, Paleontological Research 

Inst, p. 122, pi. 21, figs. 5, 5a, 1964 (Miocene, Ecuador).

Of medium size, moderately slender to moderately 

inflated. Body whorl angulated to subrounded at shoulder. Protoconch naticoid, 21/^-to 3-whorled, end marked 

by gradual appearance of sculpture. First sculptured 

whorl bearing 10 to 12 axial ribs and three or four 

spiral threads. First sculptured whorl or two angulated 

or subangulated at shoulder; remaining spire whorls 

up to penult rounded; penult gradually angulated or 

remaining rounded; body whorl sharply angulated to 

subrounded. Spire sculpture in general coarsely reticulate. Penult bearing six or seven spiral threads. Thread 

at shoulder almost invariably stronger than others. 

With few exceptions, axial ribs more widely spaced on 

later part of penult and body whorl; T to 16 (generally 

10 to 13) on body whorl and as many as 20 to 24 to a 

whorl at stage preceding wide spacing. Subduing of 

ribs on body whorl exceptional. Ribs drawn out into 

spines of varying length at shoulder on body whorls 

that are angulated. Secondary spiral thread appearing 

in some of spaces between primary threads, especially 

near shoulder and near siphonal fascicle. A widened 

rib present or absent on body whorl and preceding 

sculptured whorls.

Height (practically complete) 33 mm, diameter 20.7 

mm (figured slightly angulated, slightly spinose specimen). Height (practically complete) 35.6 mm, diameter 24 mm (figured moderately angulated, moderately 

spinose specimen).

Type: Calif. Acad. Sci. 4645.

Type locality: Between Chorrera and Cibarco, Dept. 

of Atlantico, Colombia, Tubara formation.

This variable species is widespread and locally abundant in the lower part of the Gatun formation and rare 

in the middle part. Some 140 specimens, including 54 

from locality 138c, are available in 10 collections from 

the lower part and 11 in four collections from the 

middle part. Among mature shells those that have a 

slightly or moderately angulated body whorl and 

slightly or moderately spinose ribs (pi. 54, figs. 3, 4, Y, 

8) are most abundant; those that are sharply angulated 

and strongly spinose (pi. 54, figs. 11,12) are less abundant; and those characterized by a subrounded outline 

and nonspinose ribs are rare. None of the few specimens in the third group is suitable for illustrating. 

Ribs on mature body whorls are almost invariably 

widely spaced and the spiral thread at the shoulder is 

almost invariably stronger than the others, invariably 

so on angulated whorls. It should be noted that the 

three forms Anderson named were found in strata of 

middle Miocene age in Colombia and two of them
340 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

occur together (Cancellaria karsteni and C. hettneri). 

C. codazzii is given preference, as the other two names 

were used for extreme forms. In the Darien area the 

range of variation is not as great as in Colombia and 

in the Gatun formation.

C. maldonadoi represents the subrounded, nonspinose 

form of C. codazzii. The spire whorls of C. werenfelsi 

Jung (1965, p. 552, pi. 75, figs. 9-11; middle Miocene, 

Venezuela) are more distinctly turreted.

A middle Miocene Venezuelan species, C. venezuelana H. K. Hodson (Hodson and Hodson, 1931, p. 45, 

pi. 23, figs. 1, 4), is more inflated than C. codazzii and 

has coarser spiral sculpture. C. montserratensis Maury 

(Kutsch, 1942, p. 163, pi. 9, figs. 7a, 7b), a late Miocene 

Trinidad species, also has coarser spiral sculpture and 

includes distinctly tabulated shells. In general features 

C. codazzii is similar to the type of Euclid, the living 

Panamic C. cassidiformis Sowerby (Keen, 1958, p. 

439, fig. 693), which is larger, invariably spinose, and 

its spire whorls are angulated, like the body whorl. 

The fossil species, however, is much more similar to 

another species living in Panama Bay: C. balboce Pilsbry (1931, p. 439, pi. 41, figs. 7, 8). The type of C. ~balboce is lost or misplaced and the paratype is immature. 

Twenty-two specimens were collected recently from 

dredge dumpings at Fort Amador, Canal Zone, by R. 

H. Stewart, Joanne L. Alien, and Douglas Alien. 

According to this sample, C. balboce is similar to the 

slightly or moderately angulated form of C. codazzii, 

but has fewer axial ribs on spire whorls. The type of 

C. balbocB (height 45 mm) evidently is exceptionally 

large.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 134, 

136, 136a, 137, 137a, 138a, 138b, 138c, 138d, 138e, 138f. 

Middle part, eastern area, localities 139c, 139e, 139f, 

155. Middle Miocene, Colombia, Darien area (USGS 

8429, 8430, 8453, 8476, 8477), Panama. Angostura formation (middle Miocene), Ecuador.

Cancellaria (Euclia) dinota Woodring1, n. sp. 

Plate 54, figures 1, 2; plate 56, figures 5, 6

Of medium size, moderately inflated. Body whorl 

rounded at shoulder. Protoconch naticoid, 2i/£>-whorled. 

End of protoconch marked by appearance of sculpture. 

First sculptured whorl or two angulated or subangulated at shoulder; remaining whorls rounded. First 

sculptured whorl bearing 10 to 12 axial ribs and three 

or four spiral threads. Axial ribs narrow, except on 

body whorl; more widely spaced on entire body whorl, 

or crowded and somewhat subdued on last third of 

whorl; 16 to 21 (generally 18 or 19) ribs on penult, 

13 to 18 (generally 17) on body whorl of specimens

that have ribs of essentially uniform spacing. Spiral 

threads narrow; low and subdued on late whorls; six to 

eight (generally seven) on penult. Spiral thread at 

shoulder and those between shoulder and suture rarely 

a little stronger than others. Secondary spirals appearing on penult; those 011 main part of body whorl almost 

as strong as primaries. One or two (generally two) 

widened ribs on body whorl and preceding two whorls.

Height (practically complete) 29.9 mm, diameter 18 

mm (type). Height 34.9 mm, diameter 21 mm (exceptionally large figured specimen).

Type: USNM 645728.

Type locality: 138c (USGS 21956, about 100 meters 

north of Transisthmian Highway and about 75 meters 

west of road to refinery site on Payardi Island, 

Panama; immediately east of Cativa and 100 meters 

north of locality 138), lower part of Gatun formation.

In marked contrast to the diversity shown by Cancellaria codazzii, G. dinota is fairly uniform. The spacing of axial ribs on the last third of the body whorl, 

however, is not uniform. Also on a few specimens the 

spiral thread at the shoulder and those between the 

shoulder and the suture are slightly emphasized. The 

size of the exceptionally large specimen shown on plate 

56, figures 5, 6, is the most conspicuous exception to 

uniformity. Many of the shells, including the type, 

are somewhat worn.

C. dinota is most similar to C. harpiformis Pilsbry 

and Olsson (1941, p. 23, pi. 3, figs. 1, 2), a Pliocene 

species from Ecuador. That species is larger (height 

43 mm), its spire is lower, and its sculpture is coarser.

C. dinota occurs in association with C. codazzii in the 

lower part of the Gatun formation, but has not been 

found in the middle or upper parts. It is abundant only 

at the type locality, where 61 specimens were collected. 

Immature shells may be distinguished from immature 

shells of C. codazzii by the finer sculpture of C. dinota.

Occurrence: Lower part of Gatun formation (middle 

Miocene), localities 136, 136a, 137a, 138, 138a, 138c, 

138d, 138f.

Submenus?

The following species is distinguished by its small 

size, slightly inflated spire whorls, narrowly reticulate 

sculpture, strong columellar folds, widely swinging 

siphonal fascicle and accompanying relatively wide 

umbilicus. The general facies of form and sculpture and 

the strong columellar folds suggest Cancellaria s.s., 

whereas the widely swinging siphonal fasciole suggests 

Bivetopsia Jousseaume (1887, p. 193; type (logotype, 

Cossmann, 1899, p. 9) : Cancellaria chrysostoma Sowerby, living, Panama to Peru) and Bivetiella Wenz 

(1943, p. 1,356; type (orthotype) : Cancellaria similis 

Sowerby, living, west Africa). Contrary to the view
GASTROPODS: EITLIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 341

expressed in 1928 (Woodring, 1928, p. 221), when 

Bivetopsis was inadvertently used for Bivetopsia, Bivetopsia and Bivetietta are scarcely distinguishable, except on a basis of sculptural plan. Bivetiella Marks 

(1949, p. 456) is not only a junior homonym of Bivetiella Wenz, it is also an objective junior synonym of 

that name.

Cancellaria acalypta Woodring, n. sp. 

Plate 53, figures 1, 2

Small, moderately inflated, spire whorls slightly and 

uniformly inflated. Protoconch 21/^-whorled, end 

marked by abrupt appearance of sculpture. Sculpture 

narrowly reticulate. Axial ribs wider than spiral 

threads and for most part a little more widely spaced. 

First sculptured whorl bearing nine axial ribs and 

four spiral threads. Body whorl sculptured with 20 

to 22 axial ribs and 13 or 14 spiral threads. Third 

sculptured whorl and later whorls bearing one or two 

slightly widened ribs. Aperture ovate, tapering to a 

very short siphonal canal. Columella bearing three 

folds, progressively stronger from first (basal) to 

third. Columellar folds, lirse on interior of outer lip, 

and ridges at upper end of parietal wall strong for 

size of shell. An elongate denticle alined with outer 

end of third columellar fold; a denticle near edge of 

columellar lip between second and third folds; and an 

elongate denticle extending obliquely from outer end of 

second fold. Siphonal fasciole moderately inflated, 

swinging far from outer edge of columellar lip, forming a relatively wide umbilicus.

Height: 15.6 mm, diameter 10.2 mm (type).

Type: USNM 645730; paratype, Stanford University.

Type locality 136a (Stanford University 2611, Transisthmian Highway, lat 9° 21' N. plus 1,100 feet (335 

meters), long 79° 49' W., Panama; same as USGS 

16912), lower part of Gatun formation.

Three specimens of this small cancellarid were 

found in the lower part of the Gatun formation and 

one in deposits near the base of the middle part. All 

are of approximately the same size. Despite the small 

size, the apertural features are strong, especially on 

the shell from the middle part of the formation. The 

body-whorl sculpture of that specimen is also for the 

most part more evenly reticulate than that of the 

others. The basal part of the outer lip of the type is 

missing and the lip of the other specimens is damaged.

The axial ribs of Cancellaria laqua Mansfield (1935, 

p. 26, pi. 2, fig. 5; middle Miocene, Florida), O. lavelana H. K. Hodson (in Hodson and Hodson, 1931, 

p. 44, pi. 24, fig. 12; middle Miocene, Venezuela), and 

an unnamed late Miocene Mexican species (USGS

18687) are closely spaced. The third columellar fold 

of these three species is wider than that of C. acalypta. 

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 136, 

136a. Middle part, eastern area, locality 139c.

Submenus?

Cancellaria nancellaria Woodring, n. sp. 

Plate 53, figures 3, 4

Small, moderately slender, whorls rounded. Protoconch large for size of shell, naticoid, 3-whorled, end 

marked by appearance of first axial rib. First sculptured whorl bearing eight to ten axial ribs and four 

or five spiral threads. Sculpture throughout consisting 

of narrow axial ribs (nine or ten on penult, 11 to 13 

on body whorl) and narrower spiral threads (five or 

six on penult, 11 to 13 on body whorl). Aperture of 

moderate width, tapering to a short siphonal canal. 

Columella bearing three fairly strong columellar folds. 

Denticle near outer edge of columellar lip between 

second and third (from base) folds present or absent. 

Interior of outer lip bearing moderately strong lirae. 

Siphonal fasciole slightly inflated, swinging some distance from outer edge of columellar lip, forming a 

narrow umbilicus.

Height 7.6 mm, diameter 4.5 mm (type). Height 

8.9 mm, diameter 5 mm (largest specimen).

Type: USNM 645731.

Type locality: 147b (USGS 6033c, Panama Kailroad, about 3,500 feet (1,065 meters) southeast of 

Gatun railroad station, Canal Zone), middle part of 

Gatun formation.

Twenty-eight specimens, collected at five localities 

(four in the middle part of the Gatun formation and 

one in the upper part in the eastern area) are referred 

to Cancellaria nancellaria. As usual, the lower part of 

the outer lip of the type and other specimens, except 

some of the smallest, is imperfect. The largest of these 

fossils has a height under 9 mm and almost half of 

them consist of only the protoconch and one or two 

later whorls. The type locality, as already mentioned 

(p. 69, 74, 75, 88,103, 274), is noteworthy for its many 

small specimens, both mature small species and immature shells. Small specimens are fairly common also 

at two of the other localities at which C. nancellaria 

was found (146 and 147g), both near the type locality. 

Nevertheless these three collections include moderately 

large fossils. Despite their small size, perhaps the 

largest shells of this species are mature. If they are 

mature, C. nancellaria is a minute species. The columellar folds, the short siphonal canal, and the position of the siphonal fasciole suggest Cancellaria s.s.,
342 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

but the outline and the widely spaced axial ribs are 

more similar to those of Narona, the subgenus under 

the next heading. At all events no close allies of C. 

nancellaria are recognized.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

localities 146, 147b, 147g, 153a. Upper part, eastern 

area, locality 175.

Subgenus Narona H. and A. Adams

H. and A. Adams, The genera of Recent Mollusca, v. 1, p. 277,

1854. 

Type (logotype, Jousseaume, Le Naturaliste, 2d ser., annee 9,

p. 222, 1887): Cancellaria clavatula Sowerby, living,

Mexico to Peru.

Narona, like Pyruclia and Euclia, formerly lived in 

the present Caribbean region, but survives in the 

eastern Pacific Ocean. Despite its very short siphonal 

canal, Cancellaria bullbrooki Mansfield (1925, p. 31, 

pi. 5, fig. 3), a middle Miocene species from Trinidad, 

evidently is a species of Narona that has strongly 

constricted whorls. The type doubtless is immature. 

Unlike Pyruclia and Euclia, Narona reached Florida. 

As recognized by both Gardner and Mansfield, it is 

represented there by middle and late Miocene species: 

Oancellaria atraktoides Gardner (1926-47, p. 376, pi. 

45, figs. 13, 14, 1938; middle Miocene), G. blountiana 

Mansfield (1935, p. 27, pi. 2, fig. 4; middle Miocene), 

G. agria uaquala Mansfield (1935, p. 27, pi. 2, figs. 7, 

8; middle Miocene), C. agria agria Mansfield (1930, p. 

48, pi. 3, fig. 1; late Miocene), and G. coensis Mansfield (1930, p. 49, pi. 3, figs. 3, 4, late Miocene).

Cancellaria (Narona) barystoma Woodring, n. sp. 

Plate 53, figures 5, 6

Small, slender, distinctly turreted. First two sculptured whorls weakly angulated at shoulder, later 

whorls rounded. Protoconch small, naticoid, 2*4- 

whorled, end marked by appearance of sculpture. First 

sculptured whorl bearing nine narrow axial ribs and 

two very narrow spiral threads. Axial ribs of late 

whorls swollen, 13 on penult and 12 on body whorl. 

Spiral threads of late whorls narrow, 10 on penult. 

Secondary spiral threads on main part of body whorl. 

Two ribs on penult and three on body whorl varicose. 

Aperture narrow, tapering to a short siphonal canal. 

Lirse on interior of outer lip strong. Columella bearing two strong folds. Inner edge of columellar lip 

slightly inflated below lower fold and bearing a denticle between the folds. Siphonal fascicle slightly inflated, close to outer edge of columellar lip.

Height 16.3 mm, diameter 7.8 mm (type).

Type: USNM 645732.

Type locality: 138d (USGS 22016, about 100 meters 

north of Transisthmian Highway and about 75 meters 

west of road to refinery site on Payardi Island, Panama ; immediately east of Cativa and 100 meters north 

of locality 138), lower part of Gatun formation.

In essential features Gancellaria ~barystoma is similar 

to the type species of Narona. The fossil species, however, is distinguished by its smaller size, more rounded 

late whorls, more numerous axial ribs and spiral 

threads, and above all by the much stronger columellar folds and lirse on the interior of the outer lip. 

No other species of Narona is known to have such 

strong apertural features, but those features may not 

be fully formed on some of the fossil species.

Though G. ~barystoma is represented by 13 specimens (12 from the lower part of the Gatun formation 

and one from a horizon near the base of the middle 

part), the type is the only mature shell. The next 

largest has a height of 10 mm. As is to be expected, 

its apertural folds and lirse are not as strong as those 

of the type. Two of the immature shells are not as 

slender as the others. The edge of the outer lip of the 

type is missing except at and near its insertion.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 137, 

138, 138a, 138b, 138c, 138d. Middle part, eastern area, 

locality 139c.

Cancellaria (Narona) decaptyx Brown and Pilsbry

Cancellaria decaptyx Brown and Pilsbry, Acad. Nat. Sci. Phila. 

Proc., v. 63, p. 346, pi. 24, figs. 5, 6, 1911 (Miocene, Canal 

Zone).

This species, collected at the Gatun Locks excavation, is not recognized in the collections at hand. The 

type is the only available specimen.

Like Cancellaria barystoma, C. decaptyx is small and 

slender (height 11 mm, diameter 5 mm). It is distinguished from G. barystoma by its more slender outline and sculpture. The first sculptured whorl bears 

three spiral threads, and on the penult and body whorls 

the threads are more numerous and more closely spaced 

than those of G. barystoma (11 on penult). The columellar folds and lirse on the interior of the outer lip 

are of about the same strength as those on a specimen 

of G. barystoma of comparable size.

Type: Acad. Nat. Sci. Phila. 1701.

Type locality: Gatun Locks excavation, Canal Zone, 

middle part of Gatun formation.

Occurrence: Middle part of Gatun formation (middle Miocene), Gatun Locks excavation.
GASTROPODS: EULIMIDAE, MARGESTELLIDAE TO HELMINTHOGLYPTIDAE 343

Subgenus Charcolleria Olsson

Olsson, Bull. Am. Paleontology, v. 27, no. 106, p. 61, 1942. 

Type (orthotype): Cancellaria (Charcolleria) perdiciana Olsson, 

Miocene, Colombia.

The large, fusiform, reticulately sculptured shells of 

Charcolleria are so different from those of Cancellaria 

s.s. that at first glance generic rank seems to be amply 

justified. Nevertheless Charcolleria evidently is allied 

to Massyla (H. and A. Adams, 1854, p. 278; type 

(monotype): Cancellaria corrugata Hinds, living, 

Ecuador to Peru) and Massyla is generally treated as 

a subgenus of Cancellaria. The type of Massyla is a 

rare species and no specimens are available. According to Hinds' (1844-45, p. 42, pi. 12, figs. 1, 2, 1844) 

illustrations, it is fairly slender and, like Charcolleria, 

has two columellar folds and narrowly reticulate 

sculpture. It is, however, less than half as large 

(height about 18 mm), as Charcolleria, less slender, 

and has weaker spiral sculpture. Usage of Massyla for 

European Miocene species (Sacco, 1894, p. 67, pi. 3, 

figs. 80a, 80b; Cossmann, 1899, p. 39-11, pi. 2, figs. 13, 

14) is unjustified.

Oharcolleria has been found so far in the lower 

Miocene of Colombia, the middle Miocene of Colombia and the Caribbean part of Panama, the middle 

and upper Miocene of Venezuela, the upper Miocene 

or Pliocene of Ecuador, and the Pliocene of southwestern Costa Rica near the boundary of Panama. It has 

no known predecessors or descendants in the Caribbean region or elsewhere.

It is surprising that in many features the two known 

species of Charcolleria are similar to a Pliocene species 

of southeastern United States: Cancellaria venusta 

Tuomey and Holmes (Olsson and Harbison, 1953, p. 

179, pi. 28, figs. 8, 8a, 8b), which was assigned to 

Massyla by Olsson and Harbison. Cancellaria propevenusta Mansfield (1930, p. 47, pi. 17, fig. 2; upper 

Miocene, Florida) may be a predecessor of Cancellaria 

venusta and perhaps Cancellaria propevenusta arose 

from Cancellaria runchcena Gardner (1926-47, p. 375, 

pi. 45, figs. 8, 9, 1938; middle Miocene, Florida). The 

resemblance of Charcolleriarto Cancellaria venusta is 

attributed to convergence.

Cancellaria (Charcolleria) terryi Olsson 

Plate 54, figures 5, 6, 9, 10

Cancellaria (Charcolleria) terryi Olsson, Bull. Am. Paleontology, v. 27, no. 106, p. 62, pi. 8, fig. 1, 1942 (Pliocene, 

Costa Rica). Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst, p. 124, pi. 

22, fig. 2, 1964 (Miocene, Ecuador). Jung, Bull. Am. 

Paleontology, v. 49, no. 223, p. 556, pi. 75, figs. 17-19, 

1965 (Miocene, Venezuela).

Cancellaria (Charcolleria) sp., Olsson, Neogene mollusks from 

northwestern Ecuador, Paleontological Research Inst., 

p. 124, pi. 22, figs. 1, la, 1964 (Miocene, Canal Zone).

Large, slender, fusiform, whorls strongly inflated. 

Protoconch and earliest sculptured whorl missing or 

worn. Sculpture of remaining whorls strongly and 

evenly reticulate, forming squares, or more generally 

rectangles that are slightly elongate spirally. Penult 

whorl bearing 27 to 37 axial ribs and nine or ten spiral 

cords. Axial ribs slightly narrower and more closely 

spaced on irregularly spaced varixlike slight swellings 

on late whorls of large shells. Microscopic axial threads 

visible on unworn late whorls. Aperture elongate, tapering to a short siphonal canal. Columella bearing 

two moderately strong folds. Siphonal fascicle slightly 

inflated, swinging some distance from edge of columellar lip.

Height (incomplete) 53.5 mm, diameter 24 mm 

(larger figured specimen). Height 42.3 mm, diameter 

19 mm (smaller figured specimen).

Type: Paleontological Research Inst. 4045.

Type locality: Quebrada Penites, Puntarenas Province, Costa Rica, Charco Azul formation.

Collections from the lower and middle parts of the 

Gatun formation include 16 specimens of this large 

fusiform cancellarid. Eight of them were collected at 

locality 138c and one or two at each of the other seven 

localities. The sculpture of all these specimens is 

strongly reticulate and essentially uniform. It is 

coarser than the sculpture of the type of Charcolleria: 

Cancellaria perdiciana (Olsson, 1942, p. 61, pi. 8, fig. 

5), an early Miocene moderately deep-water species 

from Colombia. The coarse sculpture is the basis for 

the identification of the Gatun species as Cancellaria 

terryi. The identification means that Cancellaria terryi 

survived later in eastern Pacific waters than in western 

Atlantic waters.

Cancellaria terryi is twice as large as Cancellaria 

venusta (height 22 to 26 mm) the Pliocene species 

already mentioned and has more inflated spire whorls 

and stronger axial sculpture. Cancellaria propevenusta 

also already mentioned is low spired and inflated 

in outline; otherwise it is similar to Cancellaria 

venusta, aside from the difference in size. The type of 

Cancellaria propevenusta has a height of 35.8 mm, but 

specimens in Druid Wilson's upper Miocene collections 

from peninsular Florida reach a height of 46 mm. 

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138c, 

138d. Middle part, eastern area, localities 139f, 140, 

155,155a, 155b, 157. Cantaure formation (middle Miocene), Venezuela. Punta Gavilan formation (late Miocene), Venezuela. Esmeraldas formation (late Mio8-278 O - 70 - 4
344 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

cene), Ecuador. Charco Azul formation (Pliocene), 

Costa Rica.

Genus Aphera H. and A. Adams

H. and A. Adams, The genera of Kecent Mollusca, v. 1, p. 277,

1854. 

Type (monotype): Cancellaria tessellata Sowerby, living, Gulf

of California to Peru.

Aphera tessellata, a remarkable cancellarid, is the 

sole surviving species of Aphera. During Miocene 

time the genus had a more extensive distribution. It 

occurs in the lower Miocene of southeastern Costa 

Rica; the middle Miocene of the Dominican Republic, 

the Caribbean part of Panama, southeastern Costa 

Rica, Mexico, Florida, and Peru; and the upper Miocene of Peru. In other words, Aphera (as well as 

Pyruclia, Euclia, and Narona] is one of the many 

genera or subgenera that formerly lived in the Caribbean region, but now survive in the eastern Pacific 

Panamic region.

Miocene European species that have been referred 

to Aphera (Sacco, 1894, p. 66-67, pi. 3, figs. 76-79; 

Cossmann, 1899, p. 18, pi. 1, fig. 11) lack the distinctive apertural features of that genus.

Aphera islacolonis (Maury) 

Plate 56, figures 1, 2

Cancellaria tessellata Sowerby, Gabb, Am. Philos. Soc. Trans., 

n. ser., v. 15, p. 236, 1873 (Miocene, Dominican Republic).

Cancellaria (Aphera) islacolonis Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 65, pi. 10, figs. 12, 12a, 12b, 1917 

Miocene, Dominican Republic).

Cancellaria islacolonis Maury, Olsson, Idem, v. 9, no. 39, p. 86, 

pi. 6, fig. 12, 1922 (Miocene, Costa Rica) ; assigned to 

subgenus Aphera.

Cancellaria ellipsis Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 73, 

p. 333, pi. 22, figs. 8, 9, 1922 (Miocene, Dominican 

Republic).

Small, elliptical, low-spired. Protoconch naticoid, 

21/£-whorled, end marked by gradual appearance of 

sculpture. Sculpture reticulate. Spiral cords strongly 

overriding axial ribs, wider than the ribs on spire 

whorls, of about same width on body whorl. Penult 

whorl sculptured with 30 to 35 axial ribs and five or 

six spiral cords. On spire whorls width and spacing of 

axial ribs irregular at irregular intervals corresponding 

to varices on other cancellarids; on body whorl axial 

ribs closely spaced and subdued near outer lip. Aperture elongate, narrow. Outer lip somewhat thickened 

and slightly ascending; liree on its interior strongly 

thickened just within aperture. Columella bearing two 

strong folds. Apertural shield thick; traces of underlying sculpture reflected on parietal part; low, irregular, elongate denticles on part opposite end of columellar folds. Upper part of parietal wall bearing an 

elongate denticle.

Height: 16.8 mm, diameter 10 mm (figured specimen).

Type: Cornell Univ.

Type locality: Bluff 3, Cercado de Mao (long bluff 

on left bank of Rio Mao opposite Hata Viejo, about 

5 km above ford at Cercado de Mao), Dominican Republic, Cercado formation.

Aphera islacolonis is a rare species in the lower part 

of the Gatun formation and still rarer in the middle 

part. The figured specimen is the largest of the five 

that have been found. The protoconch of a shell in a 

Stanford University collection (locality 138a) is 

slightly tilted, like that of some specimens of A. tessellata. The single shell from the middle part of the 

Gatun is incomplete and partly exfoliated. The reflection of axial ribs on the parietal part of its apertural shield is relatively strong.

As compared with topotypes and other specimens 

from the Dominican Republic, the axial ribs of these 

few Gatun fossils are wider and fewer (some 30 as 

compared with some 40 on the penult whorl), their 

outer lip is thicker, and their lirse on the interior of 

the outer lip are thicker just within the aperture. 

Perhaps these coarser features indicate that the Panama fossils lived in shallower water than those in the 

Dominican Republic and they may eventually be 

given subspecific rank. Olsson's figured specimen and 

the type of Pilsbry's Cancellaria ellipsis are not entirely mature. In Mexico (USGS 9995, near Santa 

Rosa, Vera Cruz) A. islacolonis reaches a slightly 

larger size than elsewhere (height 18.5 mm).

A. islacolonis is more similar to A. tessellata than 

to the other described species of Aphera. That living 

species is larger (height 26.5 mm), more slender, and 

has a higher spire. It is doubtful whether A. wigginsi 

Emerson and Hertlein (1964, p. 362, figs. 5d, e; Pleistocene, Isla Montserrate, Gulf of California) is to be 

distinguished from A. tessellata.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138a, 

138c, 138f. Middle part, western area, locality 161c. 

Uscari shale (early Miocene part), Costa Rica. Middle 

Miocene deposits, Costa Rica, Mexico. Cercado formation (middle Miocene), Dominican Republic.

Genus Trigonostoma Blainville

Blainville, Manuel de malacologie et de conchyliologie, p. 652, 

1827.

Type (monotype and tautotype): Delphinula trigonostoma Lamarck (= Triaona pellucida Perry), living, tropical western Pacific Ocean.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 345

Petit (1967, p. 217) has pointed out that Lamarck's 

well known name for the type species is a junior 

synonym of Trigona pettucida.

Trigonostoma cf. T. scalatella (Guppy)

Small, slender, scalariform, sutural area sloping 

toward shoulder. Protoconch and spire whorls up to 

later part of penult missing. Body whorl sculptured 

with eight strongly arched axial ribs and 18 spiral 

threads, swollen on crest of ribs. Spiral threads fading 

out in sutural area. Near base a faint secondary thread 

appearing in two of spaces between spiral threads. 

Aperture and umbilicus narrow. Lirse on interior of 

outer lip strong just within aperture. Columella bearing two folds and a faint basal swelling.

Estimated height 12 mm, diameter 6.8 mm.

Two fragmentary shells demonstrate the presence 

of a small, slender species of Trigonostoma in the 

Gatun formation. The description is based on the 

better of the two (locality 139c). The other fragment 

presumably represents the same species.

In general outline this unnamed, apparently new, 

species is similar to the somewhat larger Miocene Jamaican Trigonostoma scalatella (Guppy) (Woodring, 

1928, p. 224, pi. 13, fig. 1). The sutural area of that 

species is horizontal, its spiral sculpture consists of 

closely spaced primary and secondary threads, and its 

aperture is wider. T. funiculatum (Hinds) (1844-45, 

p. 43, pi. 12, figs. 5, 6, 1844; living, Baja California, 

Gulf of California) is more inflated, and has secondary spiral threads and a wider aperture.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, locality 138e. 

Middle part, eastern area, locality 139c.

Trigonostoma cf. T. insulare (Pilsbry and Johnson)

Cancellaria (Trigonostoma) aff. Cancellaria ~bullata Sowerby, 

Toula, K. k. Geol. Reichsanstalt Jahrb, v. 61, p. 504, 

pi. 30, fig. 10, 1911 (Miocene, Canal Zone).

Toula's specimen of this Gatun species is missing in 

his collection and no other is available. According to 

his description and illustration, the sculpture of the 

penult whorl is finely reticulate. The body whorl is 

sculptured with 14 narrow, widely spaced axial ribs 

and fairly strong spiral threads. The columella bears 

three folds.

Pilsbry and Johnson thought this species may be 

their Trigonostoma insulare (Pilsbry and Johnson, 

1917, p. 163; Pilsbry, 1922, p. 334, pi. 22, fig. 11) and 

Olsson thought it probably is his T. toroense (Olsson, 

1922, p. 84, pi. 6, fig. 4). The early finely reticulate 

sculpture of the Gatun species suggests that of T.

toroense, but the ribbing on the body whorl is more 

like that of T. insulare.

T. insulare is found in the Dominican Republic, 

presumably in the Cercado or Gurabo formation, and 

T. toroense in the late Miocene Limon formation of 

the Bocas del Toro area of northwestern Panama.

Occurrence: Middle part of Gatun formation (middle Miocene), Gatun Locks excavation.

Trigonostoma n. sp. 

Plate 63, figures 14, 15

Of medium size, moderately inflated. Sutural area 

depressed, forming a deep U-shaped gutter. Earliest 

preserved sculpture consisting of weak axial ribs and 

stronger spiral threads. Penult and body whorls sculptured with low, narrow-crested, widely spaced axial 

ribs (14 on body whorl) and subdued, closely spaced 

primary and secondary spiral threads. Outer lip broken 

back; interior of preserved part smooth. Umbilicus 

moderately wide. Columella bearing two slender folds.

Height (almost complete) 20.2 mm, diameter 

14.7 mm.

The Chagres sandstone yielded a poorly preserved 

specimen of this species. Both the outer and columellar 

lips are broken back and much of the outer shell material is corroded. The deep U-shaped sutural gutter; 

the distinct, though low and narrow-crested, axial 

ribs; and the distinct, though subdued, spiral threads 

indicate it is a new species. The axial ribs of Trigonostoma woodringi Jung (1965, p. 557, pi. 76, figs. 1, 2; 

middle Miocene, Venezuela) are wider.

Occurrence: Chagres sandstone (late Miocene), locality 208.

Family CONIDAE

Genus Conns Linne

Linne, Systema naturae, 10th ed., p. 712, 1758.

Type (logotype, Children, Quart. Jour. Sci. Lit. Art, v. 16, p. 

69, 1823): Conus marmoreus Linne, living, western Pacific Ocean.

Though many attempts have been made to recognize 

subgenera, or genera, in the traditional genus Conus, 

they have not been notably successful. The species 

living in both the western Atlantic and eastern Pacific 

Oceans have been reviewed recently (Clench, 1942; 

Hanna, 1963, respectively).

The genus has been found in the Gatuncillo, Bohio, 

Caimito, Culebra, La Boca, and Gatun formations, and 

in the Chagres sandstone, but the only adequate representation is in the Gatun. With the exception of two 

specimens of medium size, the formations older than 

the Gatun yielded only small specimens and the species 

themselves evidently are small. The exceptions are an
346 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

unidentified specimen from the Culebra formation (locality 110) and another from upper member of the 

Alhajuela formation (locality 89). In addition to the 

briefly described Caimito, Culebra, and La Boca species, the collections from those formations include a 

considerable number of unidentified specimens, most 

of which are, in fact, unidentifiable.

The Gatun formation contains 16 species and subspecies of Conus: eight in the lower part, 14 in the 

middle part, seven in the upper part in the eastern 

area, and six in the upper part in the western area. 

Eight, the largest number in any collection, were recovered at localities 138c and 155. The large number in 

the middle part is not unusual in the middle Miocene 

of the Caribbean region. Twelve occur in southeastern 

Costa Rica, 16 in the Bowden formation of Jamaica, 

and 16 in the Gurabo formation of the Dominican Republic. Four of the Gatun species are closely related to 

living species and one is identified as a living species. 

As shown in the following table, the four closely allied 

species are living in the eastern Pacific Panamic region, 

whereas the species still living is a Caribbean species.

Species of Conus from Gatun formation and closely allied or 

identical species living in Panamic and Caribbean regions

Species in Gatun formation

C. recognitus Guppy ___. 

C. spurius Gmelin _ ___

C. molis Brown and 

Pilsbry. 

C. consobrinus SowerbyC. imitator Brown and 

Pilsbry.

Closely allied or identical living species

Panamic region

C. patricius Hinds __

C. fergusoni 

Sowerby. 

C. emersoni Hanna 

C. arcuatus Broderip 

and Sowerby.

Caribbean region

C. spurius 

Gmelin.

Despite the large number of species in the Gatun 

formation, Conus planiliratus so far has not been found 

in the Canal Zone. That species is widespread in middle 

Miocene formations in the Caribbean region, is abundant in the late Miocene L/imon formation of southeastern Costa Rica, occurs in the Pliocene Moin formation of that area, and may still be living in western 

Atlantic waters. It has been pointed out that the living 

Caribbean species C. stimpsoni is similar to C. planiliratus (Woodring, 1928, p. 211). The recently described C. austini (Render and Abbott, 1951, p. 22, fig. 

7) is even more similar. In fact, it is doubtful whether 

C. austini can be distinguished consistently from C. 

planiliurats by any feature other than the slightly 

larger size of the Recent form.

The status of C. tortuosopunctatus, an alleged Gatun 

species, is discussed on page 358.

Conus recognitus Guppy 

Plate 55, figure 4

Conus solidus Sowerby, Geol. Soc. London Quart. Jour., v. 6, 

p. 45, 1850 (Miocene, Dominican Republic). Guppy, Idem, 

v. 22, p. 287, pi. 16, fig. 1, 1866 (Miocene, Jamaica). Not 

Conus solidus Gmelin, 1791, or Conus solidus Sowerby, 

1834(?).

Conus recognitus Guppy, Sci. Assoc. Trinidad Proc., pt 3, p. 

171, 1867 (Miocene, Dominican Republic, Jamaica) ; reprint, Bull. Am. Paleontology, v. 8, no. 35, p. 50, 1921. 

Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 45, pi. 7, 

fig. 9, 1917 (Miocene, Dominican Republic). Pilsbry, 

Acad. Nat. Sci. Phila. Proc., v. 73, p. 327, pi. 19, fig. 2, 

1922 (Miocene, Dominican Republic). Olsson, Bull. Am. 

Paleontology, v. 9, no. 39, p. 46, pi. 2, fig. 9, 1922 (Miocene, Costa Rica). Maury, Brazil Servigo, Geol. y 

Mineral. Mon. 4, pi. 11, fig. 20, 1925 (Miocene, Dominican 

Republic). Anderson, Calif. Acad. Sci. Proc., 4th ser., 

v. 18, no. 4, p. 109, 1929 (Miocene, Colombia).

Conus (Lithoconus) recognitus Guppy, Pfiug, Acta Humboldtiana, Geol. Palseontologica Ser., no. 1, p. 59, pi. 18, figs. 

12-15, 1961 (Miocene, Dominican Republic).

Conus pyriformis Reeve, Gabb, Am. Philos. Soc. Trans., n. ser., 

v. 15, p. 229, 1873 (Miocene, Dominican Republic).

Of medium size, moderately slender, pyriform. Spire 

low, its profile only slightly concave, owing to absence 

of earliest whorls. Shoulder rounded, but marked by 

obscure spiral thread. Earliest whorls missing, earliest 

preserved whorls corroded. Anal fasciole slightly convex, bearing faint microscopic spiral lineation. Lower 

two-thirds of body whorl sculptured with low spiral 

threads, gradually replaced by faint lineation on upper 

third.

Height (almost complete) 39 mm, diameter (body 

whorl broken back) 20.8 mm (figured specimen).

Type material: Lectotype, British Museum (Natural 

History) G83971.

Type locality: Valley of Rio Yaque del Nbrte, 

Dominican Republic, Miocene.

A damaged small specimen, found in the upper part 

of the Gatun formation in the western area, is the only 

pyriform cone in the collections at hand. It would be 

more pyriform, if the outer lip were not so far broken 

back.

This species is identified as Conus recognitus, a name 

proposed, without any discussion of the features of the 

species, for Sowerby's junior homonym C. solidus. My 

discussion (Woodring, 1928, p. 203,205) of those names 

was based on a cast, kindly forwarded in 1926 by the 

late L. R. Cox, of the British Museum (Natural History), It bears the number 12802, the number for C. 

solidus in Sherborn's register of the Heneken collection 

of Miocene fossils from the Dominican Republic described by Sowerby. When I had an opportunity to 

examine the Heneken collection in 1958, two small
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 347

cones were under that number. The larger of the two 

(height 39.2 mm, diameter 23.2 mm) was used in making the cast, a duplicate of which was in the tray with 

the specimens. As was evident from the excellent cast, 

this small cone is not a pyriform species, but is a representative of the C. spurius group. In 1961, however, 

Pflug published under the same Sherborn register 

number illustrations of two pyriform cones in the 

Heneken collection, the larger of which (height 65 mm, 

diameter 34 mm) was designated the lectotype of C. 

recognitus/ that is, the lectotype of C. solidus, which 

automatically is the lectotype of C. recognitus. Pflug 

did not mention the two small specimens formerly 

under the number 12802. Though no pyriform cones 

were seen in the Heneken collection in 1958, Pflug's 

lectotype designation is accepted, principally because 

it agrees with Guppy's 1866 identification of a Miocene 

Jamaican pyriform cone as C. solidus. Guppy was the 

first paleontologist to deal with Sowerby's species and 

he had access to the Heneken collection when he was in 

London. As indicated by Sherborn's remarks in his 

register (translated on page 8 of Pflug's publication), 

much can happen to a collection that has passed 

through so many hands. Since 1867 the name C. recognitus has been used for a pyriform species, with the 

exception of my 1928 action.

In the collections from the Dominican Republic in 

the U.S. National Museum C. recognitus is represented 

by small specimens, comparable in size to the Gatun 

fossil, from the Baitoa and Cercado formations. The 

large lectotype was found presumably in the Gurabo 

formation. The occurrence in that formation was recorded by Maury.

A large broad-shouldered, low-spired, pyriform cone, 

collected by Gabb in the Dominican Republic, was 

named C. williamgabM (Maury, 1917, p. 36, pi. 5, 

fig. 2). It is known to occur in the Baitoa formation. 

The Bowden formation of Jamaica yielded a similar 

pyriform species, C. apium (Woodring, 1928, p. 202, 

pi. 9, fig. 3), characterized by a low thread at the edge 

of the shoulder.

C. recognitus has an age range of late early Miocene 

to middle Miocene. As was recognized long ago, it has 

no living allies in Caribbean waters, but is similar to 

the eastern Pacific species C. patricius Hinds (Hanna, 

1963, p. 49, pi. 8, pi. 10, figs. 4, 5), appropriately designated G. pyrifonnis by Reeve. Pilsbry pointed out that 

the tuberculate shoulder of C. recognitus disappears at 

an earlier stage than that of C. patricius. In fact, good 

preservation of the first few post-protoconch whorls is 

needed to show the early tuberculate shoulder of C. 

recognitus. The early tuberculate shoulder of C. patricius is well shown on Hinds' illustration of the type

of that species, an immature shell (Hinds, 1844-45, p. 

7, pi. 1, figs. 1, 2,1844). The largest recorded C. recognitus is half as large as the exceptionally large C. patricius recently illustrated by Hanna (1963, pi. 8; 

height 140 mm). C. patricius is the type of the recently 

proposed su'bgenus Pyruconus (Olsson, 1967, p. 21).

Occurrence: Upper part of Gatun formation, western area (middle Miocene), locality 182. Baitoa formation (late early Miocene), Dominican Republic. Cercado and Gurabo formation (middle Miocene), Dominican Republic. Bowden formation (middle Miocene), 

Jamaica. Middle Miocene deposits, southeastern Costa 

Rica. Tubara formation (middle Miocene), Columbia.

Conus musaensis Olsson 

Plate 57, figure 2

Conus mustensis Olsson, Bull. Am. Paleontology, v. 9, no. 39, 

p. 47, pi. 1, figs. 22, 24, 1922 (Miocene, Costa Rica).

Small, moderately wide at shoulder, shoulder 

rounded or subrounded at maturity. Spire of moderate 

height, its profile slightly concave. Protoconch conspicuous, slender, cylindrical, li/^-whorled. Early postprotoconch whorls sharply carinate at shoulder. Carina 

gradually suppressed with further growth. Anal fasciole flat. Lower half to two-thirds, or all (especially 

on immature shells), of body whorl sculptured with 

flat spiral bands, separated by generally narrower 

channels. Spiral bands of immature shells nonpustulose or weakly pustulose.

Height 20.6 mm, diameter 10 mm (figured specimen).

Type material: Lectotype (herewith designated), 

specimen represented by Olsson's fig. 24), Paleontological Research Inst. 20887.

Type locality: Rio Banana, Limon Prov., Costa Rica, 

middle Miocene.

This small species occurs in the middle part of the 

Gatun formation west of Gatun Lake, where 19 specimens were collected. It is the first occurrence to be recorded outside the type area in southeastern Costa 

Rica. Immature shells are notably different from mature shells, owing to the carinate shoulder at an early 

stage.

The small size and general appearance suggest C. 

jaspideus Gmelin, of the present Caribbean fauna, especially the round-shouldered form, for which Reeve's 

name C. pygmceus is used, either as a subspecies of C. 

jaspideus (Clench, 1942, p. 12, pi. 7, figs. 1, 2), or as 

an infrasubspecific form of that species (Abbott, 1958, 

p. 89, pi. 3, fig. j). That the resemblance is superficial 

is indicated by the noncylindrical, more inflated protoconch and somewhat concave anal fasciole of C. jaspideus.
348 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Occurrence: Middle part of Gatun formation, western area (middle Miocene), localities 161c, 161d, 170. 

Middle Miocene deposits, southeastern Costa Eica.

Conus spurius Gmelin 

Plate 55, figure 7

Conus spurius Gmelin, Systema naturae, 13th ed., p. 3396, 1791 

(living, locality unknown). Clench, Johnsonia, v. 1, no. 6, 

p. 19, pi. 10, figs. 4, 5,1942 (living, Bahamas, West Indies).

Conns proteus Hwass, variety B, in Bruguiere, Encyclopedic 

methodique, Histoire naturelle des vers, v. 1, p. 682, pi. 

334, fig. 2, 1792 (living, West Indies). Pilsbry and Brown, 

Acad. Nat. Sci. Phila. Proc., v. 69, p. 32 (list), 1917 

(Miocene, Colombia). Maury, Bull. Am. Paleontology, 

v. 5, no. 29, p. 42, pi. 6, fig. 11, 1917 (Miocene, Dominican 

Republic). Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 73, 

p. 331, 1922 (Miocene, Dominican Republic). Olsson, 

Bull. Am. Paleontology, v. 9, no. 39, p. 43, pi. 2, figs. 3, 4, 

1922 (Miocene, Costa Rica, Panama).

Conus (Lithoconus) proteus Hwass, Woodring, Carnegie Inst. 

Washington Pub. 385, p. 204, pi. 9, fig. 4, 1928 (Miocene, 

Jamaica).

Conus leoninus Hwass, Gabb, Acad. Nat. Sci. Phila. Jour., 2d 

ser., v. 8, p. 359, 1881 (Pliocene, Costa Rica).

Of medium size and moderate width at shoulder, 

shoulder abruptly rounded. Spire low, practically flattopped, except part formed by earliest whorls. Protoconch missing, earliest preserved whorls worn. Anal 

fasciole flat, bearing weak spiral threads on early unworn whorls. Color pattern consisting of well separated 

spiral rows of brownish rectangles, or crude rectangles, 

of uniform width in a row, but of varying width from 

row to row.

Height 45.8 mm, diameter 28.5 mm (figured specimen).

Type locality: Presumably West Indies, living.

Conus spurius, formerly designated G. proteus, is 

represented by one specimen collected at the Gatun 

Third Locks site. The anal fasciole, inside the slightly 

bulging shoulder, is not concave. Though it generally 

is concave on both living and fossil specimens, that 

feature is variable. The color pattern, though bleached, 

is preserved, as it generally is on fossil specimens of 

this species, which ranges back to at least middle Miocene. The color pattern of the Gatun fossil is like that 

of G. spurius atlanticus Clench (1942, p. 20, pi. 10, 

figs. 1-3), which was given a range extending from the 

west coast of Florida to Venezuela, along the continental border. The same pattern is shown by Miocene 

Jamaican shells, but is less distinct on the Jamaican 

specimen illustrated in 1928. Whatever may be decided 

about living shells, it is improbable that the color pattern can be used to distinguish geographic races in G. 

spurius of Miocene age.

Occurrence: Middle part of Gatun formation, eastern area (middle Miocene), locality 155. Middle Miocene deposits, southeastern Costa Eica, Colombia. Bowden formation (middle Miocene), Jamaica. Cercado 

and Gurabo formations (middle Miocene), Dominican 

Eepublic. Deposits of late Miocene and Pliocene age, 

Bocas del Toro area, Panama, Limon Peninsula, Costa 

Eica. Living, Florida, Bahamas, and West Indies.

Conus species

A small poorly preserved cone (restored height about 

24 mm, diameter 13.5 mm) from the Caimito formation of the Gatun Lake area has a moderately low, 

evenly tapering spire and a wide shoulder. The shoulder is abruptly rounded and the anal fasciole is flat. 

A few low spiral threads are visible on the lower part 

of the body whorl. The outline is like that of Conus cf. 

C. marginatus Sowerby of Hubbard (1920, p. 162, pi. 

24, fig. 16), which occurs in limestone of early Miocene 

age in Puerto Eico (Aymamon limestone of present 

terminology). Conus marginatus Sowerby (1850, p. 44; 

not Conus marginatus J. de C. Sowerby, 1840), as identified by Guppy (1876, p. 528, pi. 29, fig. 5), has relatively strong spiral sculpture on the body whorl.

Occurrence: Caimito formation, Gatun Lake area 

(late Oligocene), locality 55b.

Conus bravoi Spieker 

Plate 56, figures 10, 11

Conus molis var. bravoi Spieker, Johns Hopkins Univ. Studies 

in Geology, no. 3, p. 41, pi. 1, fig. 6, 1922 (Miocene, Perti).

Conus (Dendroconus) bravoi Spieker, Olsson, Bull. Am. Paleontology, v. 19, no. 68, p. 151, pi. 16, figs. 1, 3, 4, 1932 

(Miocene, Peru). Marks, Idem, v. 33, no. 139, p. 137, 

1951 (Miocene, Ecuador).

Conus molis Brown and Pilsbry, Barrios, Colombia Servicio 

Geol. Nac., Bol. Geol., v. 6, nos. 1-3, (Informe 1082), p. 

294, pi. 12, fig. 8, 1960 (Miocene, Colombia).

Conus riosantiagensis Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst, p. 85, 

pi. 13, figs. 1, la, Ib, 1964 (Miocene, Ecuador).

Moderately large, broad-shouldered. Edge of shoulder marked by angulation, generally pronounced, but 

of varying strength. Spire low, profile smoothly conical, except part formed by strongly emerging early 

whorls. Protoconch missing. Early post-protoconch 

whorls worn, but showing suggestion of tuberculate 

shoulder. Succeeding whorls overlapping to edge of 

shoulder, or not quite to edge. Anal fasciole flat or 

somewhat concave, smooth (except for growth lines), 

or bearing faint to weak spiral threads. Spiral sculpture on lower part of body whorl faint or absent on 

mature shells, weak on immature shells.
GASTROPODS: EULIMIDAE, MARGESTELLIDAE TO HELMINTHOGLYPTIDAE 349

Height (incomplete) 64 mm, diameter 44 mm (larger 

figured specimen). Height 59.4 mm, diameter 36.6 mm 

(smaller figured specimen).

Type: Peabody Museum, Yale University.

Type locality: Zorritos district, Peru, Zorritos formation.

This Peruvian and Ecuadorean species is fairly abundant in the lower part of the Gatun formation and is 

represented in the middle part by two specimens. Fourteen, ranging in height from 23.5 to 57 mm, were collected at locality 138e and 23 others at four other 

lower Gatun localities.

Conus bravoi may be recognized by its stubby lowspired outline, and flat or somewhat concave anal fascicle, which lacks spiral sculpture or is faintly to 

weakly sculptured. Three immature shells from locality 

138c have a more distinctly concave anal fascicle than 

others. A few of the earliest post-protoconch whorls 

evidently have a tuberculate shoulder, but this feature 

is not clearly shown. The angulation at the shoulder is 

weak on the large specimen illustrated on plate 56, 

figure 11, as it is on C. riosantiagensis. It is also weak 

on the single specimen from the middle part of the 

Gatun. In the middle Miocene Tubara formation of 

Columbia C. bravoi reaches a height of 112 mm. Many 

of the specimens from that region have a faintly angulated, or even rounded shoulder.

A relatively elongate, moderately high-spired shell 

(locality 136a) is doubtfully referred to C. bravoi. The 

shoulder angulation is exposed on late spire whorls, 

but the overlap of the succeeding whorl is not uniform. 

This shell may be abnormal.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 136, 

136a, 137a, 138c, 138e, 1381 Middle part, eastern area, 

localities 155, 159d. Tubara formation (middle Miocene), Colombia. Progreso and Angostura formations 

(middle Miocene), Ecuador. Zorritos, Cardalitos, and 

Montera formations (middle Miocene), Peru.

Conus acolus Woodring, n. sp. 

Plate 55, figure 3

Small, broad-shouldered, edge of shoulder angulated. 

Spire of moderate height, profile evenly conical or 

almost evenly conical. Protoconch missing. Earliest 

post-protoconch whorls worn, shoulder angulation exposed on later whorls. Anal fascicle concave, marked 

only by growth lines. Lower part of body whorl sculptured with widely spaced spiral grooves.

Height 35 mm, diameter 20.8 mm (type).

Type: USNM 645741; 2 paratypes, Stanford University.

Type locality: 136a (Stanford University 2611, 

Transisthmian Highway, lat 9° 21' N. plus 1,100 feet 

(335 meters); long 79° 49' W., Panama; same as USGS 

16912), lower part of Gatun formation.

Conus acolus is a small species based on nine specimens in three collections from the lower part of the 

Gatun formation. It may be distinguished from immature shells of C. bravoi, with which it is associated, by 

its more evenly conical spire, more concave anal fasciole, and stronger spiral sculpture on the lower part 

of the body whorl.

C. cercadensis Maury (1917, p. 43, pi. 7, fig. 4; Cercado formation, Dominican Republic) has a more 

rounded shoulder; its late whorls overlap to the edge 

of the shoulder of the preceding whorl; and its spiral 

sculpture covers a larger part of the body whorl, although in reduced form. In outline C. acolus suggests 

a small version of C. bermudensis Clench (1942, p. 34, 

pi. 13, fig. 4), a living western Atlantic species from 

Bermuda and the east coast of Florida.

Occurrence: Lower part of Gatun formation (middle 

Miocene), localities 136, 136a, 138c. Middle Miocene 

deposits, Chiriqui area (USGS 7955, identification 

doubtful), Panama.

Conus cf. C. sauridens Conrad

The silicified fossils of the Rio Casaya area include 

a small incomplete, elongate, low-spired Conus that 

probably is allied to C. sauridens, a species of southeastern United States ranging in age from middle Eocene to early Oligocene (Palmer, 1937, p. 458, pi. 71). 

The shoulder is sharply carinate and the anal fascicle 

is concave. The sharply carinate shoulder shows on 

spire whorls. There is a suggestion of small tubercles 

on the shoulder of the earliest preserved whorls. No 

spiral sculpture is visible on the anal fascicle, but the 

granular silica is too coarse to reproduce low threads 

like those of C. sauridens. The restored height is 30 

mm, the diameter 15 mm.

A similar late Eocene and late Eocene or early Oligocene cone from Peru has been named C. chircensis 

(Olsson, 1930, p. 39, pi. 5, figs. 1, 2, 4,10). C. chircensis, 

treated as a subspecies of C. sauridens, has been identified in deposits of late Eocene age in Columbia (Clark, 

in Clark and Durham, 1946, p. 46, pi. 24, figs. 11, 16, 

18,19).

Occurrence: Gatuncillo formation, Rio Casaya area 

(middle Eocene), locality 38.

Conus cf. C. sulculus Ball

The upper part of the Bohio formation on Barro 

Colorado Island and the Caimito formation at a nearby 

locality yielded poorly preserved remains of a small
350 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

species of Comas. It is fairly slender and has a sharply 

angulated shoulder. The spire is of moderate height 

and its profile is concave. Late whorls overlap to, or 

not quite to, the shoulder angulation. The anal fascicle 

is slightly concave and bears (at least on the specimen 

of best preservation) weak spiral threads. The largest 

of 14 specimens has an almost complete height of 22 

mm and a diameter of 13 mm.

This unnamed species is smaller and more slender 

than Conus sulculus (Gardner, 1926-47, p. 359, pi. 43, 

fig. 3, 1938), of the early Miocene Chipola formation 

of Florida.

Occurrence: Upper part of Bohio formation (late 

Oligocene), locality 42d. Caimito formation, Gatun 

Lake area (late Oligocene), localities 56, 57.

Conus cf. C. planiceps Heilprin

Four lots of poorly preserved small cones from the 

La Boca formation suggest a species similar to Conus 

planiceps Heilprin (1887, p. 110, pi. 15, figs. 48, 48a), 

of the early Miocene Tampa limestone of Florida. The 

late whorls form a low, almost flat-topped spire, from 

which the earliest preserved whorls (missing on many 

of the specimens) abruptly rise. The shoulder is 

broadly angulated and the whorls overlap to, or almost 

to, the shoulder. The anal fascicle is barely concave 

and is sculptured with weak spiral threads. The largest 

specimen has an estimated height of 32 mm and a 

diameter of 19.5 mm.

C. planiceps reaches a height of 46 mm. As noted by 

Mansfield (1937, p. 80), the shoulder of the specimen 

from Tampa illustrated by Dall in 1915, p. 37, pi. 6, 

figs. 1, 2) is sharply angulated and shows on spire 

whorls, but on other specimens in the same lot it is 

less sharply angulated and fails to show on spire 

whorls. Ball's 1890 illustrations (Dall, 1890-1903, p. 

25, pi. 11, figs. 5, 5a, 1890) are poorly drawn or represent a different species. His brief remarks may be 

taken to indicate that the specimen was collected at 

Martin Station, Florida, by Willcox. Druid Wilson 

suggests that this fossil may be in the collections of 

the Philadelphia Academy of Natural Sciences.

A small cone from deposits of middle Miocene age 

in the Tehuantepec area of Mexico, recently identified 

as C. planiceps (Perrilliat Montoya, 1963, p. 27, pi. 6, 

figs. 3, 10), was described as bearing small tubercles 

on the shoulder of early spire whorls. Such tubercles 

are absent on C. planiceps.

Occurrence: La Boca formation (early Miocene), 

localities lOlh, 115a, 115b, 116a.

Conus molis Brown and Pilsbry 

Plate 55, figures 8-10

Conus molis Brown and Pilsbry, Acad. Nat. Sci. Phila. Proc., 

v. 63, p. 343, pi. 23, fig. 1, 1911 (Miocene, Canal Zone). 

Pilsbry and Brown, Idem, v. 69, p. 32 (list), 1917 (Miocene, Colombia). Olsson, Bull. Am. Paleontology, v. 9, 

no. 39, p. 42, pi. 2, figs. 1, 2(1), 1922 (Miocene, Costa 

Rica, Panama, Canal Zone). Weisbord, Idem, v. 14, no. 

54, p. 56, pi. 6, fig. 1, 1929 (Miocene, Colombia). Anderson, Calif. Acad. Sci. Proc., 4th ser., v. 18, no. 4, p. 109, 

1929 (Miocene, Colombia).

tConus molis Brown and Pilsbry, Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 36, 1917 (Miocene, Dominican Republic).

Conus concavitectum Brown and Pilsbry, Acad. Nat. Sci. Phila. 

Proc., v. 63, p. 341, pi. 23, figs. 5, 6, 1911 (Miocene, Canal 

Zone). Olsson, Bull. Am. Paleontology, v. 9, no. 39, p. 43, 

1922. (Miocene, Canal Zone, Panama). Anderson, Calif. 

Acad. Sci. Proc., 4th ser., v. 18, no. 4, p. 110, 1929 (Miocene, Colombia).

Conus (Lithoconus) concavitectum Brown and Pilsbry, Cossmann, Jour. Conchyliologie, v. 61, p. 43, pi. 4, figs. 3, 4, 

1913 (Miocene, Canal Zone).

fConus domingensis Sowerby?, Brown and Pilsbry, Idem, p. 

341, 1911 (Miocene, Canal Zone).

Not Conus molis Brown and Pilsbry, Barrios, Colombia Servicio 

Geol. Nac., Bol. Geol., v. 6, nos. 1-3 (Informe 1082), 

p. 294, pi. 12, fig. 8,1960 (Miocene, Colombia); = C. travoi 

Spieker.

Exceptionally large, elongate, moderately wide at 

shoulder. Edge of shoulder generally abruptly angulated. Spire of moderate height, its profile concave. 

Protoconch consisting of 2i/£ slightly bulging whorls. 

Early half of first post-protoconch whorl bearing axial 

riblets, angulated shoulder appearing on later half. 

Angulated shoulder of first five or six post-protoconch 

whorls exposed; first three or four slightly tuberculate. Anal fascicle generally concave, bearing spiral 

threads. Growth lines of fascicle generally exaggerated 

on some whorls. Lower part of body whorl, or entire 

whorl, bearing faint spiral sculpture, of decreasing distinctness upward. Spiral threads on lower part of very 

young shells faintly pustulose. Color pattern faintly 

showing on some immature shells, consisting of spiral 

rows of brownish crude rectangles, much like the pattern of Conus spurius.

Height (practically complete) 160 mm, diameter 90 

mm (larger figured specimen).

Type: Princeton University.

Type locality: Gatun Locks excavation, Canal Zone, 

middle part of Gatun formation.

Conus molis, based on a mature shell, is given precedence over C. concavitectum, the type of which is immature.

This large species the largest American fossil species is found throughout the Gatun formation. A total
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 351

of 67 specimens, ranging in height from 23.5 to about 

180 mm, is available. The largest occur in the upper 

part of the formation. That shown on plate 55, figure 

8, was collected in the western area. A still larger 

crushed shell from the eastern area would have a height 

of about 180 mm, if it were complete.

With the exception of two specimens, the basic features of these fossils are essentially uniform. The two 

exceptions are in a collection from the lower part of 

the Gatun (locality 136a). The shoulder of those specimens is less sharply angulated t]jan that of the others 

and their anal fascicle is flat or almost flat. Though 

the shoulder and fascicle suggest G. cemulator (the 

next species described), these two fossils are larger than 

that species and have the elongate outline of C. molis.

The distribution of C. molis is clustered around the 

southeastern border of the Miocene Caribbean Sea: in 

Costa Kica, Panama, and Columbia. It has a close Miocene ally in Ecuador and Peru: C. cacuminatus Spieker 

(1922, p. 40, pi. 1, fig. 5), which has more strongly 

tuberculate early whorls, a more deeply concave anal 

fascicle, and is not known to be of comparable size. C. 

haytensis Sowerby (Pflug, 1961, p. 60, pi. 16, figs. 1-5; 

Cercado and Gurabo formations, Dominican Republic) 

 the most similar fossil species in the West Indies 

is wider at the shoulder, and has a less concave anal 

f asciole and more strongly tuberculate early whorls. It 

is likely that Maury's C, molis is an immature C. Jiaytensis. On the contrary, Brown and Pilsbry's C. Jiaytensis and C. domingensisf, both from the Gatun formation, are likely to be immature C. molis. The stubby 

outline and flat anal f asciole of the small middle Miocene Venezuelan cone identified as G. aff. G. molis 

(Jung, 1965, p. 575, pi. 78, figs. 1, 2) indicate that it 

is more similar to G. (Emulator than to G. molis.

No similar species is living in the Caribbean Sea, but 

G. fergusoni Sowerby (Hanna, 1963, p. 42, pi. 4, fig. 2), 

living in the eastern Pacific Ocean, is closely allied, as 

pointed out by Hanna and Strong (1949, p. 295) and 

Hanna (1963, p. 44). Though the two species are similar in size and outline, the anal fascicle of C. molis is 

generally more concave and more strongly sculptured, 

and its early whorls are less distinctly tuberculate.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 136a, 137a, 138c, 138d. Middle part, eastern area, 

localities 139e, 139f, 146, 147b (identification doubtful), 147f, 147h (identification doubtful), 151,153,155, 

155a, 155b, 155c, 157, 158 (identification doubtful), 

159,159b. Upper part, eastern area, localities 171 (identification doubtful), 172, 175, I76a, 177, 177a, 177b, 

I77c; western area, locality 185. Middle Miocene, deposits, Costa Rica. Tubara formation (middle Miocene), Colombia. Limon formation (late Miocene), 

Bocas del Toro area, Panama.

Conus aemulator aemulator Brown and Pilsbry 

Plate 55, figures 5, 6; plate 56, figures 4, 8

Conus (emulator Brown and Pilsbry, Acad. Nat. Sci. Phila. 

Proc., v. 63, p. 342, pi. 23, fig. 9, 1911 (Miocene, Canal 

Zone). Pilsbry and Brown, Idem, v. 69, p. 32 (list), 1917 

(Miocene, Colombia).

Not Conus proteus (emulator Brown and Pilsbry, Rutsch, 

Schweizer. Palseont. Gesell. Abh., v. 55, no. 1, p. 104, pi. 9, 

figs. 7-11, 1934 (Miocene, Venezuela) ; = C. (Emulator 

manzanillcensis Mansfield.

Conus Uaytensis Sowerby, Brown and Pilsbry, Acad. Nat. Sci. 

Phila. Proc., v. 63, p. 341, 1911 (Miocene, Canal Zone).

Conus veatchi Olsson, Bull. Am Paleontology, v. 9, no. 39, p. 44, 

pi. 2, figs. 5, 8, 1922 (Miocene, Panama, Canal Zone). 

Woodring and Mansfield, in Geology of Republic of 

Haiti, p. 175, pi. 15, figs. 1, 2, 1924 (Miocene, Haiti). 

Anderson, Calif. Acad. Sci. Proc., 4th ser., v. 18, no. 4, 

p. 108,1929 (Miocene, Colombia). Trechmann, Geol. Mag., 

v. 72, no. 858, p. 545, pi. 21, fig. 10, 1935 (Miocene, Carriacou). Olsson, Neogene mollusks of northwestern Ecuador, Paleontological Research Inst., p. 82, 1964 (Miocene, 

Ecuador).

f Conus sp., Toula, K. k. Geol. Reichsanstalt Jahrb., v. 58, p. 

710, pi. 25, fig. 18, 1909 (Miocene, Canal Zone). Engerrand and Urbina, Soc. Geol. Mexicana BoL, v. 6, pi. 58, 

fig. 16, 1910 (reproduction of Toula's illustration).

Of medium size, stubby, wide at shoulder. Edge of 

shoulder angulated. Spire low, practically flat-topped, 

except part formed by earliest whorls. Protoconch and 

earliest post-protoconch whorls not well preserved. 

Angulated shoulder of first three or four post-protoconch whorls exposed; the shoulder slightly tuberculate. Anal fascicle practically flat or slightly concave, 

bearing spiral threads. Lower part of body whorl bearing weak spiral sculpture; the threads faintly pustulose on very young shells.

Height 45 mm, diameter 28 mm (larger figured 

specimen).

Type: Acad. Nat. Sci. Phila. 1691.

Type locality: Gatun Locks excavation, Canal Zone, 

middle part of Gatun formation.

The moderate size, stubby outline, flat-topped late 

spire whorls, and spirally sculptured anal fasciole are 

diagnostic features of Conus (emulator. The type (pi. 

55, figs. 5, 6), photographed by A. A. Olsson, is immature (height 22.7 mm) and therefore does not show the 

flat-topped late spire whorls. The largest specimen 

examined (height 59 mm) was found on Rio Chico, in 

Darien (USGS 8433). The less elongate outline, flattopped late spire whorls, and flatter anal fasciole distinguish mature shells of G. emulator from immature 

shells of G. molis. The two species are associated at 

many localities and in many features are similar.
352 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

C. (Emulator is fairly widespread in the Gatun formation, but so far has not been found in the upper part 

of the formation in the eastern area. The 42 available 

specimens are essentially uniform, with the exception 

of two from the upper part of the Gatun in the western 

area the only specimens in the upper Gatun collections. The edge of the shoulder of those two shells is 

less sharply angulated than that of the others. Toula's 

Conus sp. is missing in his collection. His illustration 

suggests an immature C. (Emulator.

This species has a wider distribution than C. molis 

and, unlike C. molis, occurs on both sides of Central 

America. It has also a greater time range: late early to 

late Miocene. C. manzanillasnsis Mansfield (1925, p. 12, 

pi. 2, figs. 5, 10; San Jose calcareous silt member of 

Manzanilla formation, Trinidad), is assigned to subspecific rank under C. (Emulator. It is smaller than the 

nominate subspecies and has faintly pustulose spiral 

sculpture on the lower part of the body whorl, like 

immature specimens of the nominate subspecies. The 

anal fascicle of the type is very weakly sculptured, but 

the sculpture is stronger on other specimens. The largest in a collection of three (height 36 mm) from the 

San Jose calcareous silt member on Guaracarita River 

(USGS 21098) is high-spired, like Eutsch's Venezuelan specimens.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 136a, 138,138a, 138c, 138d, 138e, 1381 Middle part, 

eastern area, localities 139c, 139e, 139f, 147h, 155, 155a, 

155b, 155c, 159, 159d, 160 (identification doubtful). 

Upper part, western area, locality 182. Baitoa formation (early Miocene), Dominican Republic. Thomonde 

formation (early Miocene), Haiti. Deposits of early 

Miocene age, Carriacou, Grenadine Islands. Deposits 

of early (?) Miocene age, Ecuador. Cercado and Gurabo formations (middle Miocene), Dominican Republic. Tubara formation (middle Miocene), Colombia. 

Middle Miocene deposits, Darien area (USGS 8429, 

8433, 8477, 8479, 8492), Panama. Limon formation 

(late Miocene), Bocas del Toro area, Panama.

Conus cf. C. catenatus Sowerby

Small, moderately slender, angulated at shoulder. 

Spire high, its profile concave. Angulated and tuberculate shoulder appearing on first post-protoconch 

whorl. Tubercles gradually disappearing on third 

whorl. Last few whorls overlapping almost to shoulder 

of preceding whorl. Anal fascicle flat or slightly concave, bearing weak spiral threads or none. Spiral sculpture of body whorl weak, faintly to distinctly pustulose or nonpustulose.

Height 25.7 mm, diameter 12 mm (largest specimen).

Four small cones in two collections from the Gatun 

Third Locks site are identified as Conus cf. C. catenatus. Without much doubt they are immature. In some 

features they suggest C. catenatus (Pilsbry, 1922, p. 

328, pi. 22, figs. 3, 4; Woodring, 1928, p. 213, pi. 11, 

figs. 4, 5), a middle Miocene species from the Dominican Republic and Jamaica. At the growth stage represented by the last few whorls of the Gatun fossils the 

shoulder of C. catenatus is not as strongly angulated.

Occurrence: Middle part of Gatun formation, eastern 

area (middle Miocene), localities 155, 155a.

Conus consobrinus consobrinus Sowerby 

Plate 56, figures 3, 7, 9

Conus consobrinus Sowerby, Geol. Soc. London Quart. Jour., 

v. 6, p. 45, 1850 (Miocene, Dominican Republic). Maury, 

Bull. Am. Paleontology, v. 5, no. 29, p. 39, pi. 6, figs. 5, 6, 

1917 (Miocene, Dominican Republic). Pilsbry, Acad. Nat. 

Sci. Phila. Proc., y. 73, p. 330, pi. 20, figs. 7, 7a, 7b, 1922 

(Miocene, Dominican Republic).

Conns (Leptoconus) consobrinus Sowerby, Woodring, Carnegie 

Inst. Washington Pub. 385, p. 214, pi. 11, figs. 6, 7, 1928 

(Miocene, Jamaica). Perrilliat Montoya, Mexico Univ. 

Nac., Inst. Geologia Paleontologia Mexicana, no. 8, p. 26, 

pi. 4, figs. 3, 4, 1960 (Miocene, Mexico). Pflug, Acta 

Humboldtiana, Geol. Palseontologica Ser., no. 1, p. 62, 

pi. 17, figs. 1-10, 1961 (Miocene, Dominican Republic).

?Conus consobrinus Sowerby?, Anderson, Calif. Acad. Sci. Proc., 

4th ser., v. 18, no. 4, p. Ill, 1929 (Miocene, Colombia).

Not Conus consobrinus Sowerby, Gabb, Acad. Nat. Sci. Phila. 

Jour., 2d ser., v. 8, p. 359, 1881 (Pliocene, Costa Rica) ; 

= C. consobrinus ultimug Pilsbry and Johnson.

Not Conns consobrinus Sowerby, Brown and Pilsbry, Acad. Nat. 

Sci. Phila. Proc., v. 63, p. 341, 1911 (Miocene, Canal 

Zone) ; = C. tortuosostriatus Toula.

Not Conus (Chelyconus) consobrinus Sowerby, Cossmann. Jour. 

Conchyliologie, v. 61, p. 46, pi. 3, figs. 17, 18, 1913 (Miocene, Martinique).

Conus (Conospira) lavillei Cossmann, Jour. Conchyliologie, v. 

61, p. 38, pi. 4, figs. 1, 2, 1913 (Miocene, Canal Zone).

Conus granasonatoides Maury, Bull. Am. Paleontology, v. 5, 

no. 29, p. 39, pi. 6, fig. 7, 1917 (Miocene, Dominican 

Republic).

Of medium size, elongate, moderately wide at shoulder. Shoulder sharply angulated. Anal fascicle slightly 

sloping, producing an almost square shoulder. Spire 

high and its profile almost smoothly conical, or moderately high and its profile slightly concave. Protoconch and earliest post-protoconch whorls missing. 

Shoulder of all except last, or last two, spire whorls 

strongly tuberculate; tubercles gradually disappearing. 

Anal fascicle concave, bearing faint or distinct spiral 

threads. Lower part of body whorl weakly sculptured; 

the threads faintly pustulose on immature shells.

Height 61.7 mm, diameter 30.3 mm (larger figured 

specimen).
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 353

Type material: Lectotype, British Museum (Natural 

History) G83962.

Type locality: Valley of Rio del Nbrte, Dominican 

Republic, Miocene.

Conus consobrinus consobrinus has a meager representation in the Gatun formation: two specimens from 

the lower part, two from the middle part, and one from 

the upper part in the western area. Cossmann's C. lavillei, collected along the canal at Hindi, where the upper 

part crops out, is identified as an exceptionally slender, 

immature specimen of this form. The two shells from 

the lower part, one of which is shown on plate 56, 

figures 3, 7, and one from a horizon near the base of 

the middle part are relatively low-spired an exceptional feature. The large shell from the middle part 

in the western area (pi. 56, fig. 9) is the only one that 

is fully mature. Its spire is somewhat worn. Gatun 

specimens are almost square-shouldered, as are most 

others from mainland localities. Those from Jamaica 

are round-shouldered and so are most of those from 

the Dominican Republic.

Bose's illustration of the Mexican C. scalice (Bb'se, 

1906, p. 51, pi. 5, figs. 41, 42) suggests an immature 

C. consobrinus. His type (height 15.2 mm), however, 

lacks spiral sculpture on the anal fascicle. That C. 

consobrinus consobrinus is found in deposits of middle 

Miocene age in the Tehuantepec area is shown by Perillat Montoya's illustration and by a specimen in the 

collections of the IT. S. National Museum (USGS 

10346).

The high, or moderately high, strongly tuberculate 

spire is a diagnostic feature of C. consobrinus. The type 

material (lectotype) has been designated and illustrated recently by Pflug. The nominate subspecies occurs in deposits of middle Miocene age. The lineage is 

continued by an almost square-shouldered form that 

has a tuberculate shoulder throughout, even on the 

body whorl of specimens that have a height of 50 mm. 

This form is C. consobrinus ultimus Pilsbry and Johnson (Pilsbry, 1922, p. 330, pi. 20, fig. 8). It first appears 

in the late Miocene Limon formation of southeastern 

Costa Rica and continues in the Pliocene Moin formation of that area. C. torcensis Olsson (1922, p. 48, pi. 2, 

fig. 7) may be treated as a weakly sculptured late Miocene subspecies. G. emersoni Hanna (1963, p. 25, pi. 1, 

fig. 2), dredged off Cape San Lucas, Baja California, 

is probably a descendant of G. consobrinus. It is roundshouldered and lacks tubercles on the shoulder of the 

body whorl, and therefore is similar to G. consobrinus 

consobrinus.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, locality 138c. Middle part, eastern area, locality 139d; western area, locality 161b. Upper part, eastern area, Cossmann's record; western area, locality 185. Middle Miocene deposits, Darieii area (small race, USGS 8430, 

8477), Panama. Agueguexquite formation (middle 

Miocene), Tehuantepec area, Mexico. Bowdeii formation (middle Miocene) Jamaica. Gurabo formation 

(middle Miocene), Dominican Republic.

Conus symmetricus Sowerby 

Plate 57, figures 13, 14

Conus symmetricus Sowerby, Geol. Soc. London Quart. Jour., 

v. 6, p. 44, pi. 9, fig. 1, 1850 (Miocene, Dominican Republic). Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 36, 

pi. 7, figs. 7, 7a, 1917 (Miocene, Dominican Republic). 

Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 73, p. 328, pi. 20, 

figs. 2, 2a, 2b, 1922 (Miocene, Dominican Republic). Not 

Conus symmetricus Sandberger, 1862.

Conus (Leptoconus) symmetricus Sowerby, Pfiug, Acta Humboldtiana, Geol. Paleontologica Ser., no. 1, p. 63, pi. 18, 

figs. 1-11, 1961 (Miocene, Dominican Republic).

Conus symmetricus semiobsoletus Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 37, pi. 7, fig. 8, 1917 (Miocene, Dominican Republic).

Conus symmetricus domingensis Sowerby, Sowerby, Maury, 

Idem, p. 37, pi. 4, fig. 10 (Miocene, Dominican Republic).

fConus (Lithoconus) sp., Woodring, Carnegie Inst. Washington 

Pub. 385, p. 204, 1928 (Miocene, Jamaica).

Small, wide at shoulder, shoulder sharply angulated. 

Spire low, its profile slightly concave. Protoconch 

worn, consisting of about two whorls. Earliest postprotoconch whorls somewhat worn. Shoulder of about 

first three exposed, obscurely tuberculate. Later whorls 

overlapping to shoulder of preceding whorl. Anal fasciole flat, or almost flat, sculptured with spiral threads. 

Body whorl sculptured with primary and secondary 

spiral threads. Primary threads bearing pustules, arranged in axial series, or not so arranged.

Height 20.4 mm, diameter 11.8 mm (figured specimen).

Type material: Lectotype, British Museum (Natural 

History) G83696.

Type locality: Valley of Rio Yaque del Norte, 

Dominican Republic, Miocene.

Two specimens, both immature and both collected at 

the Gatun Third Locks site, are referred to this small, 

broad-shouldered, delicately sculptured, pustulose species. The larger of the two is illustrated. On the early 

part of the body whorl of this specimen slight swelling 

along the alinement of the pustules produces slight 

axial wrinkles.

Though the unidentified Conus from Bowden, Jamaica, may be a corroded Conus symmetricus, this is 

the first unequivocal record for that species beyond the 

Dominican Republic, where it is abundant in the Gurabo formation. Sowerby's illustration is unmistakable.
354 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Pflug recently designated and illustrated the lectotype. 

For the most part height at maturity is not more than 

35 mm. On the largest shells (height about 40 mm) 

the sculpture fades out on the upper part of the body 

whorl: Maury's C. symmetricus semiobsoletus. The 

spire is low and may be so low that the shell is almost 

flat-topped: Maury's G. symmetricus domingensis. Pflug 

(1961, p. 61) preferred to hold in abeyance designation 

of a lectotype of C. domingensis Sowerby (1850, p. 45). 

Occurrence: Middle part of Gatun formation, eastern area (middle Miocene), localities 155a, 155c. Gurabo formation (middle Miocene), Dominican Republic.

Conns taphrus Woodring, n. sp. 

Plate 57, figures 1, 7

Small, moderately wide at shoulder, edge of shoulder angulated. Protoconch and earliest post-protoconch 

whorls missing. Earliest preserved whorls carinate and 

obscurely tuberculate; tubercles obscure owing to wear. 

Succeeding whorls overlapping to edge of shoulder of 

preceding whorl, bearing a narrow channel adjoining 

suture. Anal fascicle flat, bearing growth threads and 

microscopic spiral lineation. Lower part of body whorl 

weakly sculptured. Entire whorl showing microscopic 

spiral lineation.

Height (almost complete) 28.2 mm, diameter 15 mm 

(type).

Type:USNM645T48.

Type locality: 161d (USGS 8366, Cuts west of 

Gatun Dam, station 3a, Canal Zone), middle part of 

Gatun formation.

The type the only specimen was collected at a 

locality in the middle part of the Gatun formation west 

of Gatun Dam. Though it may not be mature, the 

sutural channel is a distinctive feature.

Occurrence: Middle part of Gatun formation, western area (middle Miocene), locality 161d.

Conns aff. C. chipolanus Ball

Formations of late Oligocene and early Miocene age 

yielded 19 cones of minute, small, or medium size: nine 

from the upper part of the Bohio formation of Barro 

Colorado island, one from the Caimito formation of 

the Gatun Lake area, one from the Culebra formation, 

and eight from the La Boca formation. The Bohio 

fossils are minute shells, three of which show a slender 

protoconch of three whorls. If the first few post-protoconch whorls are sculptured, the sculpture is obscure. 

A carinate shoulder appears at an early stage and 

shows on the remaining whorls of the high spire. One 

of the La Boca specimens shows a similar crushed protoconch. The lower part of the body whorl of all these 

fossils bears distinct spiral sculpture. The largest, the

Culebra specimen (locality 104b) has an almost complete height of 31.3 mm and a diameter of 15.2 mm. Its 

shell material, like that of the others, is replaced by 

calcite.

Though these fossils are unsatisfactory, they are allied to C. chipolanus^ which is discussed under the next 

heading, and show that in the Canal Zone the lineage 

of G. imitator (the next species described) extends 

back to late Oligocene time. Similar small cones occur 

in deposits of late Oligocene age in Florida and Alabama : Mansfield's G. aff. G. imitator (Mansfield, 1938, 

p. 101, fig. 4; 1940, p. 203). In southeastern United 

States, however, the lineage is unknown later than 

early Miocene time.

Occurrence: Upper part of Bohio formation (late 

Oligocene), locality 42d. Caimito formation, Gatun 

Lake area (late. Oligocene), locality 56. Culebra formation (early Miocene), locality 104b. La Boca formation (early Miocene), localities lOlh, 115a, 116a.

Conns imitator imitator Brown and Pilsbry 

Plate 55, figures 1, 2

Conus imitator Brown and Pilsbry, Acad. Nat. Sci. Phila. Proc., 

v. 63, p. 342, pi. 23, fig. 4, 1911 (Miocene, Canal Zone). 

Pilsbry and Brown, Idem, v. 69, p. 32,1917 (list, Miocene, 

Colombia). Olsson, Bull. Am. Paleontology, v. 9, no. 39, 

p. 45, pi. 2, fig. 6, 1922 (Miocene, Canal Zone, Panama, 

Costa Rica). Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 73, 

p. 327, 1922 (Miocene, Dominican Republic). Anderson, 

Calif. Acad. Sci. Proc., 4th ser., v. 18, no. 4, p. 108, 1929 

(Miocene, Colombia). Perrilliat Montoya, Mexico Univ. 

Nac., Inst Geologla, Paleontologia Mexicana, no. 14, p. 

27, pi. 6, figs. 6, 7,1963 Miocene, Mexico). Barrios, Colombia Servicio Geol. Nac., Bol. Geol., v. 6, nos. 1-3 (Informe 

1082), p. 295, pi. 12, fig. 6, 1960 (Miocene, Colombia).

Not Conus imitator Brown and Pilsbry, Li, Geol. Soc. China 

Bull., v. 9, p. 275, pi. 8, figs. 72 (mislabelled 70), 72a, 

1930 (Miocene, Panama Bay;=C'. maliogani Reeve, fide 

Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 83, p. 434, 1931, 

living, Panama Bay).

Conus (Leptoconus) imitator lius Woodring, Carnegie Inst. 

Washington Pub. 385, p. 209, pi. 10, figs. 5, 6, 1928 (Miocene, Jamaica).

fConus cf. imitator lius Woodring, Olsson, Neogene mollusks 

from northwestern Ecuador, Paleontological Research 

Inst, p. 84, 1964 (Miocene or Pliocene, Ecuador).

Conus dalli Toula, K. k. Geol. Reichsanstalt Jahrb., v. 61, p. 

509, pi. 31, fig. 23a-d, 1911 (Miocene, Canal Zone). Maury, 

Bull. Am. Paleontology, v. 5, no. 29, p. 48, pi. 7, fig. 15, 

1917 (Miocene, Dominican Republic). Alaneaster-Ibarra, 

Asoc. Mexicana Geologos Petroteros Bol., v. 2, p. 570, fig. 

17, 1950 (Miocene, Mexico). Not Conus dalli Stearns, 

1873.

Conus (Lithoconus) dalli Toula, Cossmann, Jour. Conehyliologie, v. 61, p. 41, pi. 3, figs. 30, 31, pi. 4, figs. 7, 8, 1913 

(Miocene, Oanal Zone).

Of medium size, moderately wide at shoulder, shoulder strongly carinate. Spire high, its profile almost
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 355

smoothly conical. Protoconch slender, consisting of 

three whorls, slightly enlarging in diameter. Earliest 

part of first post-protoconch whorl bearing one or 

several axial riblets, followed by appearance of carinate shoulder, generally tuberculate on first three or 

four whorls. Tubercles axially compressed and extending in diminished form to suture of succeeding whorl. 

Tubercles rarely absent or practically absent. Except 

on earliest whorls, succeeding whorl generally missing 

carinate shoulder by a considerable margin; rarely just 

missing it. Anal fascicle distinctly concave to almost 

flat, bearing faint microscopic spiral sculpture and 

conspicuous growth threads, exaggerated at fairly regular intervals, especially on early whorls. Lower part of 

body whorl bearing fairly strong or strong spiral sculpture. On immature shells spiral sculpture generally 

limited to lower part of whorl, rarely extending in 

subdued form to shoulder.

Height 49 mm, diameter 23.7 mm (largest specimen 

in collections at Washington). Height 45.4 mm, diam- 

'eter 22 mm (larger figured specimen).

Type: Acad. Nat. Sci. Phila. 1688.

Type locality: Gatun Locks excavation, Canal Zone, 

middle part of Gatun formation.

Conus imitator imitator occurs throughout the Gatun 

formation and its range may continue into the Chagres 

sandstone. The one Chagres specimen is too incomplete 

for unequivocal identification. Seventy of these cones 

are available. The largest (height 55 mm), in Thompson's collection at Balboa, was found in the middle 

part of the Gatun at locality 155, the largest number 

(20) in the upper part in the eastern area at locality 

177b. As was evident from his illustrations, Toula's 

C. dalli, a junior homonym, is C. imitator imitator. 

The specimen (height 26.7 mm, diameter 12.2 mm) 

shown in his figure 23b is herewith designated the 

lectotype.

Forty-five of these fossils show the early post-protoconch whorls, which are tuberculate on all except 

four. On the four exceptions the tubercles are absent, 

although a few faint axial riblets are discernible on the 

earliest part of the first of these whorls. The nontuberculate shells were collected at lower Gatun localities. 

On the contrary, four other lower Gatun shells, one of 

which is shown on plate 55, figure 2, are tuberculate. 

The Miocene Jamaican C. imitator lius was named on 

the grounds that it is nontuberculate grounds that 

are unacceptable on the basis of the Gatun samples. As 

a matter of fact, Olsson (1922, p. 154) reported that 

Jamaican specimens in his collection are tuberculate. 

The Ecuadoran fossils identified as C. cf. C. lius are 

poorly preserved.

The overlap of whorls on the preceding whorl is of

variable extent. The carina on the shoulder of the preceding whorl is generally missed by a considerable margin, but it may be just missed. The overlap difference 

is slight in vertical dimension, but the effect on spire 

profile is pronounced. The sculpture on the lower part 

of the body whorl consists of narrow grooves separating wider bands, or of narrow threads separated by 

wider spaces. The threads are flat or slightly shelflike.

C. imitator imitator may be recognized by its slender, 

cylindrical protoconch, moderate size, sharply carinate 

shoulder, strong growth threads on the anal fascicle, 

emphasized at fairly regular intervals, and distinct 

spiral sculpture on the lower part of the body whorl. 

It is inferred to be descended from C. chipolanus Ball 

(Gardner, 1926-47, p. 360, pi. 43, fig. 6, 1938), or from 

the late Oligocene and early Miocene form identified as 

C. aff. C. chipolanus. It is found along the west and 

south borders of the present Caribbean Sea in deposits 

of middle and late Miocene age, and in the middle 

Miocene of the Darien area, Jamaica, and the Dominican Republic. C. sophus Olsson (1932, p. 154, pi. 16, 

figs. 6, 8, 9; middle Miocene, Peru and Ecuador) may 

be treated as a small (height 15 to 32 mm) subspecies 

of C. imitator. The early postprotoconch whorls are 

nontuberculate (Olsson, 1932, p. 154) or only the first 

is tuberculate (Olsson, 1964, p. 85). A still smaller 

subspecies (height 8 mm), which is nontuberculate 

and has a 2-whorled protoconch, is found in the early 

Miocene Baitoa formation of the Dominican Republic 

and the equivalent Thomonde formation of Haiti. C. 

springvalensis Mansfield (1925, p. 11, pi. 1, figs. 3, 6; 

upper Miocene, Trinidad) is considered to be a slightly 

tuberculate or nontuberculate subspecies of C. imitator 

that lacks exaggerated growth threads on the anal 

fascicle and is consistently sculptured with narrow 

grooves on the lower part of the body whorl. The type 

is immature and poorly preserved. The weakly denticulate carina mentioned by Mansfield is the result of 

corrosion.

C. aff. C. imitator (Jung, 1965, p. 579, pi. 78, fig. 12; 

middle Miocene, Venezuela), which has a strongly flaring outer lip and heavy shoulder carina, is not closely 

related to C. imitator.

The C. chipolanus-imitator lineage can be traced 

with reasonable assurance to C. arcuatus Broderip and 

Sowerby (Hanna, 1963, p. 40, pi. 3, figs. 4-6), now living at moderate depths (15 to 50 fathoms) in the eastern Pacific Ocean, from the Gulf of California to 

Panama. C. arcuatus tends to be more slender and to 

have a higher spire and more steeply sloping anal fasciole. A complete protoconch of the living species is 

not available, but the last whorl of the protoconch 

suggests that it is slender.
356 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

C. imitator imitator is more similar to G. arcuatus 

than to the western Atlantic species C. floridanus Gabb 

(Clench, 1942, p. 27, pi. 13, figs. 1, 2). Late whorls of 

the western Atlantic species overlap to the carina of 

the preceding whorl or reach it, and the anal fascicle 

of late whorls is flat or practically flat; the protoconch 

is blunt and l^-to 2-whorled. A small middle Miocene Costa Bican form has been named C. -floridanus 

costaricensis (Olsson, 1922, p. 45, pi. 3, figs. 3, 9). In 

the type region it is associated with C. imitator imitator. Very young specimens of the two species may be 

distinguished by their protoconchs. Perhaps C. 'floridanus represents a branch of the chipolanus-imitator 

lineage adapted to shallower water than C. arcuatus: 

a few fathoms to 25 fathoms, exceptionally up to 40 

fathoms. Since Miocene time C. floridunus, like Antillophos candei (p. 265), has withdrawn northward from 

the Caribbean Sea.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 136a, 137, 137a, 138c, 138d, 138f. Middle part, 

eastern area, localities 155, 155a, 155b, 159, 159d; western area, locality 161a. Upper part, eastern area, localities 171, 172, 173, 175,177, 177a, 177b, 177d, 178; western area, locality 182. Chagres sandstone (late Miocene), locality 208 (Conus imitator?}. Middle Miocene 

deposits, southeastern Costa Eica, Tehuantepec area, 

Mexico, Colombia, Darien area (USGS 8477), Panama. 

Bowden formation (middle Miocene), Jamaica. Cercado and Gurabo formations (middle Miocene), Dominican Republic). Limon formation (late Miocene), 

Bocas del Toro area, Panama, Limon Peninsula, Costa 

Rica.

Conus multiliratus mnltiliratus Bose

Plate 57, figures 3, 4

Conus agassisi multiliratus Bose, Inst. Geol. Mexico Col. 22, 

p. 49, pi. 5, figs. 34-38 (multistriatus in explanation of 

plate), 1906 (Miocene, Mexico).

Conus (Leptoconus) multiliratus Bose, Marks, Bull. Am. Paleontology, v. 33, no. 139, p. 139, 1951 (Miocene, Ecuador).

conns multistriatus Bose, Olsson, Idem. v. 9, no. 39, p. 54, pi. 1, 

figs. 21, 23,1922 (Miocene, Panama).

Conus gaza Johnson and Pilsbry, in Brown and Pilsbry, Acad. 

Nat. Sci. Phila. Proc., v. 63, p. 342, pi. 23, figs. 2, 3, 1911 

(Miocene, Dominican Republic, Canal Zone). Pilsbry and 

Brown, Idem, v. 69, p. 32 (list), 1917 (Miocene, Colombia). Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 46, 

pi. 7, fig. 12, 1917 (Miocene, Dominican Republic).

Not Conus multiliratus gasa Johnson and Pilsbry, Spieker, 

Johns Hopkins Univ., Studies in Geology, no. 3, p. 37, 

1922 (Miocene, Peru).

Conus (Leptoconus) multiliratus gaza Johnson and Pilsbry, 

Woodring, Carnegie Inst. Washington Pub. 385, p. 212, 

pi. 11, fig. 3, 1928 (Miocene, Jamaica). Oinomikado, Geol. 

Soc. Japan Jour., v. 46, p. 626, pi. 29, fig. 30, 1939 (Miocene, Colombia).

Conus (Leptoconus) planiliratus Sowerby, Cossmann, Jour.

Conchyliologie, v. 61, p. 48, pi. 3, figs. 24-26 (not fig. 27;

= C. planiliratus Sowerby), 1913 (Miocene, Canal Zone,

Jamaica). 

Conus marginatus Sowerby, Maury, Bull. Am. Paleontology, v.

5, no. 29, p. 46, pi. 7, fig. 11, 1917 (Miocene, Dominican

Republic).

Small, biconical, wide at shoulder, shoulder strongly 

carinate. Spire high, its profile almost smoothly conical. Protoconch slender, cylindrical, 2i/£- to 3-whorled. 

Earliest part of first post-protoconch whorl bearing 

three to five axial riblets, followed by appearance of 

carinate shoulder, tuberculate on about first three 

whorls. Except on earliest whorls, succeeding whorl 

overlapping almost to carina. Anal fascicle slightly 

concave, bearing growth threads exaggerated at fairly 

regular intervals. Entire body whorl sculptured with 

strong, narrow, straplike spiral cords. On unworn 

shells exaggerated axial growth threads conspicuous 

in channels between cords, weaker or absent on cords 

themselves. Cords of immature shells rarely bearing 

low tubercles.

Height (almost complete) 27.2 mm, diameter 14.4 

mm (figured specimen).

Type: Apparently lost, but may eventually be found 

at the Instituto de Geologia, Universidad Nacional de 

Mexico.

Type locality: Tuxtepec, Oaxaca, Mexico, Miocene.

Though this strongly sculptured cone occurs in the 

three divisions of the Gatun formation but not in the 

upper part in the western area more than one or two 

specimens were collected at only two localities: nine at 

locality 138c and 10 at locality 139c. It is characterized 

by its broad-shouldered, biconical outline, and exaggerated growth threads on the anal fascicle and in the 

channels between the straplike spiral cords. The protoconch and spire whorls are like those of Conus imitator imitator, except that spiral threads are not discernible on the anal fascicle. One immature shell 

(height 17.8 mm, locality 138c) shows low tubercles on 

the spiral cords.

Contrary to the view that was adopted in 1928, C. 

gaza is treated as a synonym of C. multiliratus 'multiliratus. Though no specimens of the nominate subspecies 

are available, Bose's illustrations show the essential features of C. gaza. C. multiliratus 'multiliratus is recognized in middle Miocene formations in the Tehuantepec area, Panama, northeastern and southwestern 

Columbia, Jamaica, the Dominican Republic, and possibly in Ecuador, and in deposits of late Miocene age 

in northwestern Panama. A comparable form of the 

species, perhaps the same as that in Ecuador, occurs in 

the middle Miocene of the Darien area. It lacks strong 

growth threads between the spiral cords, but the ab-
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 357

sence of that feature may be a matter of preservation 

and perhaps should not be overemphasized.

C. multiliratus walli Mansfield (1925, p. 13, pi. 2, figs. 

1, 9; middle Miocene part of Brasso formation) may 

be recognized as a Trinidad race that is not as broadshouldered as the nominate subspecies. The type, which 

is somewhat worn, and the other shells in the type lot 

show scattered low tubercles on the spiral cords and 

slight undulations on the shoulder carina of the body 

whorl. Spieker's two specimens from the Zorritos formation of Peru, identified as C. multilirat-us gaaa,, are 

no longer available. When they were examined many 

years ago they were found to have weaker spiral cords 

and less distinct growth threads than C. multiliratus 

multiliratus. C. multiliratus spiekeri Olsson (1932, p. 

153, pi. 16, figs. 5, 7), also from the Zorritos formation, 

is sculptured with heavy tuberculate spiral cords and 

may be given specific rank. C. turbinopsis Gardner 

(1926-47, p. 361, pi. 43, fig. 12, 1938; Shoal Eiver formation, Florida) has a turbinate outline and less conspicuous growth threads between the spiral cords. It 

has been identified in the Tehuantepec area (Perrilliat 

Montoya, 1960, p. 27, pi. 4, figs. 5, 6).

C. clarki Kehder and Abbott (1951, p. 22, fig. 6), 

dredged off Louisiana at a depth of 29 fathoms, is 

sculptured with exaggerated growth threads on the 

anal fascicle and between the spiral cords. Its outline, 

however, is turbinate and the shoulder of all the postprotoconch whorls is tuberculate. C. frisbeyce Clench 

and Pulley (Clench, 1953, p. 369, pi. 184, fig. 1; Campeche Banks, Yucatan) evidently is C. clarki.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 138c, 138d. Middle part, eastern area, localities 

139c, 139e, 151, 159d. Upper part, eastern area, localities 173, 175, 176, 176a. Middle Miocene deposits, 

Tehuantepec area, Mexico, northeastern and southwestern Columbia. Bowden formation (middle Miocene), 

Jamaica. Cercado and Gurabo formations (middle Miocene). Dominican Kepublic. Limon formation (late 

Miocene), Bocas del Toro area, Panama.

Conus burckhardti burckhardti Bose 

Plate 57, figures 19, 20

Conus burckhardti Bose, Inst. Geol. Mexico Bol. 22, p. 50, pi. 5, 

figs. 39, 40, 1906 (Miocene, Mexico). Olsson, Bull. Am. 

Paleontology, v. 9, no. 39, p. 224, pi. 3, figs. 4, 5, 1922 

(Miocene, Panama, Costa Rica). Maury, Idem, v. 10, no. 

42, p. 187, pi. 34, fig. 5, 1925 (Miocene, Trinidad). Rutsch, 

Schweizer. Palseont. Gesell. Abh., Band 55, no. 1, p. 102, 

pi. 4, figs. 3, 4, 1934 (Miocene, Venezuela).

fConus burckhardti Bose?, Anderson, California Acad. Sci. 

Proc., 4th ser., v. 18, no. 4, p. 110, 1929 (Miocene, Colombia).

fConus burckhardti Bose, Alenchaster-Ibarra, in Masson and 

Alencaster-Ibarra, Asoc. Mexicana, Geb'logos Petroleros 

Bol., v. 3, p. 210, fig. 25, 1951 (Miocene, Mexico). Alencaster-Ibarra, in Rios Macbeth, Idem, vol. 4, p. 347 (list), 

pi. 21, figs. 14, 15, 1952 (Miocene, Mexico).

fConns (Leptoconus) burckhardti Bose, Perrilliant Montoya, 

Mexico Univ. Nac., Inst. Geologia Paleontologia Mexicana, no. 8, p. 26, pi. 3, figs. 20, 21, 1960 (Miocene, 

Mexico).

Conus tortuosopunctatus Toula, Olsson, Bull. Am. Paleontology, 

v. 9, no. 39, p. 54, pi. 3, figs. 6, 11, 1922 (Miocene, Canal 

Zone).

Of medium size, moderately slender, high-spired. 

Shoulder moderately carinate. Outline of spire smoothly conical, except part formed by earliest whorls. Protoconch slender, 2%- to 3-whorled. Early half of 

first post-protoconch whorl sculptured with arcuate 

axial riblets. Carinate and tuberculate shoulder appearing on late half of first whorl. Tubercles gradually 

disappearing on second to fourth whorl. Carinate 

shoulder exposed on spire whorls; generally just exposed on late whorls of mature shells. Anal fascicle 

slightly concave, bearing exaggerated growth threads 

at fairly regular intervals and faint spiral lineation, 

or more distinct, though faint, spiral striae. Body 

whorl sculptured with moderately wide, straplike 

spiral cords, wider on upper part of whorl than on 

lower part, separated by channels, narrower on upper 

part than on lower. Variable number of spiral cords, 

ranging from a few at base to those on lower half 

or more of body whorl, weakly tuberculate. Tubercles 

located on posterior part of wide cords, extending 

practically across narrow cords. Channels between 

cords bearing closely spaced exaggerated axial growth 

threads, producing a punctate effect in narrow channels.

Height (practically complete) 42.3 mm, diameter 

17.7 mm (larger figured specimen). Height (practically complete) 33.7 mm, diameter 14.8 mm (small figured specimen).

Type: Institute de Geologia, Universidad Nacional 

de Mexico.

Type locality: Tuxtepec, Oaxaca, Mexico, Miocene.

Thirty cones from 10 localities in the Gatun formation are identified as Conus burckhardti burckhardti: 

22 from the middle part, four from the upper part in 

the eastern area, and four from the upper part in 

the western area. Though they have the same plan of 

outline and sculpture, they present a considerable 

range of variation, chiefly affecting the sharpness of 

the shoulder carina, the absence or presence of spiral 

striee on the anal fascicle, and the development of 

tubercles on the spiral cords. One shell from the upper 

part of the Gatun in the eastern area and another
358 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

from the upper part in the western area have distinct, 

though faint, spiral sculpture on the anal fasciole. 

The strongest tubercles strong on all the spiral cords 

 is shown by an immature shell (height 23.7 mm) 

found in the upper part in the eastern area. The protoconch and spire whorls are like those of O. imitator 

imitator and O. multUiratus multUiratus.

The type of C. ~burkhardti is moderately slender and 

immature (height 27.3 mm, diameter 10.5). Immature 

Gatun shells and three small shells (largest, height 

18.8 mm, diameter 7.9 mm) in Perrilliat Montoya's 

lot, collected 200 kilometers east of the type locality, 

agree well with it. All the larger specimens in Perrilliat Montoya's collection, however, are less slender and 

their spiral bands, beginning with that below the 

shoulder are tuberculate, as shown in her illustrations. 

If the type represents a cone that develops into this 

stubby tuberculate form, C. ~burckhardti ~burcJchardti 

is not a suitable name for the Gatun fossils. Mature 

shells from the type locality are needed.

Beyond the type region O. burckhardti fiurckfiardti, 

as identified in the present report, has been recognized 

in middle Miocene formations in southeastern Costa 

Rica, Panama, possibly Colombia, and Trinidad, and 

in deposits of late Miocene age in the Bocas del Toro 

area, Panama, and Venezuela. Cossmann's illustration 

of O. marginatus Iboussaci (Cossmann, 1913, p. 46, 

pi. 3, fig. 16) indicates the occurrence of a similar cone 

in the Miocene of Martinique.

C. harveyensis Mansfield (1930, p. 33, pi. 1, fig. 12; 

upper Miocene, Florida) lacks tubercles on the carina 

of early post-protoconch whorls and on the body-whorl 

spiral cords. The type evidently the only specimen 

is small (height 20.9 mm). The type of C. corrugatm 

Gardner (1926-47, p. 360, pi. 43, fig. 9, 1938; lower 

and middle Miocene, Florida; not C. corrugatus Sowerby, 1870) is even smaller (height 15.5 mm), but the 

largest of some 100 topotypes is comparable in size 

to the type of O. harveyensis. It is doubtful whether 

these two species are distinguishable. Though Gardner's name is a junior homonym, a substitute name 

should not be proposed for it without adequate study 

of the available material.

Though C. ~burckhardti burcJchardti is more similar 

to the eastern Pacific species O. tornatus Broderip 

(Hanna, 1963, p. 39, pi. 7, figs. 10-13) than to any 

known to be living in the Caribbean region, the similarity and presumably the relationship is not close. 

The overlap of the late whorls of O. ~burckkardti ~burckhardti is greater and its anal fasciole is flatter.

C. tortuosopunctatus Toula (1911a, p. 507, pi. 31, 

fig. 20) is rejected as a Gatun fossil. The type is a 

chalky shell and the matrix in and near the aperture

consists of quartz grains. Both the type of preservation and the matrix, as well as the species itself, are 

unlike anything known in the Canal Zone.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

localities 144, 147b (identification doubtful), 155,155a, 

155b, 156 (identification doubtful), 157. Upper part, 

eastern area, localities 171, 175; western area,, localities 

182, 182a, 185. Middle Miocene deposits, Tehuantepec 

area, Mexico, southeastern Costa Rica, Trinidad. 

Limon formation (late Miocene), Bocas del Toro area, 

Panama. Punta Gavilan formation (late Miocene), 

Venezuela.

Conns bnrckhardti harrisi Olsson

Plate 57, figure 17

Conus harrisi Olsson, Bull. Am. Paleontology, v. 9, no. 39, p. 53, 

pi. 3, fig. 1, 1922 (Miocene, Costa Rica). Rutsch, Schweizer, Palseont. Gesell. Abh. v. 55, no. 1, p. 104, pi. 9, figs. 

4-6, 1934 (Miocene, Venezuela). Olsson, Neogene mollusks 

from northwestern Ecuador, Paleontological Research 

Inst, p. 88, pi. 11, fig. 6, 1964 (Miocene, Ecuador).

Resembling Conus ~burcJchardti ~burckhardti, but outline more slender. Carinate and tuberculate shoulder 

appearing near end of first post-protoconch whorl 

and spiral cords on upper part of mature body whorl 

wider. Tubercles gradually disappearing on second or 

third whorl. Spiral sculpture on anal fasciole limited 

to faint lineation.

Height 37.7 mm, diameter 13.5 mm (figured specimen).

Type: Paleontological Research Inst. 20899.

Type locality: Rio Banana, Limon Prov., Costa Rica, 

middle Miocene.

This slender subspecies of Conus Iburckhardti is represented by 22 specimens from the lower and middle 

parts of the Gatun formation. An exceptionally slender shell from the upper part in the eastern area is 

doubtfully identified, as the shoulder and the spire 

whorls are missing. The slender outline and greater 

width of the spiral cords on mature body whorls are 

the most diagnostic features distinguishing O. ~burckhardti harrisi from the nominate subspecies. The other 

differences mentioned in the brief description are of 

no significance. In both Costa Rica and Venezuela C. 

~burckhardti harrisi reaches a larger size (height about 

60 mm) than in the Canal Zone. Both subspecies are 

in the collection from locality 155, but that collection 

is not narrowly controlled stratigraphically.

C. alaqucensis Mansfield (1935, p. 18, pi. 1, fig. 7; 

upper Miocene, Florida) is wider at the shoulder and 

sharply carinate.

Occurrence: Lower, middle, and possibly upper parts 

of Gatun formation (middle Miocene). Lower part,
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 359

localities 136, 136a, 138a, 138c, 138d. Middle part, 

eastern area, localities 139c, 139e, 155. Upper part, 

eastern area, locality I77d (identification doubtful). 

Deposits of early (?) Miocene age, Ecuador. Middle 

Miocene deposits, southeastern Costa Rica. Punta Gavilan formation (late Miocene), Venezuela.

Conus cf. C. peruvianus Olsson

A fragmentary body whorl from the Caimito formation of Barro Colorado Island reveals the presence of 

a cone that has a strongly tuberculate shoulder and 

narrow spiral grooves on the lower part of the whorl. 

If this fossil were complete, its height would be about 

18 mm. In general features, so far as it goes, it is 

similar to Conus peruvianus Olsson (1930, p. 40, pi. 5, 

figs. 13-15), a late Eocene Peruvian species.

Occurrence: Caimito formation, Gatun Lake area 

(late Oligocene), locality 54h.

Conns tortuoso&triatus Toula 

Plate 57, figures 5, 6

Conus (Chelyconus} tortuosostriatus Toula, K. k. Geol. Reichsanstalt Jahrb., v. 61, p. 508, pi. 31, fig. 22a, b, 1911 (Miocene, Canal Zone). Olsson, Bull. Am. Paleontology, v. 9, 

no. 39, p. 49, pi. 1, fig. 15, 1922 (Miocene, Panama).

Conus (Hcmiconus) tortuosostriatus Toul'a, Cossmann, Jour. 

Conchy liologie, v. 61, p. 40, pi. 3, figs. 28, 29,1913 (Miocene, 

Canal Zone).

Not Conus tortuosostriatus Toula, Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 41, pi. 6, fig. 9, 1917 (Miocene, Dominican Republic) ;= C. gabbi Pilsbry and Johnson.

Conus tortuosopunctatus Toula, Cossmann, Jour. Conchy liologie, 

v. 61, p. 47 (discussion), pi. 3, figs. 22, 23, 1913 (Miocene, 

Canal Zone).

Conus granozonatus Guppy, Brown and Pilsbry, Acad. Nat. Sci. 

Phila. Proc., v. 63, p. 341, 1911 (Miocene, Canal Zone).

Conus consobrinus Sowerby, Brown and Pilsbry, Idem (Miocene, Canal Zone).

Of medium size, slender to inflated, shoulder angulated and tuberculate. Spire moderately high, its 

profile conical, except part formed by earliest whorls. 

Protoconch not completely preserved, consisting of 

about two slender whorls. Early half of first postprotoconch whorl sculptured with relatively coarse 

axial riblets. Angulated and coarsely tuberculate shoulder appearing on late half of first whorl. Except on 

first few whorls, tubercles flattened parallel to shoulder 

and whorls overlapping to edge of shoulder. Tubercles 

continuing, in reduced form, on shoulder of body 

whorl. Anal fascicle slightly concave, bearing exaggerated growth threads and more or less distinct spiral 

sculpture. Body whorl sculptured with narrow, relatively widely spaced, flattened spiral cords. Weak 

tubercles present or absent on cords. Channels between

cords bearing exaggerated axial growth threads. Some 

channels, especially near top of whorl, on some shells 

bearing a secondary spiral thread.

Height (incomplete) 28.2 mm, diameter 15.7 mm 

(inflated figured specimen). Height (not quite complete) 26.3 mm, diameter 11.7 mm (slender figured 

specimen).

Type: Tech. Hochschule, Vienna.

Type locality: Presumably Gatun Locks excavation, 

Canal Zone, middle part of Gatun formation.

Ten specimens of this species are in the collections 

now being studied. They were found in the upper 

part of the Gatun formation in the eastern area and 

in the Chagres sandstone. One presumably from Mount 

Hope, like other species presumed to have been collected at that locality by "Bland?" including the type 

of PJios metuloides (p. 267), is mislabelled "Ponton, 

Dominican Eepublic." These few specimens show the 

presence or absence of secondary spiral threads and 

of weak tubercles below the shoulder. The type has a 

height of 30.4 mm, and a diameter of 14 mm.

Seven names have been proposed for Miocene Central American, West Indian, and northern South 

American cones that are more or less similar to Conus 

mcgintyi Pilsbry (1955), a fairly deep-water species 

dredged off the coast of Florida and Cuba at depths of 

70 to 240 fathoms. The name C. mazei Deshayes has 

generally been used for that species (Clench, 1942, 

p. 17, pi. 9, figs. 1, 2). The slender, high-spired species 

C. larvatus Pilsbry and Johnson (Pilsbry, 1922, p. 332, 

pi. 21, fig. 10; Miocene, Dominican Eepublic; not C. 

larvatus Gmelin, 1791) is more similar to C. mcgintyi 

than any other of these fossil forms. Perhaps some of 

the Miocene species other than C. "larvatus" have been 

too narrowly defined. Nevertheless, the late whorls of 

C. tortuosostriatus overlap to the edge of the shoulder 

of the preceding whorl and the spacing of the spiral 

cords is relatively wide. These features distinguish it 

from C. gracilissimus Guppy (Woodring, 1928, p. 216, 

pi. 12, figs. 3, 4; Miocene, Jamaica), C. gabbi Pilsbry 

and Johnson (Pilsbry, 1922, p. 332, pi. 21, figs. 8, 9; 

Miocene, Dominican Republic), and G. stibarus Woodring (1928, p. 217, pi. 12, fig. 5, Miocene, Jamaica). 

In addition, C. stibarus has fewer and coarser shoulder 

tubercles.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

Gatun Locks site (Toula's and Brown and Pilsbry's 

records). Upper part, eastern area, localities 175, 176, 

177, 177b. Chagres sandstone (late Miocene), locality 

208. Limon formation (late Miocene), Bocas del Toro 

area, Panama.

368-278 O - 70 - 5
360 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Family TTJRRIDAE

Powell's (1966) recent monograph is indispensable 

in dealing with this highly diversified family. His 

subfamily classification is adopted in the present 

account.

The reaction against the early usage of a few generic names to cover the entire family has gone so far 

that turrid generic names are almost as narrowly 

restricted as ammonite generic names. As a minor 

contribution toward bringing the family more into 

line with other gastropod families, six names proposed 

at the generic level (Woodring, 1928, p. 145-201) are 

now reduced to subgeneric rank: Polystira, Fusiturricula, Oryoturris, Saccharoturris, Miraclathurella, and 

Lioglyphostoma. Perhaps other genera named in 1928 

may be downgraded, but they are not involved in the 

present report.

Turrids have been found in all the fossiliferous formations, except the Gatuncillo formation and the Cucaracha formation, the latter of which for the most part 

is nonmarine. As shown in the following table, however, 50 of the 70 species and subspecies, including all 

the mangilenes and daphnellines, occur in the Gatun 

formation.

Species and subspecies of turrid gastropods

Subfamily

Turrinae _ _

Clavinae

Mangeliinae _ _ _

Daphnellinae__ __

Eocene

or Oligocene

§

0 03 

(H g

rt S

p«2

§^-N

S^

03 W

§

1

1

Oligocene

a

 *g

g

o

2

o

pq

1

1

a

.0

"o3

g

O

o

.t3

R

03

O

?

3

7

Miocene

a

_o

"o3

0

o

03

ft

S

O

1

1

d

.2

03

Q

O

C3

§

m03

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7,

4

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 8

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18

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50

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CO "S

s

n

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4

7

Three of the species listed one from the Culebra 

formation and two from the La Boca formation are 

merely mentioned. In addition to those listed, all the 

formations, except the Gatun and Chagres, contain 

unidentifiable turrid material.

Subfamily TURRIWAE 

Genus Gemmula Weinkauff

Weinkauff, Deuitschen Malakozoologischen Gesell. Jahrbiicher, 

v. 2, p. 287, 1875.

Type (logotype, Cossmann, Essais paleoconchologie comparee, pt. 2, p. 62, 1896) : Pleurotoma, gemmata Hinds \P. 

gemmata Reeve, 1843, not P. gemmata Conrad, 1835,rr 

Gemmula hindsiana Berry], living, Baja California and 

Gulf of California.

The type species has been illustrated by Harris (1937, 

p. 10, pi. 1, figs. 33, 33a) and Keen (1958a, p. 250, 

pi. 30, figs. 1-4).

Gemmula, a genus dating back to Paleocene time, is 

considered by Powell (1964, p. 243) to be the main 

turrine stem. Its distribution has been greatly restricted 

since early Tertiary time. Though it is chiefly a westtern Pacific genus at the present time, the type species 

survives in the eastern Pacific Ocean and G. periscelida (Dall) (1889, p. 74, pi. 32, fig. 2) in the western 

Atlantic.

Three fragments of a few small whorls from the 

Culebra formation are listed as Gemmula sp., and 

molds of a moderately large species from the La Boca 

formation are identified as O. cf. O. vaningeni.

Gemmula cf. G. arnica Casey

Small, slender. Protoconch missing. Peripheral band 

coarsely gemmate. Subsutural primary spiral thread 

fairly strong, except on body whorl near aperture. 

Three widely spaced primary spiral threads on main 

part of body whorl below peripheral band. Fine secondary spiral threads in space between peripheral band 

and subsutural primary thread. Pillar broken, but 

aperture apparently short.

Height (incomplete) 16.2 mm (estimated complete 

height 23 mm), diameter 8.8 mm (largest specimen).

A small, slender species of Gemmula is represented 

in the shallow-water facies of the Caimito formation 

of the Gatun Lake area by four specimens. All are 

incomplete and their shell material is replaced by 

calcite. This species is larger and more strongly sculptured than G. arnica (Casey, 1903, p. 270; Harris, 1937, 

p. 13, pi. 1, figs. 26-28), of the early Oligocene Eed 

Bluff clay of Mississippi.

Harris and Palmer (1946-47, p. 420, 1947) assigned 

G. arnica to Ooronia (Gregorio, 1890, p. 23; type 

(logotype, Cossmann, 1896, p. 78) : Pleurotoma acutirostra Conrad, Eocene, Alabama), which they treated 

as a subgenus of Cremmula. Pleurotoma acutirostra was 

identified by Harris (1937, p. 12, pi. 1, fig. 22) as a 

form of Pleurotoma cMldreni Lea. Though G. arnica, 

like other Oligocene and Eocene species of Cremmula, 

is smaller than the late Tertiary and living species and 

has a shorter aperture, its strong and strongly gemmate 

peripheral band favors assignment to Cremmula.

Occurrence: Caimito formation, Gatun Lake area 

(late Oligocene), locality 56.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 361

Gemmula vaningeni (Brown and Pilsbry) 

Plate 57, figures 11, 18, 22; plate 65, figure 1

Pleurotoma (Gemmula) vaningeni Brown and Pilsbry, Acad. 

Nat. Sci. Phila. Proc., v. 64, p. 505, pi. 22, fig. 4, 1913 

(Miocene, Canal Zone). Cossmann, Jour. Conchyliologie, 

v. 61, p. 19, pi. 2, figs. 21, 22, 1913 (Miocene, Canal Zone). 

Mansfield, Florida Dept. Conservation, Geol. Bull. 12, 

p. 10, 11 (lists), 1935 (Miocene, Florida).

Turris (Gemmula) vaningeni (Brown and Pilsbry), Oinomikado, Geol. Soc. Japan Jour., v. 46, p. 624, pi. 29, fig. 14, 

1939 (Miocene, Colombia). Marks, Bull. Am. Paleontology, v. 33, no. 139, p. 126, pi. 8, fig. 11, 1951 (Miocene, 

Ecuador).

Gemmula vaningeni (Brown and Pilsbry), Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst, p. 93, pi. 20, fig. 12,1964 (Miocene, Ecuador).

Hemipleurotoma eileta Gardner, U.S. Geol. Survey Prof. Paper 

142, p. 290, pi. 38, figs. 27, 28, 1938 (Miocene, Florida).

Moderately large, high-spired, slender. Protoconch 

relatively slender, 4-whorled. First whorl emerging so 

abruptly that apex appears to be broken. First whorl 

to first li/£ whorls smooth, remainder sculptured with 

axial riblets of gradually increasing strength. A faint 

spiral thread visible at lower edge of sculptured protoconch whorls on some specimens, especially on last 

whorl or two. End of protoconch marked by suppression of axial riblets and appearance of spiral sculpture. 

Sculpture dominated by coarsely gemmate peripheral 

band, both edges slightly raised. Subsutural primary 

spiral forming a wide-based, narrow-crested collar. 

Intermediate and late spire whorls bearing a primary 

spiral thread between peripheral band and succeeding 

whorl. Similar threads on body whorl. Secondary spiral 

threads of variable strength, generally strongest between peripheral band and subsutural collar; present 

or absent on flanks of collar. Fine exaggerated growth 

threads conspicuous, especially between peripheral 

band and subsutural collar. Anal sinus deep, apex 

U-shaped. Aperture long.

Height (not quite complete) 38.8 mm, diameter (incomplete) 12.3 mm (largest figured specimen).

Type: Acad. Nat. Sci. Phila. 3855.

Type locality: Gatun Locks excavation, middle part 

of Gatun formation.

Though Gemmula is rare in the Miocene deposits of 

the Caribbean region, G. vaningeni is represented in 

the Gatun formation by 65 specimens. Only one, however, was found in the lower part of the formation 

and none in the upper part in the western area. Locality 175, in the upper part in the eastern area, 

yielded 27, whereas other collections contain nine or 

less. The type (height 19.5 mm) is immature. Its 

siphonal canal and outer lip, as on most of the specimens, are broken. The largest shell would have, an

estimated height of 45 mm, if it were complete. 

Twenty-six of these fossils show the protoconch, but 

on many of them at least some of the protoconch 

whorls are somewhat worn.

Two immature shells are labeled, in Dall's writing, 

types of Pleurotoma pontonensis Dall and are alleged 

to have been collected by "Bland?" at Ponton, in the 

Dominican Kepublic. This record, like that for the type 

lot of Phos metuloides (p. 267), is presumed to be an 

erroneous record for the present Mount Hope. If the 

rejected record is ignored, Gemmula is unknown in the 

Dominican Republic. Pleurotoma pontonensis, mentioned by Mansfield (1925, p. 15) and Gardner (1926- 

47, p. 290, 1938), is a nude name, as recognized by 

Mansfield.

The features of G. vaningeni are basically uniform, 

although the strength of the secondary spiral threads 

is somewhat variable. Three of the 27 specimens from 

locality 175 are malformed. The malformation takes the 

form of a swelling on the body whorl at and near the 

edge of the outer lip between the anal sinus and the 

siphonal canal. In apertural view the effect is an insinuation of the lip, but the growth lines are not 

insinuated. With further growth the swelling forms 

an almost closed tube, which is later sealed off and 

followed by another swelling (pi. 57, figs. 11, 18). The 

anal sinus of the smallest of these shells is partly covered by the attached valve of an anomid (?). The attachment, however, may have taken place after death 

of the gastropod. At all events the anal sinus of the 

other two shells shows no indication of blocking. The 

same kind of malformation is shown by other turrids. 

Those that have come to my attention are as follows: 

species in the genera Gemmula (Glibert, 1954, p. 7, 

pi. 2, figs. 3a, b; MacNeil, 1960, p. 104; Powell, 1964, 

p. 253); Pinguigemmula (MacNeil, 1960, p. 103, 104; 

Powell, 1964, p. 277, 279, pi. 215, figs. 5, 6); Ptychosyrinx (Powell, 1964, p. 289, 290, 291, pi. 223, fig. 1, 

pi. 224; Okutani, 1964, p. 425, pi. 4, fig. 9); Aforia 

(Dall, 1918, p. 319); and a species assigned to Drillia 

(Melvill, 1917, p. 150). Though various explanations 

for the malformation have been proposed, its significance, as pointed out by Powell (1966, p. 8), will not 

be known until the animal is found in such a shell. 

A. A. Olsson called my attention to the remarks concerning Aforia by Dall, who later (Dall, 1921, pi. 11, 

fig. 6) illustrated a specimen of Aforia showing the 

"anterior sulcus."

Hemipleurotoma eileta, of the Shoal River formation of Florida, is considered to be a synonym of G. 

vaningeni. The largest Florida fossils are a little 

smaller than Gatun shells and some, including the 

type, have a slightly narrower and slightly less coarse-
362 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

ly gemmate peripheral band. Incomplete shells indicate a maximum height of about 35 mm, although the 

type, which is not quite complete, has a height 24.3 mm. 

Mansfield listed the Florida form as G. vaningeni. The 

largest of his three specimens from USGS locality 

12044 closely resembles Gatun fossils of intermediate 

size.

Small shells (maximum estimated height 17 mm) 

from the early middle Miocene part of the Brasso formation of Trinidad (Globorotalia fohsi zone, USGS 

21234, 21230) appear to represent immature shells of 

G. vaningeni or a small form of that species. Two 

other small shells from the same part of the Brasso 

formation (USGS 9219) were identified by Mansfield 

(1925, p. 15) as Turris vaningeni var. machapoorensis 

(Maury). Their relatively low peripheral band and 

subsutural collar suggest Maury's (1925, p. 191, pi. 32, 

figs. 1, 14) [unnumbered on plate] Drillia vaningeni 

var. sanctiandrece. The status of Mansfield's and 

Maury's specimens is uncertain.

Illustrations of G. hindsiana, the type of the genus, 

suggest that it is smaller and more slender than G. 

vaningeni, and has a narrower and lower sutural collar. 

Nevertheless G. vaningeni is much more closely related 

to G. hindsiana than to G. periscelida, a deep-water 

species and the only species surviving in western Atlantic waters. The type locality of that species is off 

the Gulf of Morrosquillo, on the Caribbean coast of 

Colombia. Ball's record, "near Monosquillo," was based 

on a cataloger's error in transcription.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, locality 138c. Middle part, eastern area, localities 147b, 

147g, 147h, 151, 155, 155a, 155b, 159d. Upper part, 

eastern area, localities I73a, 175, 176, 177, I77c, 178. 

Middle Miocene deposits, Darien area (USGS 8429, 

8452, 8477), Panama. Subibaja formation (early Miocene) and Picaderos formation (middle Miocene), 

Ecuador. Middle Miocene deposits, southwestern Columbia. Early middle Miocene part of Brasso formation, Trinidad. Shoal River formation (middle Miocene), Florida.

Gemmula machapoorensis (Maury)

Drillia vaningeni var. machapoorensis Maury, Bull. Am. Paleontology, v. 10, no. 42, p. 191, pi. 32, figs. 5, 9, 1925 

(Miocene, Trinidad).

Turris brasscensis Mansfield, U.S. Natl. Mus. Proc., v. 66, art. 

22, p. 14, pi. 2, figs. 7, 8, 1925 (Miocene, Trinidad).

Hemipleurotoma Utropis Gardner, U.S. Geol. Survey Prof. 

Paper 142, p. 290, pi. 38, fig. 29, 1938 (Miocene, Fla.).

Protoconch and first two post-protoconch whorls 

available. Protoconch large, 5-whorled, rapidly enlarging. First whorl emerging abruptly from apex. At

least last three whorls sculptured with narrow, closely 

spaced axial riblets. Lower half of these whorls also 

sculptured with still narrower spiral threads. A stronger sutural thread on last whorl. Last few axial riblets 

coarser and more widely spaced than others, followed 

by abrupt suppression of riblets and appearance of 

strong spiral sculpture. Both edges of peripheral band 

of post-protoconch whorls raised to form distinct spiral 

threads. Subsutural primary spiral forming a strong, 

wide-based, narrow-crested collar. Exaggerated, uniformly spaced growth threads strong on flanks of subsutural collar.

Height (almost complete) 6.4 mm, diameter 3 mm.

Lectotype (herewith designated) : Paleontological 

Research Inst. 1006.

Type locality: Thirteenth mile post on Guaico-Tamana Road, Trinidad, "Machapoorie Miocene" [Brasso 

formation].

The only specimen, from the lower part of the 

Gatun formation, is immature. Though the protoconch 

whorls are partly corroded, the features mentioned in 

the description are discernible. Despite the unpromising material, the identification of this distinctive 

species is made with considerable confidence on the 

combined basis of the protoconch and the post-protoconch sculpture. The protoconch is large, rapidly enlarging, and bears both axial and spiral sculpture. The 

spiral thread at both edges of the peripheral band 

and the strong, wide-based subsutural collar are diagnostic features of the post-protoconch sculpture. With 

further growth, as shown by shells from Trinidad and 

Florida, a secondary spiral thread appears on both 

flanks of the subsutural collar and exaggerated growth 

threads on parts of the shell other than the flanks of 

the subsutural collar. Also as shown by growth lines 

on shells from Trinidad and Florida, the anal sinus is 

typically that of Gemmula.

G. machapoorensis evidently is a small species. 

Maury estimated 19 mm for the maximum height, 

which is about the same as that for Florida shells. The 

largest Trinidad shell in the USNM collections has a 

height of 12.2 mm. The protoconch is not preserved 

on any of the nine specimens from the Shoal River 

formation of Florida. It is preserved, however, on all 

of the 18 specimens in the USNM Trinidad collections 

and, though somewhat worn, on the lectotype. The 

number of protoconch whorls is six or seven, and 

sculpture appears on the first or second. The Globorotalia fohsi zone of the Brasso formation yielded all 

this Trinidad material (USGS localities 8302, 9212, 

19856, 19870, 21234). According to H. G. Kugler, who 

supplied the zonal data for these localities, and also 

for 21230 cited under G. vaningeni, the areal geology
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 363

shows that even the float collections (8302, 9212) were 

derived from that zone.

Mansfield commented on the similarity of Trinidad 

and Florida specimens. It seems remarkable that G. 

vaningeni and G. machapoorensis are associated at such 

widely separated localities as Trinidad, Panama, and 

Florida.

Occurrence: Lower part of Gatun formation (middle Miocene), locality 138c. Early middle Miocene part 

of Brasso formation, Trinidad. Shoal Eiver formation 

(middle Miocene), Florida.

Genus?

"Gemmula" species 

Plate 48, figures 19, 20

Small, moderately slender, biconic. Protoconch missing. Peripheral band wide, coarsely gemmate. Suture 

bordered by a gemmate collar, less coarsely gemmate 

than peripheral band. Other spiral sculpture consisting of narrow primary and secondary threads, faintly 

noded by conspicuous, exaggerated growth threads. 

Aperture long, narrow. Siphonal canal virtually absent; aperture ending in a shallow siphonal notch. 

Siphonal fasciole slightly inflated. Anal sinus moderately deep, apex U-shaped, located on peripheral band.

Height (almost complete) 20.7 mm, diameter 18.2 

mm (figured specimen).

This unnamed species is represented by a defective 

specimen and a minute tip from the moderately deepwater facies of the Caimito formation on Barro Colorado Island. It is to be assigned to an apparently 

unnamed turrine genus that has a Gemmula-\ike anal 

sinus and gemmate peripheral band, but, unlike Gemmula, has a long, narrow aperture, siphonal notch, 

and siphonal fasciole. Though the anterior end of the 

aperture is defective, growth lines show the shallow 

siphonal notch. The gemmate sutural collar is a noteworthy feature. The outline suggests Scobinella or a 

small slender Bathytoma.

Occurrence: Caimito formation, Gatun Lake area 

(late Oligocene), localities 54g, 54j.

Genus Pleuroliria Gregorio

Gregorio, Annales Geol. Paleontologie, pt. 7, p. 38, 1890.

Type (orthotype): Pleurotoma supramirifica Gregorio, locality 

not specified (presumed to be Pleurotoma cochlearis Conrad, Oligocene, Mississippi).

Should Gregorio's type designation be considered 

equivocal, Cossmann unequivocally cited Pleurotoma 

supramirifica as the type four years later (Cossmann, 

1893 (1894), p. 43).

Harris (1937, p. 8, pi. 1, figs. 5, 5a, b) reproduced 

Gregorio's illustrations of Pleurotoma supramirifica.

There is no reasonable doubt that it is the Oligocene 

species Pleurotoma cochlearis. The known Eocene 

species are much smaller.

Submenus Polystira Woodring

Woodring, Carnegie Inst. Washington Pub. 385, p. 145, 1928. 

Type (orthotype): Pleurotoma albiaa Perry, living, southern 

Florida, Gulf of Mexico, and West Indies.

Bartsch (1934, p. 8) claimed that Pleurotoma albida 

is a western Pacific species and that Murex virgo 

Wood is the proper name for the type species of 

Polystira. Despite Perry's statement that his species 

is "from the South Seas, being frequently found at 

New Zealand and Lord Howe's Island" (Perry, 1811, 

explanation of pi. 32, fig. 4), his illustration is considered to be a reasonably satisfactory representation 

of the western Atlantic species later named Murex 

virgo. Pleurotoma albida is not recognized in the western Pacific Ocean (Powell, 1964, p. 315).

Though Polystira was named at the generic level, 

it is proposed to treat it as a subgenus of Pleuroliria. 

It is interpreted to be the direct descendant of Pleuroliria and the two groups have similar basic features. 

Polystira reaches a larger size than Pleuroliria s.s. and 

has a less rapidly enlarging protoconch. The protoconch of Polystira generally has fewer whorls and 

fewer whorls sculptured with axial riblets. The number of protoconch whorls and of sculptured protoconch whorls, however, overlap in the two groups and 

not all of the species of Polystira are large. Some 

species of Polystira, including the type species, have 

a blunt protoconch, but that of others is acute. In 

other words, too much reliance was placed on type 

species when Polystira was named a procedure that 

has been justly criticized (MacNeil, 1960, p. 100).

Both Pleuroliria s.s. and Polystira embrace American species. Pleuroliria s.s. is of Eocene and Oligocene age. The age range of Polystira is early Miocene 

to the present time and the living species occur in 

western Atlantic and eastern Pacific waters. They are 

characterized by strong, coarse, exaggerated growth 

threads, at least on early post-protoconch whorls, if 

not also on later whorls. This may seem to be a minor 

feature, but it can be traced back through the fossil 

species of Polystira to the Oligocene species that 

doubtless is the type species of Pleuroliria.

Pleuroliria albidoides (Gardner) (1926-47, p. 287, 

pi. 38, fig. 24, 1938; Chipola formation, Florida) is 

the earliest species of Polystira that is known to have 

a blunt protoconch. That species was illustrated by 

Ball (1915, p. 38, pi. 5, fig. 13) as Turns albida without any indication that the specimen is a Chipola 

fossil, not a Tampa fossil. The specimen he illustrated
364 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

(height 49 mm) would serve as a better namebearer 

than the minute shell (height 10.6 mm) chosen as the 

type because it shows the protoconch. One of the 

species of Polystira that has an acute protoconch 

Pleuroliria tenagos, the next species described was 

designated as the type of Josephina (Gardner, 1945, 

p. 246), proposed as a section of Pleuroliria. That 

proposal followed my suggestion (Woodring, 1928, 

p. 145), which overemphasized the distinction between blunt and acute protoconchs. The protoconch 

illustrated by Gardner (1926-47, p. 288, pi. 38, fig. 23, 

1938) as the protoconch of Polystira virgo is that of a 

small shell (height 11.6 mm) dredged off Grenada at 

a depth of 73 fathoms. It presumably is one of the 

relatively small living West Indian species, for which 

six names have been proposed.

LopJiiotoma (Casey, 1904, p. 130; type (logotype, 

Woodring, 1928, p. 146) : Pleurotoma tigrina Lamarck 

(= P. acuta Perry), living, western Pacific Ocean) is 

the western Pacific analog of Polystira. As pointed 

out by Powell (1942, p. 49, figs. E2, E3; 1964, p. 303, 

pi. 173), Polystira may be distinguished by its shallower, more V-shaped anal sinus. If, however, so much 

emphasis is to be placed on the anal sinus, the western 

Atlantic species described by Dall as Pleurotoma 

albida var. tellea (Dall, 1889, p. 73) is not a typical 

species of Polystira. The sinus of mature shells of that 

species (but not of immature shells) is decidedly asymmetric and shallow. This unusual species unusual in 

mature sculpture, as well as in mature anal sinus 

has been illustrated recently (Abbott, 1954, p. 268, pi. 

13, fig. m), but has not yet been adequately described. 

It has been dredged at depths of 35 to 220 fathoms off 

southern Florida and in the eastern Gulf of Mexico.

Pleuroliria (Polystira) tenagos (Gardner) 

Plate 57, figures 12, 23, 24; plate 65, figure 2

Pleurotoma albida Perry, Brown and Pilsbry, Acad. Nat Sci.

Phila. Proc., v. 63, p. 343, 1911 (Miocene, Canal Zone). 

Turns albida (Perry), Olsson, Bull. Am. Paleontology, v. 9,

no. 39, p. 58, pi. 4, figs. 1, 2, 1922 (Miocene, Panama). 

Pleurotoma haitcnsis Sowerby, Cossmann, Jour. Conchyliologie,

v. 61, pi. 16, pi. 2, figs. 1-4, 1913 (Miocene, Canal Zone). 

Polystira (Pleuroliria) tenagos Gardner, U.S. Geol. Survey

Prof. Paper 142, p. 288, pi. 38, figs. 25, 26, 1938 (Miocene,

Fla.). 

f Pleuroliria tenagos (Gardner), Gardner, Geol. Soc. America

Mem. 11, p. 246, 1945 (Miocene, Mexico).

Of medium size to moderately large, moderately 

slender to moderately inflated. Profile of late whorls 

slightly to decidedly modified by peripheral carina. 

Protoconch acute, 3- to 4-whorled (generally 3 to 3i/£). 

Last whorl, or a little less, to last two (generally last 

to last 11/4) bearing axial riblets. Spiral sculpture of

post-protoconch whorls dominated by peripheral 

carina of variable strength; both edges slightly raised. 

Subsutural primary spiral thread strong, single or 

double. On intermediate and late spire whorls one or 

two fairly strong primary spiral threads between 

peripheral carina and succeeding whorl. Secondary 

spiral threads on intermediate and late whorls, except 

on pillar. Exaggerated, uniformly spaced growth 

threads conspicuous on unworn early whorls, more 

subdued on intermediate and late whorls, except those 

representing stages of arrested growth at irregular 

intervals. Anal sinus moderately shallow, V-shaped, 

located on peripheral carina. Interior of outer lip lirate.

Height 58.8 mm, diameter 16.6 mm (largest figured 

specimen). Estimated restored height 75 mm, diameter 

20.8 mm (largest specimen).

Type: USNM 351134.

Type locality: USGS 3742, Shell Bluff, Shoal Kiver, 

Walton County, Fla., Shoal Eiver formation.

Pleuroliria tenagos is widespread in the three parts 

of the Gatun formation. It is locally abundant in the 

lower and middle parts, as shown by 37 specimens in a 

lower Gatun collection (locality 138c) and 24 in a 

middle Gatun collection (locality 161a). The largest 

specimens occur in the upper part of the formation 

in both eastern and western areas. Though they are 

incomplete, they indicate a height of about 75 mm. 

On his plate 4, figures 1, 2, Cossmann illustrated an 

incomplete large specimen from the upper part in 

the eastern area.

The degree of inflation, the whorl profile, and details of sculpture, especially the strength of the subsutural primary spiral thread, are variable. The strongest modification of whorl profile, due to angulation by 

the peripheral carina, is shown, but not uniformly 

shown, by shells from the lower part of the Gatun 

(pi. 57, fig. 12). A slightly modified profile is illustrated by the upper Gatun specimen of plate 57, figure 24; and an intermediate profile by the middle 

Gatun specimen of plate 57, figure 23. The fossils of 

the three parts, however, do not uniformly show a 

graded profile series. Ten of the 14 smallest shells 

(maximum height 22 mm) in the largest lower Gatun 

collection are exceptionally slender and have a low 

peripheral carina. They presumably would have a 

stronger carina at a later growth stage.

Twenty-four of the some 230 available specimens 

show the acute protoconch, but none of the six specimens from the upper part of the Gatun in the western 

area has a preserved protoconch. A minute middle 

Gatun shell (height 5.2 mm) the only shell from locality 161c has a blunt protoconch of two whorls, the 

last half whorl of which bears axial riblets. This fossil
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 365

is listed as Pleuroliria sp. It may eventually be found 

to be an aberrant specimen of P. tenagos, but that seems 

to be unlikely. At all events, its status is uncertain 

until mature features are known.

In Florida P. tenagos is not so variable. The Florida 

form is most like moderately slender Gatun fossils of 

intermediate whorl profile. The type is small (height 

37.5 mm), but fragments suggest a height comparable 

to that of the largest lower and middle Gatun specimens.

P. tenagos is not similar to species of the same age 

in the Dominican Republic and Jamaica: P. haitensis 

(Sowerby) (Pflug, 1961, p. 69, pi. 20, figs. 2, 3, 5-13) 

and P. barretti (Guppy) (Woodring, 1928, p. 146, 

pi. 4, figs. 6, 7; Pflug, 1961, p. 70, pi. 20, figs. 1, 4), 

respectively. The protoconch of both of those species 

is blunt. The low peripheral carina of P. haitensis has 

no effect, or a slight effect, on whorl profile. P. ~barretti, a close relative of the living West Indian P. 

aTbida, is larger than P. haitensis and P. tenagos. Its 

peripheral carina strongly affects whorl profile. The 

contrast between P. haitensis and P. barretti is well 

shown by Pflug's illustrations on the same plate. P. 

andersoni Olsson (1964, p. 90, pi. 14, figs. 3-3b), a 

large early Miocene Columbian species, has exceptionally strong secondary spiral sculpture. Its protoconch 

is unknown.

P. tenagos occurs in the middle Miocene deposits of 

the Darien area (USGS 8428, 8429, 8430, 8477, 8479) 

and in the Limon formation, of late Miocene age, in 

the Bocas del Toro area. Its distribution may be found 

to be more extensive. The middle Miocene Venezuelan 

fossils identified as P. barretti (Jung, 1965, p. 561, 

pi. 76, figs. 5, 16), for example, should be compared 

with P. tenagos. Their protoconch, however, is not 

known.

The moderate or moderately large size, acute protoconch, and relatively narrow primary spiral sculpture 

show that P. tenagos is not closely related to P. albida. 

The relationship to the living eastern Pacific species 

P. oxytropis (Sowerby) and P. picta (Reeve) (Olsson, 

1964, p. 90, pi. 14, figs. 1, la, Ib; p. 90, pi. 14, figs. 2, 2a, 

respectively), especially P. picta, is closer. That species 

is slender, and shows a moderately modified whorl 

profile, and weak subsutural primary spiral thread.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 136, 138, 138a, 138c, 138d, 138e, 138f. Middle 

part, eastern area, localities 139c, 139d, 139e, 139f, 142, 

146, 147b, 147g, 147h, 151, 153a (identification doubtful), 154,155, 155a, 155b, 155c, 156, 157, 159d; western 

area, localities 161a, 161b, 161c (Pleuroliria sp.). Upper part, eastern area, localities 172, 173, 175, 176,

I76a, 177c; western area, localities 182, 182a, 183, 185. 

Guajalote formation (late early Miocene), northeastern 

Mexico (identification doubtful). Shoal River formation (middle Miocene), Florida. Middle Miocene deposits, Darien area, Panama. Limon formation (late 

Miocene), Bocas del Toro area, Panama.

Pleuroliria (Polystira) ecuadoriana (Olsson) 

Plate 62, figure 26

Polystira oxytropis ecuadoriana Olsson, Neogene mollusks from 

northwestern Ecuador, Paleontological Research Inst, 

p. 91, pi. 14, figs. 5, 5a, 1964 (Miocene, Ecuador).

Moderately large, rapidly enlarging in diameter. 

Profile of all except early post-protoconch whorls 

strongly modified by peripheral carina. Protoconch 

missing. Space between peripheral carina and preceding whorl wide and concave. Subsutural primary spiral 

thread of intermediate whorls low, reduced practically 

to vanishing point on last preserved whorl. On spire 

whorls primary spiral thread between peripheral 

carina and succeeding whorl moderately strong. Secondary spiral threads of variable strength. Exaggerated growth threads strong on early and intermediate 

whorls. Anal sinus, as shown by growth lines, forming 

an open V on peripheral carina. Outer lip broken back.

Height (incomplete) 39.5 mm, diameter (incomplete) 22.5 mm (figured specimen).

Type: USNM 643960.

Type locality: Punta Gorda, Esmeraldas Province, 

Ecuador, Esmeraldas formation.

The Chagres sandstone yielded the illustrated incomplete, worn shell, which indicates a height of about 

TO mm, and also a fragment of a few worn early postprotoconch whorls. This recently described species is 

noteworthy for its rapidly enlarging outline, strong 

peripheral carina, and weak subsutural primary spiral 

on the last whorl or last few whorls. Six specimens in 

a collection from the Valiente Peninsula, in the Bocas 

del Toro area of northwestern Panama (USGS 8322), 

all more or less incomplete, show the same features. 

The fossiliferous strata in the Valiente Peninsula are 

considered to be of late Miocene age.

Though Pleuroliria ecuadoriana was named as a 

subspecies of the living Panamic species P. oxytropis 

(Sowerby), the large size and rapidly enlarging outline justify specific rank. It is doubtful whether the 

slender specimen illustrated by Olsson (1964, pi. 14, 

fig. 5b) is P. ecuadoriana.

Occurrence: Chagres sandstone (late Miocene), locality 208. Limon formation (late Miocene), Bocas del 

Toro area, Panama. Esmeraldas formation (late Miocene), Ecuador.
366 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Subfamily TTTRRICTJXIH'AE 

Genus Zemacies Finlay?

Finlay, New Zealand Inst. Trans., v. 56, p. 252, 1926. 

Type (orthotype): Zemacies elatior Finlay, Miocene, New 

Zealand.

The following two early Tertiary turriculine species 

are noteworthy for their slender outline, long narrow 

aperture, and flat, closely spaced, narrow spiral bands. 

They represent Zemacies or a related genus. Though 

both evidently are new species, the material is too 

imperfect to name. In New Zealand Zemacies has an 

age range of Paleocene to Pliocene (Powell, 1966, 

p. 36).

Zemacies? species a

Plate 48, figure 13

Of medium size, slender, subangulated at s.noulder. 

Protoconch corroded, apparently consisting of about 

three rapidly enlarging whorls. Shoulder bearing low, 

axially elongate swellings, progressively more subdued 

on late whorls. Spiral sculpture consisting of flat, 

closely spaced, narrow bands. Subsutural band narrow, an adjoining band wider. Anal sinus apparently 

on narrow, concave space between shoulder and suture. Aperture long, narrow.

Height (incomplete) 19.3 mm (estimated restored 

height 27 mm), diameter (incomplete) 8.5 mm (estimated restored diameter 9 mm).

The marine member of the Bohio (?) formation at 

Vamos Vamos yielded two imperfect specimens of 

this species.

Occurrence: Marine member of Bohio (?) formation 

(late Eocene or early Oligocene), locality 40d.

Zemacies? species b 

Plate 48, figure 21

Of medium size, slender, late whorls subangulated 

at shoulder. Protoconch and early post-protoconch 

whorls missing. Late whorls bearing widely spaced, 

obliquely elongate nodes on shoulder. Spiral sculpture 

consisting of flat, narrow, closely spaced bands. Subsutural band relatively wide. Anal sinus located 011 

narrow, concave space between shoulder and suture. 

Aperture long, narrow.

Height (incomplete) 32 mm (estimated restored 

height 38 mm), diameter (incomplete) 11.6 mm (estimated restored diameter 13 mm).

This strongly noded species is represented by an incomplete specimen from the moderately deep-water 

facies of the Caimito formation on Barro Colorado 

Island.

Occurrence: Caimito formation, Gatun Lake area 

(late Oligocene), locality 54n.

Genus Pleurofusia Gregorio

Gregorio, Annales Geol. Paleontologie, pt. 7, p. 33, 1890. 

Type (orthotype): Pleiirotoma (Pleurofusia) longirostropsia 

Gregorio, locality and age not specified.

Though Gregorio did not specify a locality or age 

for Pleurotoino, longlrostropsis, he presumably 

thought it is a Claiborne species; that is, according to 

present terminology, a species from the middle Eocene Gosport sand at Claiborne, Alabama. Dr. K. Y. 

W. Palmer, Director of the Paleontological Research 

Institution, informs me that the Institution has received from the University of Palermo (the repository of Gregorio's collections) three specimens labelled 

Pleurotoma longirostropsis, but none resembles the 

illustration published by Gregorio as plate 2, figure 

26. Harris, who reproduced Gregorio's figure 26 (Harris, 1937, p. 51, pi. 10, fig. 6) was unable to identify 

this species. No species closely resembling Gregorio's 

illustration is known to occur at Claiborne or elsewhere in the Eocene or Oligocene deposits of the southeastern states. Gregorio, of course, may have had a 

rare species or his drawing may be faulty. A minute 

specimen in the USNM collection from Claiborne, both 

ends of which are broken (height 4.9 mm), has the 

anal sinus in a strongly constricted sutural area 

marked with conspicuous growth threads, as shown in 

Gregorio's figure, but lacks the two strong peripheral 

spiral threads on spire whorls and the subangulated 

early whorls of that illustration. Casey (1904, p. 152) 

thought that "a specimen in the cabinet of Mr. Aid-- 

rich, from the lower Claiborne" greatly resembles 

Gregorio's illustration. A turrid in the Aldrich collection labelled "Pleurofusia longirostropsis Greg., per 

Casey, Lee Co., Texas, L. C., Eocene," doubtless is the 

specimen Casey had in mind. It is not the species represented by Gregorio's illustration. Gregorio compared 

his species with Pleurotoma, servata Conrad, an Oligocene species from the Byram marl, cited in the third 

following paragraph; indeed, despite the lack of similarity, he thought it may be a variety of the Oligocene species.

It is unfortunate that nothing more than Gregorio's 

drawing is available for the type species of Pleurofusia. Nevertheless, it is proposed to treat the name 

as the name for a genus embracing small to moderately large turriculine turrids that have a Fusinuslike outline and sculpture, and an asymmetrical sinus 

adjoining the suture. Such species are found in American faunas since Eocene time, and now live in western 

Atlantic and eastern Pacific waters at low latitudes.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 367

At present it is uncertain what species other than 

Pleurotoma longirostropsis are to be referred to the 

subgenus Pleurofusia s.s. The protoconch of the type 

species is unknown, and some post-protoconch features 

appear to be unique. Eocene species of Pleurofusia 

have been described by Harris (1937, p. 50-53) and 

Harris and Palmer (1946-47, p. 429-431, 1947). 

Whether the species that have a short pillar should 

be referred to that genus is debatable; Pleurotoma 

collaris Casey (1903, p. 270; Harris and Palmer, 1946- 

47, p. 431, pi. 59, figs. 13-15, 1947), for example, seems 

to be a clavine species. The protoconch of not all of 

the long-pillared Eocene species have been described. 

Those that are known are blunt, smooth, and about 

2-whorled.

Pleurofusia servata (Conrad) (1847 (1848), p. 284; 

Harris, 1937, p. 50, pi. 10, fig. 4, not fig. 5; Harris and 

Palmer, 1946-47, pi. 59, fig. 21, 1947), from the Oligocene Byram marl, has an acute, 3i/£- to 4-whorled 

protoconch. The later part of the last whorl bears four 

to six axial riblets. Pleurofusia olivia (Casey) (1903, 

p. 269), from the Oligocene Ked Bluff clay, has a 

similar protoconch. This type of protoconch was assumed to be typical for Pleurofusia when Fusiturricula 

(Woodring, 1928, p. 165; type (orthotype) : Turris 

(Surcula) fusinella Dall, living, Gulf of Panama) was 

proposed as a generic name for species that have a 

range of Miocene to the present time, but that assumption was unjustified. The status of Fusiturricula, 

which has a blunt or moderately acute, li/£- to 2%- 

whorled, smooth protoconch (last half whorl rounded 

or slightly bulging), is uncertain, on account of the 

uncertainty concerning Pleurofusia s.s. If it is to be 

retained, it rates subgeneric rank under Pleurofusia. 

The type still is the only dredged specimen of the 

type species of Fusiturricula. It is a small, thin-shelled, 

deep-water shell dredged at a depth of 153 fathoms. 

Though it may be immature (height 17.2 mm), it is not 

much smaller than Pleurofusia ence (Bartsch) (1934, 

p. 13, pi. 3, figs. 1, 2, 10; height 25 mm), a similar, thinshelled, deep-water species.

P. ence is represented by eight specimens, taken in 

hauls at five stations on the south border of the Puerto 

Eico Trench, off Puerto Eico, and the Virgin Islands, 

at depths of 240 to 360 fathoms. Three of these thin 

shells, including the type, as shown in Bartsch's figure 

2, have a perfectly preserved anal sinus, showing the 

wide forward sweep of the lower limb. Until this species and P. fenimorei (Bartsch) (cited under P. cf. P. 

fenimorei) were described, Pleurofusia was unknown 

in the western Atlantic Ocean.

Subgenus? 

Pleurofusia species 

Plate 48, figure 27

Of medium size, apex and pillar broken. Sculpture 

consisting of wide, low axial swellings (four on penult 

whorl), overridden by two moderately strong peripheral spiral threads. Thread (later two threads) on spire 

whorls between peripheral threads and succeeding 

whorl, and corresponding threads on base of body 

whorl almost as strong as those on periphery. Subsutural thread and that at base of subsutural area 

narrower. Anal sinus on depressed subsutural area.

Height (incomplete) 18.7 mm (estimated restored 

height 30 mm), diameter 9 mm (figured specimen).

The description of this unnamed species is based 

on an incomplete fossil, replaced by calcite, collected 

from the shallow-water facies of the Caimito formation in the Gatun Lake area at locality 56. In general 

features it resembles the Gatun species Pleurofusia 

fusinus, but lacks secondary spiral threads. Its protoconch and outer lip are, of course, unknown.

A poorly preserved fragment of two whorls from 

the La Boca formation is listed as Pleurofusia? sp.

Occurrence: Caimito formation, Gatun Lake area 

(late Oligocene), locality 56.

Pleurofusia acra Woodring, n. sp. 

Plate 57, figure 8; plate 65, figure 3

Of medium size, Fusinus-like in outline and gross 

sculpture, pillar long and slender. Protoconch acute, 

3i/^-whorled, last two whorls carinate below middle 

of whorl. Post-protoconch whorls bearing closely 

spaced, wide axial swellings below anal fascicle (8 on 

body whorl), disappearing on body whorl below shoulder. Axial swellings overridden by spiral threads (5 

on penult whorl). Similar threads on body whorl below 

lower end of axial swellings undulated by low, elongate nodes, except near end of pillar, where the threads 

are smooth. Anal fascicle of late whorls bearing weak 

secondary spiral threads and weak, retractive subsutural swellings. Outer lip broken. Apex of anal 

sinus, as shown by growth lines, close to lower edge 

of anal fasciole.

Height (practically complete) 22.4 mm, diameter 

7.5 mm (type).

Type: USNM 645764.

Type locality: 155c (USGS 16915, Gatun Third 

Locks excavation, east side of excavation, 1 mile (1.6 

km) north of Gatun Lake, Canal Zone), middle part 

of Gatun formation.

Though the type of this species, which may be immature, is the only specimen, it combines three un-
368 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

usual features: acute, polygyrate, carinate protoconch; 

retractive subsutural swellings; and noded spiral 

threads on the lower part of the body whorl. Pleurofusia fusineUa Ball (1908, p. 261, pi. 14, fig. 7, living, 

Gulf of Panama), the type of Fusiturricula, has an 

acute, polygyrate protoconch. Its last half whorl bulges 

at the periphery, but is not sharply carinate. The retractive subsutural swellings and the faintly noded 

spiral threads on the lower part of the body whorl of 

P. acra appear to be unique among American fossil 

and living species of Pleurofusia.

Occurrence: Middle part of Gatun formation (middle Miocene), eastern area, locality 155c.

Pleurofusia cf. P. fenimorei (Bartsch)

Two incomplete shells from the Chagres sandstone 

represent an undescribed species of Pleurofusia. The 

protoconch, early post-protoconch whorls, and end of 

the pillar are missing on both. The periphery of the 

whorls is strongly inflated and the anal fasciole correspondingly strongly constricted. The strong axial 

swellings (9 on body whorl) are separated by spaces 

slightly wider than the swellings themselves. The 

spiral sculpture consists of primary (4 on penult 

whorl) and secondary threads. The estimated restored 

height is between 25 and 30 mm, the diameter 10 mm.

The whorl outline and its few, strong, widely spaced 

swellings suggest Pleurofusia fenimorei (Bartsch, 

1934, p. 7, pi. 2, figs. 4, 5), a large species (height 71 

mm) dredged on the south border of the Puerto Rico 

Trench at a depth of between 80 and 180 fathoms. At

the same diameter the fossils have two more swellings 

and their primary spiral threads are fewer and 

stronger. The same dredge haul yielded another shell 

of P. fenimorei that is smaller than the type, but in 

better condition than the somewhat worn type.

P. fenimorei is the type of Fusisyrinx (Bartsch, 

1934, p. 7). The only essential difference between Fusisyrinx, as now known, and the type of Fusiturricula 

(cited in the discussion under Pleurofusia) lies in the 

number, width, and spacing of the axial ribs or swellings. The protoconch of Fusisyrinvo, however, is unknown.

Occurrence: Chagres sandstone (late Miocene), 

locality 208.

Subgenus Cruziturricula Marks

Marks, Bull. Am. Paleontology, v. 33, no. 139, p. 129, 1951. 

Type (orthotype): Turricula (Pleurofusia) cruziana Olsson, 

Miocene, Peru, Ecuador.

Cruziturricula is distinguished from other groups of 

species assigned to Pleurofusia by its blunt or acute 

protoconch, the last whorl or less of which is carinate 

below the middle of the whorl and bears fine arcuate 

axial riblets above the carina. The axial sculpture consists of swellings rather than ribs and is present or 

absent on late whorls. As shown in the following 

table, the fossil species have a lirate outer lip, whereas 

the lip of the living species is nonlirate. Also as shown 

in the table, Cmziturricula has an age range of Oligocene to present time and survives in the eastern Pacific Ocean.

Species of Pleurofusia, subgenus Cruziturricula

Age and locality

Living, 

Panamic province

Middle and late 

Miocene, Trinidad

Middle Miocene, 

Panamd

Middle Miocene, 

Ecuador

Early Miocene, 

Ecuador; middle 

Miocene, Peru

Early Miocene, 

Florida

Oligocene,

Mississippi

Species

Pleurofusia panthea (Dall)

P. arcuata (Reeve)

P. fusinus sanctidavidis 

(Maury)

P. fusinus fusinus (Brown and 

Pilsbry)

P. latira (Olsson) (=P. 

fusinus fusinus)

P. cruziana (Olsson)

P. glypta (Gardner)

P. vicksburgensis (Casey)

Protoconch

Unknown, except later part of last whorl; carinate, 

axial riblets above carina.

Blunt, 2-whorled; last whorl carinate, axial riblets 

above carina.

Blunt, l%- to 2-whorled; last % to IK whorls carinate; 

axial riblets above carina.

Blunt, 1%- to 2-whorled; last % to 1 whorl carinate; 

axial riblets above carina.

Blunt, 2-whorled; last half whorl carinate, axial riblets 

above carina.

Tip broken, evidently blunt, about 2-whorled; last % 

whorl carinate, axial riblets above carina.

Blunt, 1^-whorled; last }£ whorl carinate, axial riblets 

above carina.

Acute, 3- to 4-whorled; last }i to % whorl carinate, 

axial riblets above carina.

Interior of outer 

lip

Nonlirate.

Nonlirate.

Lirate.

Lirate.

Lirate.

Lirate.

Lirate.

Lirate.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 369

Marks assigned the two living Panamic species to 

C'ruziturricula, but under erroneous names. His "Turricula" arcuata Dall evidently is an inadvertent error 

for "Reeve": the species described by Reeve (1843, pi.

3. fig. 15, and accompanying text) as Pleurotoma arcuata. For "Turricula" lavinia Dall, Marks intended 

to mean Turricula (Surcula) panthea Dall (1919, p.

4. pi. 1, fig. 6 [not fig. 5 as printed]), not Turricula 

(Surcula) lavinia Dall (Idem, p. 4, pi. 1, fig. 5 [not fig. 

6 as printed]). Marks was misled by errors in Dall's 

publication, involving transposition of figure numbers 

on page 4 and in the explanation of plate 1. Turricula 

lavinia is a clavine species, not turriculine. Marks was 

not the first paleontologist to be misled by these errors. 

Olsson (1922, p. 55, pi. 4, fig. 6) named a Miocene 

Costa Rican species as Turricula lavinoides on account 

of the supposed relation to the alleged Turricula lavinia. The Miocene species, however, is a species of 

the living Panamic genus Knefastia. I am indebted to 

Dr. Iv. V. W. Palmer for forwarding Marks' illustrated specimen of Cruziturricula cruziana, and to Dr. 

L. G. Hertleiii for forwarding the shell illustrated by 

Marks as Cruziturricula arcuata (Dall).

Pleurofusia (Cruziturricula) fusinus fusinus 

(Brown and Pilsbry)

Plate 57, figures 16, 21; plate 65, figure 4

Drillia fusinus Brown and Pilsbry, Acad. Nat. Sci. Phila. Proc., 

v. 63, p. 344, pi. 23, fig. 7, 1911 (Miocene, Canal Zone).

Fusiturricula latira Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst, p. 94, pi. 20, 

fig. 4, 4a, 1964, (Miocene, Ecuador).

Moderately large, Fusinus-\ike in outline and sculpture ; pillar long and slender. Protoconch blunt, almost 

flat-topped, 1%- to 2-whorled, last 1/4 to 1 whorl carinate below middle of whorl, bearing fine, arcuate axial 

riblets above carina. End of protoconch marked by 

strengthening of carina to form lower peripheral spiral 

thread, and appearance of upper peripheral and subsutural spiral threads, and closely spaced, wide axial 

swellings. Upper peripheral spiral thread gradually 

strengthened to become as strong as lower. On spire 

whorls these two peripheral primary spiral threads 

slightly to decidedly stronger than subsutural primary, 

decidedly stronger than primary thread at lower edge 

of depressed sutural area, and slightly stronger than 

primary thread below periphery. Secondary spiral 

threads, generally of two ranks, on early post-protoconch and later whorls. Axial sculpture consisting of 

low, wide swellings, generally weaker on body whorl 

(or last two whorls), or narrower and lower on part 

of body whorl. Anal sinus, as shown by growth lines,

asymmetric; upper limb short, lower limb long. Interior of outer lip lirate.

Height (incomplete) 45.T mm, diameter (incomplete) 13.5 mm (larger figured specimen). Estimated 

restored height 60 mm, estimated restored diameter 

16 mm (largest specimen).

Type: Acad. Nat. Sci. Phila. 1693.

Type locality: Gatuu Locks site, Canal Zone, middle 

part of Gatun formation.

Variation, on a minor scale, affects the strength of 

primary spiral threads with reference to the two on 

the periphery and the strength of axial swellings on 

the last whorl or two. The outer lip and the long 

slender pillar are invariably broken. On the type and 

a few other shells, one of which is shown on plate 57, 

figure 16, the pillar is practically complete. The lirae 

on the interior of the outer lip appear at intervals on 

the inside of the body whorl, corresponding to former 

positions of a fully developed outer lip. The readily 

recognizable protoconch so blunt that it is almost 

flat-topped is preserved on 36 of the 141 specimens. 

If it is even slightly worn, the fine axial riblets disappear.

This turrid is widespread in the Gatun formation, 

but was not found in the upper part in the western 

area. The largest collections are those from the lower 

part at locality 138c and the upper part in the eastern 

area at locality 175: 34 and 16 specimens, respectively. 

It has been found in two other Panama areas: in 

Darien (USGS 8433) and Chiriqui (USGS 7955).

Turris albida sanctidavidis (Maury, 1925, p. 188, 

pi. 32, figs. 6, 7), which is widespread in the late 

middle Miocene San Jose calcareous silt member of the 

Manzanilla formation of Trinidad, is classified as a 

scarcely distinguishable subspecies of P. fusinus. Fifty 

specimens are represented in 14 USNM collections. It 

is considerably smaller than the nominate subspecies, 

as the maximum height ranges from 40 to an estimated 

45 mm. The axial swellings generally disappear on the 

third to fifth post-protoconch whorl. Two exceptional 

shells, however, on which they disappear on the seventh 

and ninth whorl, are indistinguishable from immature 

shells of the nominate subspecies. P. fusinus sanctidavidis occurs also in the late Miocene Melajo clay 

member of the Springvale formation of Trinidad 

(USGS 21178, 18634). On the four specimens that 

show the early sculpture the swellings disappear on 

the third or fourth whorl.

The specimen illustrated by Maury in her figure 7 

(Paleontological Research Inst. 1008) is herewith 

selected as the lectotype.

Except for its blunt protoconch of fewer whorls, 

larger size of shell, and larger apical angle, Pleura-
370 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

fusia fusinus fusinus resembles the Oligocene species 

Pleurotoma vicksburgensis Casey (1903, p. 268, Harris, 

1937, p. 51, pi. 10, fig. 3; Harris and Palmer, 1946-47, 

pi. 59, fig. 22, 1947; height 27 mm), from the Byram 

marl. The acute, polygyrate protoconch of Pleurofusia 

vicksburgensis was described by Casey, and is shown 

in Harris and Palmer's illustration, and also in Harris' 

(1937, pi. 10, fig. 5) illustration of a fossil cited as 

P. servata. Casey, however, attributed the carinate 

and axially sculptured part to the earliest post-protoconch stage. Despite the difference in protoconch, P. 

vicksburgensis and P. fusinus fusinus are considered 

to be closely related. P. latira is indistinguishable from 

P. fusinus fusinus. The axial swellings of P. cruziana 

(Olsson, 1932, p. 150, pi. 15, figs. 6, 10; Marks, 1951, 

p. 131, pi. 8, fig. 6) are more subdued than those of 

P. fusinus fusinus.

The early Miocene P. glypta (Gardner) (1926-47, p. 

293, pi. 38, fig. 34, 1938; Chipola formation, height 

25.3 mm) is not closely related to any of the other 

species of Cruziturricula. Its last protoconch whorl 

merges into the first post-protoconch whorl. The first 

three post-protoconch whorls are unicarinate, angulated by the continuation of the protoconch carina, 

and bear four weak spiral threads above the carina. 

Low axial swellings appear on the lower part of the 

third whorl. The type of P. glypta is the only specimen now in Gardner's collections, although an additional locality was cited. The protoconch of the type 

is worn. The preservation is better on two minute 

shells (USNM 644754) in C. E. Locklin's collection 

from the type locality: McClelland's farm, Calhoun 

County, Fla.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 137a, 138, 138a, 138c, 138d, 138e, 138f. Middle 

part, eastern area, localities 139b, 139c, 142, 146, 147b, 

147f, 147g, 151, 155, 155b, 155c, 157, 159, 159d; western 

area, locality 161c. Upper part, eastern area, localities 

I73a, 175, 176, I77b. Middle Miocene deposits, Darien 

and Chiriqui areas, Panama. Angostura formation 

(middle Miocene), Ecuador.

Genus leucosyrinx Ball

Ball, Harvard Coll. Mus. Comp. Zoology Bull., v. 18, p. 75,

1889. 

Type (orthotype): Pleurotomella verrillii (Ball) (Pleurotoma

(Pleurotomella) verrillii), North Carolina and eastern

Gulf of Mexico.

leucosyrinx xenica Woodring, n. sp. 

Plate 57, figure 15

Small for the genus, very slender, anal fascicle 

barely constricted, forming a steep ramp. Protoconch

blunt 1%-whorled, last whorl carinate, the carina gradually retreating toward base, end marked by gradual 

appearance of sculpture. First post-protoconch whorl 

sculptured with 10 swollen axial ribs, strong only on 

periphery, fadng out on anal fascicle, subdued toward 

base of whorl. Nine ribs on antepenult whorl. Ribs 

lower, wider, and gradually fading out on penult, absent on body whorl, except a slight swelling at beginning of whorl. Spiral sculpture obscure on first postprotoconch whorl, gradually strengthened on succeeding whorls, consisting of closely spaced, narrow threads 

of unequal width, subdued on anal fascicle. Outer lip 

broken back. As shown by growth lines, anal sinus wide, 

adjoining suture, upper limb short, lower limb long. 

Aperture long, narrow. Siphonal canal not notched, 

according to growth lines. Siphonal fascicle not inflated.

Height 15.4 mm, diameter 4.9 mm (type).

Type USNM 645768.

Type locality: 185 (USGS 8383, Caribbean coast, 

west of Eio Miguel, station 26 plus 100 feet (30 m), 

Panama), upper part of Gatun formation.

The type, the only specimen of this unusual turrid, 

was found in the upper part of the Gatun formation 

in the western area. The combination of barely constricted anal fascicle, disappearing axial sculpture, 

and fairly strong spiral sculpture is unique in the 

genus. Only two other species are known in the Tertiary Caribbean province: Leucosyrinx chloris Olsson 

(1922, p. 59, pi. 5, fig. 19; Limon formation, Bocas del 

Toro area, Panama) and L. nicoya Olsson (1942, p. 

57, pi. 12, fig. 2; Charco Azul formation, southwestern 

Panama), and none in southeastern United States.

Leucosyrinx is basically a deep-water genus. It has 

been dredged in western Atlantic waters at depths of 

90 to 1,180 fathoms, for the most part between 300 

and 800. It is improbable, however, that the fossils 

from the type locality of L. xenica represent a depth 

greater than 100 fathoms, if that much.

Occurrence: Upper part of Gatun formation (middle Miocene), western area, locality 185.

Genus Cochlespira Conrad

Conrad, Am. Jour. Conchology, v. 1, p. 19, 1865. 

Type (monotype): Pleurotoma, cristata Conrad, Oligocene, 

Mississippi.

During recent years Cochlespira engonata Conrad 

generally has been accepted as the type species. That 

species and Pleurotoma cristata were assigned to Cochlespira by Conrad. In 1913, however, Vincent (1913, 

p. 25) pointed out that Cochlespira engonata was a 

nude name when Cochlespira was proposed, and was 

not validated until three (actually a little more than 

four) months later (Conrad, 1865, p. 142; Conrad,
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 371

1865a, p. 210, pi. 21, fig. 12). Pleurotoma cristata therefore is the monotype. Cochlespira engonata is the type 

of Cochlespiropsis (Casey, 1904, p. 143; logotype, designated by Cossmann, 1906, p. 221).

Cochlespira? species

A fragment of 2^ incomplete whorls, found in the 

moderately deep-water facies of the Caimito formation 

on Barro Colorado Island, probably represents Cochlespira. The peripheral frill is slightly upturned. It is 

faintly noded, but if it had been serrate, the spines are 

gone. A faint spiral thread lies in the concavity between the frill and the suture, closer to the frill than 

to the suture. The penult whorl has a height of about 

1.5 mm.

Occurrence: Caimito formation, Gatun Lake area 

(late Oligocene), locality 54g.

Subgenus Ancistrosyrinx Ball

Dall, Harvard Coll. Mus. Comp. Zoology Bull., vol. 9, p. 53,

1881. 

Type (monotype): Ancistrosyrinx elegans Dall, living, West

Indies.

The protoconch of Ancistrosyrinx is about li/£- 

whorled and bluntly tipped, whereas the protoconch 

of Cochlespira is about 21/^-whorled and has an acute 

tip. The peripheral frill of Ancistrosyrinx is more 

strongly upturned than that of Cochlespira, producing 

a deeper concavity between the frill and the suture. 

These are minor differences. Nevertheless Harris' 

(1937, p. 45) suggestion that a group of species, ranging in age from middle Eocene to the present time, 

may be assigned to Ancistrosyrinx as a subgenus of 

Cochlespira is adopted.

Cochlespira (Ancistrosyrinx) cedonulli (Reeve) 

Plate 57, figures 9, 10

Pleurotoma cedo-nulli Reeve, Zool. Soc. London Proc., p. 185, 

1843 (Recent, Panama). Reeve, Conchologia iconica, v. 1, 

Pleurotoma, species 117, figs. 117, 117a, 1843 (living, 

Panama).

Ancistrosyrinx cedonulli reevi Olsson, Bull. Am. Paleontology, 

v. 27, no. 106, p. 53, pi. 10, fig. 4, 1942 (Pliocene, Costa 

Rica). Olsson, Neogene mollusks from northwestern 

Ecuador, Paleontological Research Inst, p. Ill, pi. 15, 

fig. 5, 1964 (Miocene or Pliocene, Ecuador).

Pleurotoma miranda Guppy, Sci. Assoc. Trinidad Proc., v. 2, 

no. 4, pt. 12, p. 178, pi. 7, fig. 19, 1882 (Miocene, Jamaica) ; 

reprint, Bull. Am. Paleontology, v. 8, no. 35, p. 99, pi. 5, 

fig. 19, 1921.

Ancistrosyrinx miranda (Guppy), Woodring, Carnegie Inst. 

Washington Pub. 385, p. 165, pi. 6, figs. 1-3, 1928 (Miocene, Jamaica). Perrilliat Montoya, Mexico Univ. Nac., 

Inst. Geologia Paleontologia Mexicana, no. 8, p. 29, pi. 4, 

figs. 11,12,1960 (Miocene, Mexico).

Ancistrosyrinx dalli Olsson, Bull. Am. Paleontology, v. 9, no. 

39, p. 60, pi. 4, fig. 16, 1922 (Miocene, Panama).

Of medium size, moderately inflated. Protoconch 

about li/^-whorled, tip broken. First post-protoconch 

whorl bearing discrete, swollen nodes. With further 

growth nodes joined and extended into flat triangular 

spines, forming serrate peripheral frill. Spines more 

or less broken 011 intermediate and late whorls. A spiral 

thread, appearing near upper base of frill on third 

post-protoconch whorl, gradually becoming stronger 

and sharply raised, and gradually shifted backward 

until at aperture it is almost halfway between frill 

and suture. An obscure spiral thread on body whorl 

emerging at upper end of aperture. Pillar bearing 

weak spiral threads. Anal sinus very deep, lying between suture and spiral cord.

Height (incomplete) 19.3 mm, diameter (spines 

broken) 10.2 mm (figured specimen).

Type: British Museum (Natural History).

Type locality: Panama Bay, living.

A minute shell from the middle part of the Gatun 

formation and the figured specimen from the upper 

part in the eastern area are referred to Cochlespira 

cedonulli.

Three living American species of the subgenus Ancistrosyrinx are recognized: Cochlespira elegans Dall 

(1881, p. 54; 1889, p. 78, pi. 38, fig. 3) and C. radiata 

Dall (1889, p. 78, pi. 12, fig. 12) in the Caribbean 

region, and C. cedonulli in the Panamic region. C. 

elegans (height 32 mm, depth range 260 to 805 fathoms) is sculptured below the frill with strong, noded 

spirals. The corresponding spiral threads of C. radiata 

(height 25 mm, depth range 22 to 170 fathoms) are 

nonnoded and moderately strong. C, cedonulli (height 

23 mm, depth range 10 to 153 fathoms) lacks spiral 

threads between the frill and the pillar, except that 

on some specimens, both living and fossil, an obscure 

thread emerges at the upper end of the aperture.

These three species appeared abruptly in the middle 

Miocene sea of the present Caribbean region. The 

Miocene representatives of both living Caribbean species, however, are smaller than living shells. Whether 

the small fossils are to be considered as a small predecessor race or as immature is uncertain until larger 

samples are available. The occurrence of only small 

specimens of C. cedonulli at some fossil localities suggests that the small specimens of all three species are 

immature. Four small specimens of C. elegans (maximum height 17 mm) have been found in Costa Rica 

Olsson's (1922, p. 60, pi. 4, fig. 17) Ancistrosyrinx elegans, variety and in the Cercado and Gurabo formations of the Dominican Republic. The Gurabo formation yielded also a small specimen (height 18 mm) of
372 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

(7. radiata. A small shell (height 12 mm) in Spencer's 

collection of deep-water Miocene fossils from Tehuantepec has a strong spiral thread emerging at the upper 

end of the aperture and the thread is flanked on both 

sides by a smooth space. Should this type of sculpture 

prove to be consistent, treatment as a subspecies of C. 

radiata would be warranted. Bose's (in Bose and Toula, 

1910, p. 251, pi. 13, fig. 26) Pleurotoma (Ancistrosyrinx] aff. radiata, from the same locality, evidently is 

not a species of Cochlespira.

C. cedonulli is fairly widespread in middle Miocene 

deposits: Jamaica, the Tehuantepec area of Mexico, 

Costa Rica, and Panama. Small Jamaican specimens 

(maximum height 13 mm) were named Pleurotoma 

mimnda. They are slender, but not more slender than 

an immature living shell of C. cedonulli (USNM 

96801). The fossils from the Tehuantepec area are 

comparable in size to Jamaican specimens; those from 

Costa Rica and Panama are of medium size (20 mm 

and about 21 mm, respectively). The largest fossils 

(height 35 to 43 mm), larger than any available living 

shell, occur in the present Panamic region: in the 

Esmeraldas formation of Ecuador and the Charco 

Azul formation of southeastern Costa Rica.

Though the pattern of former distribution and survival shown by C. cedonulli is unusual at the specific 

level, it is common at subgeneric and generic levels.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

locality 147b. Upper part, eastern area, locality I77b. 

Bowden formation (middle Miocene), Jamaica. Middle 

Miocene deposits, Tehuantepec area, Mexico, Costa 

Rica. Limon formation (late Miocene), Bocas del Toro 

area, Panama. Esmeraldas formation (late Miocene 

or Pliocene), Ecuador. Charco Azul formation (Pliocene), Costa Rica. Living, Mazatlan, Mexico, to 

Panama Bay.

Subfamily BOESONIINAE 

Genus Scobinella Conrad

Conrad, Acad. Nat. Sci. Phila. Jour., 2d ser., vol. 1, p. 120, 1848. 

Type (monotype): Scobinella cwlata Conrad, Oligocene, Miss.

In southern United States Scolinella, an extinct 

endemic American genus, has an age range of Eocene 

to Oligocene, but in the Tertiary Caribbean province 

it survived until the end of Miocene time. The Eocene 

and Oligocene species heretofore described have a 

height of 10 to 35 mm, whereas the Miocene species 

are larger: height 42 to 70 mm. The Miocene species 

have been reviewed recently by Olsson (1964, p. 114- 

117).

Scobinella morierei (Cossmann) 

Plate 58, figures 26, 27

Euchilodon Morierei Laville mss., Cossmann, Jour. Conchyliologie, v. 61, p. 34, pi. 3, figs. 6, 7, 1913 (Miocene, Canal 

Zone).

Scobinella Morierei (Laville) Cossmann, Olsson, Bull. Am. Paleontology, v. 9, no. 39, p. 79, pi. 4, figs. 3, 4, 1922 (Miocene, 

Panama).

Scobinella morierei (?) (Laville), Anderson, Calif. Acad. Sci. 

Proc., 4th ser., v. 18, no. 4, p. 131, pi. 8, figs. 6, 7, 1929 

(Miocene, Colombia).

Scobinella morierei (Cossmann), Olsson, Neogene mollusks from 

northwestern Ecuador, p. 115 (in discussion), pi. 19, figs. 

4, 4a, 1964 (Miocene, Panama).

Scobinella moricri gavilanensis Rutsch, Eclogse Geol. Helvetia, 

v. 23, p. 611, pi. 17, figs. 8, 9, 1930 (Miocene, Venezuela). 

Rutsch, Schweitzer. Palseorit. Gesell. Abh., v. 55, no. 1, 

pi. 8, fig. 17, text fig. 13, 1934 (Miocene, Venezuela).

Not Scobinella morierei (Laville) Cossmann, Maury, Bull. Am. 

Paleontology, v. 10, no. 42, p. 193, pi. 34, fig. 1, 1925 

(Miocene, Trinidad).

Not Scobinella morierei (Laville), Perrilliat Montoya, Mexico 

Univ. Nac. Inst. Geologia Paleontologia Mexicana, no. 8, 

p. 28, pi. 4, figs. 7, 8, 1960 (Miocene, Mexico).

Moderately large, slender, turreted, anal fascicle 

strongly constricted. Protoconch not preserved. Sculpture of earliest preserved whorls consisting of a noded 

sutural band; arcuate axial threads on anal fascicle; 

closely spaced protractive axial ribs, noded by two 

peripheral spiral cords and a narrower cord at base 

of whorl. Sculpture of later whorls consisting of retractive nodes on sutural band; two closely spaced 

rows of small nodes on anal fascicle (three on-body 

whorl), arranged in axially arcuate pattern; coarsely 

noded shoulder cord, bilirate on last 2i£ whorls, upper 

part wider and more coarsely noded than lower part; 

and up to body whorl two or three noded spiral cords 

similar to narrow lower part of shoulder cord. Nodes 

arranged in protractive pattern. On pillar nodes progressively suppressed toward siphonal fascicle. Anal 

sinus and siphonal notch moderately deep, as shown 

by growth lines. Interior of outer lip lirate. Columella 

bearing two widely spaced upper folds and two closely 

spaced, weak lower folds. Siphonal fascicle slightly 

inflated, bearing a few widely spaced, weak spiral 

threads.

Height (almost complete) 37 mm (estimated height 

42 mm), diameter (incomplete) 12.5 mm.

Type: Ecole des Mines, Paris.

Type locality: Mindi, Panama Canal, Canal Zone, 

upper part of Gatun formation.

The description is based on an almost complete 

specimen of moderate size from the lower part of the 

Gatun formation the only specimen in the collections 

now being studied. The type (height 42 mm) was col-
GASTROPODS: ETJLIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 373

lected at an outcrop of the upper part of the Gatun 

in the Mindi area near locality 172.

Scobinella morierei occurs also in the late Miocene 

Limon formation at Limon, Costa Rica, and on islands 

of the nearby Bocas del Toro Archipelago, Panama. 

The change from arcuate axial threads to small nodes 

on the anal fascicle takes place two or three whorls 

earlier on these late Miocene shells than on the Gatun 

specimen. Shells from Costa Rica (Olsson, 1964, pi. 

19, fig. 4a) are decidedly less turreted than the Gatun 

form; those from the Bocas del Toro area (Olsson, 

1922, pi. 4, figs. 3, 4; 1964, pi. 19, fig. 4) are somewhat 

less turreted.

The sutural band of Maury's Trinidad specimen is 

bipartate or tripartate and swollen, so that it affects 

the whorl profile. As suggested by Olsson (1964, p. 

115), the illustrations of S. morieri gamlanensis indicate no features to substantiate that subspecies. If 

Anderson's Scobinella is S. morierei the age range of 

the species is early to late Miocene.

Though the early and middle Miocene Ecuadorean 

S. onzola Olsson (1964, p. 116, pi. 19, figs. 2, 2a, 2b) 

reaches a slightly larger size (estimated height 58 mm) 

and is slightly more coarsely noded, it is very similar 

to S. morierei.

The flat-sided, biconic Mexican Scobinella, recorded 

as S. morierei, is relatively small for a Miocene species (height 38 mm). A slightly smaller, more slender, 

biconic species occurs in the Brasso formation of Trinidad (USGS 21234).

The name Scobinella morierei is to be attributed to 

Cossmann. According to his statement (1913, p. 36), 

there is no indication that he published Laville's description of the species. Where or how Cossmann 

picked up the trivial name is irrelevant.

Occurrence: Lower and upper parts of Gatun formation (middle Miocene). Lower part, locality 138a. 

Upper part, Cossmann's record. Las Perdices shale 

(early Miocene), Colombia. Limon formation (late 

Miocene), Costa Rica, and Bocas del Toro area, 

Panama. Punta Gavilan formation (late Miocene), 

Venezeula.

Scobinella aff. S. morierei (Cossmann)

Three defective specimens establish the presence of 

Scobinella in the moderately deep-water facies of the 

Caimito formation on Barro Colorado Island. So far 

as the meager material goes, this Oligocene Scobinella 

is closely allied to S. morierei, but the shoulder nodes 

are more widely spaced. The largest of these incomplete specimens has a height of 30 mm and an estimated total height of about 40 mm; that is, comparable to S. morierei and larger than any other known 

Oligocene species.

Occurrence: Caimito formation (late Oligocene), 

Barro Colorado Island, localities 54h, 54m.

Scobinella ecuadoriana Olsson 

Plate 62, figures 33, 34

Scobinella ecuadoriana Olsson, Neogene mollusks from northwestern Ecuador, p. 117, pi. 19, figs. 1, la, Ib, 1964 (Miocene, Ecuador).

The Chagres sandstone yielded an incomplete large 

Scobinella identified as S. ecuadoriana. It shows the 

most distinctive feature of that species: large nodes 

on the shoulder, immediately below the anal fascicle, 

even on the penult whorl, as well as on the body whorl. 

The interior of the outer lip is lirate and the columella bears four folds, arranged like those of 8. morierei. The height of the body and penult whorls is 43.4 

mm, indicating a total height of about 63 mm.

8. ecuadoriana is a large species, but not as large as 

the appropriately named middle Miocene S. magnifica (Gabb) (Olsson, 1964, p. 115, pi. 19, figs. 3, 3a), 

which occurs in the Dominican Republic and Jamaica 

(height 70.5 mm).

Occurrence: Chagres sandstone (late Miocene), 

locality 208. Esmeraldas formation (late Miocene), 

Ecuador.

Genus Paraborsonia Pilsbry

Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 73, p. 326, 1922.

Type (orthotype): Borsonia (Paraborsonia) varicosa (Sowerby) (Mitra varicosa Sowerby), Miocene, Dominican

Republic.

Paraborsonia is another extinct endemic American 

borsonine genus. Its age range is late Oligocene to 

late Miocene, but early Miocene species are as yet 

unknown.

Paraborsonia aff. P. brassoensis (Mansfield)

Small, biconic, anal fasciole not constricted, pillar 

short. Protoconch and about first two post-protoconch 

whorls missing. Remaining spire whorls bearing a 

sutural groove; a sutural cord sculptured with axially 

elongate nodes, separated by a groove from a wide 

band sculptured with axially elongate nodes and 

traces of obscure spiral sculpture. Noded band of 

body whorl bilirate: two low spiral threads overriding 

nodes. Remainder of body whorl up to pillar sculptured with primary and secondary noded spiral 

threads. Spiral threads on pillar not noded. Tip of 

pillar missing. Apex of anal sinus on wide noded 

band, as shown by growth lines. Interior of outer lip 

and columella inaccessible.
374 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Height (incomplete) 8.5 mm (estimated height 12 

mm), diameter (incomplete) 5.5 mm.

An incomplete, crushed specimen from the moderately deep-water facies of the Caimoto formation 

on Barro Colorado Island is identified as the earliest 

known species of Pardborsonia. If it is mature, it is 

also the smallest species. Though the shell is too fragile 

to attempt to expose the columella, in other features it 

is a Pardborsonia closely allied to the middle Miocene 

Trinidad species P. brasscensis (Mansfield, 1925, p. 30, 

pi. 5, fig. 8). The wide band of that species is bilirate 

at an earlier stage. Like Microdrittia trina (p. 389), 

P. brassoensis was based on float material, but is represented by some 40 outcrop specimens from USGS locality 21234 on Mayo River: the same locality that yielded 

an adequate sample of Microdrillia trina. P. ~brasscensis 

occurs also in the San Jose calcareous silt member of 

the Manzanilla formation (USGS 21231).

Heretofore Pardborsonia was known in the middle 

Miocene deposits of the Dominican Republic and Trinidad, and in late Miocene in the Bocas del Toro area, 

Panama. Borsonia (Pardborsonia} centaurana (Jung, 

1965, p. 572, pi. 77, figs. 11-14; middle Miocene, Venezuela) is not a Pardborsonia.

Occurrence: Caimito formation (late Oligocene), 

Barro Colorado Island, locality 54k.

Subfamily CLAVINAE 

Genus Drillia Gray

Gray, Ann. Nat. History, v. 1, p. 28, 1838.

Type (logotype, Gray, Zool. Soc. London Proc., p. 134, 1847): 

Drillia umbilicata Gray, living, West Africa.

The type species has been well illustrated by Bartsch 

(1943, p. 82, pi. 7, fig. 5, pi. 10, fig. 7).

Submenus Neodrillia Bartsch

Bartsch, Soc. Cubana Historia Nat. Felipe Poey, Mem., v. 17, 

p. 83, 1943.

Type (orthotype): Neodrillia cydia Bartsch, living, North Carolina to Florida and West Indies.

The smaller size, more slender outline, lack of an 

umbilicus, and finer spiral sculpture of Neodrillia distinguish it from Drillia s.s. Leptadrillia (Woodring, 

1928, p. 159; type (orthotype) : Turns (Surcula) parkeri Gabb, Miocene, Dominican Republic) is more 

slender than Neodrillia and has narrow axial ribs and 

no spiral sculpture, except on the pillar.

According to Pilsbry and Harbison (1933, p. 112), 

Leptadrillia "differs from Cymatosyrinx by its more 

slender shape and longer anterior canal, but there 

seems to be no other structural difference." Leptadrillia, however, lacks a constricted anal fasciole, deep 

stromboid notch, deep siphonal notch, and sharp thread

limiting the siphonal fasciole all shown by Cymatosyrinx (Dall, 1889, p. 95; type (orthotype): Pleurotoma lunata H. C. Lea, Miocene, Virginia). Cymatosyrinx is the temperate counterpart of Clathrodrillia 

(p. 381). On morphologic grounds it may be better to 

assign Cymatosyrinx lunata aclinica Tucker and Wilson (1933, p. 13, pi. 4, figs. 6, 7; Olsson and Petit, 1964, 

p. 537, pi. 82, figs. 5-5b; assigned specific rank), a 

late Miocene Florida turrid, to a subgenus under 

Clathrodrillia that has no spiral sculpture. A middle 

Miocene ally of Cymatosyrinx aclinica is found in the 

Tehuantepec area, where it has been recorded as Cymatosyrinx lunata (Perrilliat Montoya, 1963, p. 29, pi. 5, 

figs. 17, 18).

Drillia (Neodrillia) riogTirabonis eurysoma Gardner 

Plate 58, figures 17, 18; plate 64, figure 2

"Drillia" eurysoma Gardner, U.S. Geol. Survey Prof. Paper 142, 

p. 316, pi. 40, fig. 10, 1937 (1938) (Miocene, Florida).

Of medium size, moderately inflated, pillar short, 

whorls not constricted adjoining suture to form an anal 

fasciole. Protoconch acute, 3- to 31^-whorled, last 1 

to iy2 whorls bulging near base. End of protoconch 

marked by an arcuate riblet, followed by first axial 

rib. Axial ribs undulating suture except where they 

are alined and on body whorl, where they are attenuated near suture; alined on first 3 to 4^ whorls; 

practically straight on about first 3 whorls, slightly 

arcuate on remaining spire whorls, decidedly arcuate 

on body whorl; 7 or 8 on all whorls. Last rib wide 

and high, forming a hump. Spiral sculpture consisting 

of weak, closely spaced, narrow threads, not discernible 

on earliest few whorls. Outer lip broken back. Anal 

sinus moderately deep, as indicated by growth lines; 

upper limb heavily bordered. Stromboid notch very 

shallow, according to growth lines. Siphonal notch 

moderately deep. Siphonal fasciole slightly inflated, 

sculptured with spiral threads, stronger and more 

widely spaced than those elsewhere.

Height (practically complete) 11.8 mm, diameter 

(including hump) 4.6 mm (larger figured specimen).

Type: USNM 351188.

Type locality: USGS 3856, 5 to 6 miles west-northwest of Mossyhead, Walton County, Florida, Shoal 

River formation.

The larger figured specimen, on which the earliest 

part of the protoconch is missing, is the only practically complete mature shell among the six specimens 

of this Gatim turrid. Though it is identified as the 

Florida form, the axial ribs on the early whorls of the 

two available specimens from the Shoal River formation are not uniformly alined. Gardner realized that 

her species is closely related to Neodrillia riogurdbonis
GASTROPODS: ETJLIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 375

(Maury, 1917, p. 54, pi. 9, fig. 2), found in the Gurabo 

formation of the Dominican Republic. The nominate 

subspecies has one or two more axial ribs on late 

whorls and they are somewhat more attenuated near 

the suture. As on the Florida specimens, the ribs on 

early whorls are not uniformly alined. The ribs of the 

species from the late Miocene Savaneta glauconitic 

sandstone member of the Springvale formation in 

Trinidad, identified by Mansfield (1925, p. 24, pi. 3, 

fig. 8) as Drillia aff. D. riogurdbonis and by Rutsch 

(1942, p. 166) as Cymatosyrinx? sp. ind., are narrower 

and on late whorls are more decidedly attenuated near 

the suture. This species is represented in USNM collections, deposited by H. G. Kugler, by better material 

than that available to Mansfield and Rutsch. The turrids from the Bowden formation of Jamaica described 

as "Drillia" sp. and Bellaspira? sp. (Woodring, 1928, 

p. 162, 163, respectively) represent Neodrillia. Though 

the so-called Bellaspira? is poorly preserved, they 

probably are the same species a species that is larger 

and more inflated than D. riogurabonis.

The species so far mentioned, and also early Miocene 

species from the Tampa limestone of Florida and the 

Thomonde formation of Haiti (TJSGS 9945), are more 

similar to the living Panamic species D. cybele (Pilsbry and Lowe (1932, p. 46, pi. 2, fig. 6) than to the 

living Caribbean species D. cydia. The sutural area of 

the Caribbean species is constricted, forming an anal 

fascicle, and the shell is sculptured with fine, submicroscopically crimped spiral threads. The sutural 

area of D. subperpolita Bose) (in Bose and Toula, 

1910, p. 246, pi. 13, fig. 22), a late Miocene species from 

the Tehuantepec area, is constricted, but not as strongly constricted as that of D. cydia.

Abbott (1958, p. 96) was justified in placing in synonymy under D. cydia the four other living Caribbean 

species of Neodrillia named by Bartsch (1943, p. 84-89) 

and in rejecting, on the basis of Melvill's illustration, 

Bartsch's identification of the Cuban species D. euphanes Melvill (1923, p. 164, pi. 4, fig. 4). If Bartsch's 

identification were correct, D. cydia, the type species 

of Neodrillia, would be a synonym of D. euphanes.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, locality 138c. 

Middle part, eastern area, locality 139b; western area, 

locality 162. Shoal River formation (middle Miocene), 

Florida.

Drillia? (Neodrillia?) species

The upper part of the Bohio formation of Barro 

Colorado Island yielded an incomplete minute clavine 

turrid consisting of the protoconch and four postprotocoiich whorls, all replaced by calcite. The apertural face of the penult and body whorls is missing. The

protoconch is acute and 3-whorled. The post-protoconch whorls are almost flat-sided and are sculptured 

with eight strong, straight axial ribs that are alined 

or almost alined. The anal fascicle is not constricted 

and spiral sculpture is not discernible. The height, 

minus the pillar, is 4 mm, and the diameter, minus the 

outer lip, 2mm.

Despite the absence of spiral sculpture, the outline, 

protoconch, and style of ribbing point to Neodrillia. 

The absence of spiral sculpture may be due to masking 

by the replacing calcite, or the specimen may be immature. If it is Neodrillia, it is the earliest species, 

possibly a small primitive species.

Occurrence: Upper part of Bohio formation (late 

Oligocene), locality 42d.

Genus Carinodrillia Dall

Ball, U.S. Natl. Mus. Proc., v. 56, p. 17, 1919. 

Type (orthotype): Clathrodrillia halts Dall, living, Gulf of 

California to Manzanillo, Mexico.

Subgenus Carinodrillia s.s.

C aiinodrillia, like Pleurofusia, has Fusinus-\\]s& 

sculpture. It appeared in early Miocene time and survives in the western Atlantic and eastern Pacific oceans.

The subgenus Carinodrillia s.s. embraces species of 

medium to moderately large size. The type and other 

species, including C. zooki, have an acute, polygyrate 

protoconch (three to four whorls), the last 14 to 1/2 of 

which bears arcuate axial riblets. The protoconch of 

other fossil and Recent species, however, is blunt, paucigyrate (about two whorls), and near the end bears a 

few peripheral axial swellings. The siphonal canal of 

all the species described in the present report is slightly notched and the siphonal fascicle is slightly inflated.

Several western Atlantic species are characterized 

by their small size and blunt protoconch of two to 

three whorls that lack axial sculpture other than 

faint, crowded, slightly arcuate growth threads. These 

species are assigned to the subgenus Buchema Corea 

(1934, p. 1; type (orthotype): Carinodrillia (Buchema} tainoa Corea, living, Puerto Rico).

Carinodrillia (Carinodrillia) zooki (Brown and Pilsbry) 

Plate 58, figures 9-12; plate 65, figure 9

Drillia sooki Brown and Pilsbry, Acad. Nat. Sci. Phila. Proc.,

v. 63, p. 345, pi. 23, fig. 18, 1911 (Miocene, Canal Zone). 

Drillia (Crassispira) zooki Brown and Pilsbry, Cossmann, Jour.

Conchyliologie, v. 61, p. 23, pi. 3, figs. 8, 9, 1913 (Miocene,

Canal Zone). 

Not Drillia zooki Brown and Pilsbry, Li, Geol. Soc. China Bull.,

v. 9, p. 275, pi. 8, fig. 71, 1930 (Miocene, Panama Bay) ;

 Clathrodrillia alcestis Dall, fide Pilsbry, Acad. Nat.

Sci. Phila. Proc., v. 83, p. 434,1931 (Living, Panama Bay).

8-278 O - 70 - 6
376 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Of medium size, slender, Fusinus-like, aside from 

short pillar. Protoconch acute, 3- to 4-whorled, last 14 

to 1/2 whorl bearing 3 to 7 arcuate riblets terminating 

on or below the swollen, almost carinate periphery. 

End of protoconch marked by abrupt appearance of 

axial and spiral sculpture. First post-protoconch whorl 

sculptured with 6 to 8 closely spaced, wide, swollen 

axial ribs and 3 spiral threads: 1 sutural and 2 peripheral. Axial ribs on late whorls moderately swollen and 

fading out on lower part of anal fasciole, or strongly 

swollen and modifying outline of lower part of anal 

fasciole; 8 to 10 ribs on penult whorl. Spiral sculpture 

of late spire whorls consisting of a sutural thread and 

generally 3 other widely spaced primary threads overriding axial ribs; a fourth thread exposed at base of 

whorl on some specimens. Sutural thread strong on 

all whorls or relatively weak. In addition to sutural 

thread, 15 to 17 primary threads on body whorl, those 

on pillar more closely spaced and weaker than elsewhere. Microscopic secondary spiral threads and exaggerated growth threads visible on late whorls. An axial 

rib near outer lip widened to form a hump; ribs absent between hump and lip. Anal sinus deep, border 

of upper limb somewhat thickened.

Estimated restored height 25 mm, diameter (including hump) 8 mm (largest specimen). Height 19.8 mm, 

diameter (including hump) 6 mm (figured strongly 

sculptured specimen).

Type: Acad. Nat. Sci. Phila. 1695.

Type locality: Gatun Locks excavation, Canal Zone, 

middle part of Gatun formation.

The 20 specimens from the three parts of the Gatun 

formation in the eastern area identified as Carinodrillia zooki show a wide range of variation, chiefly 

affecting the strength of the axial ribs and sutural 

spiral thread. The type is an incomplete specimen that 

has subdued sculpture, like that shown on plate 58, 

figures 9, 10, whereas plate 58, figures 11, 12, shows 

strong sculpture. Other shells, however, are more or less 

intermediate. The suppression of axial ribs on the 

lower part of the anal fasciole of specimens that have 

subdued sculpture produces the effect of a wide fasciole.

This slender species is more similar to C. haliostrephis (Dall) (1889, p. 86, pi. 13, fig. 3), dredged 

northwest of Alacran Eeef off Yucatan at a depth of 

84 fathoms, than to any of the described fossil species. 

The primary spiral threads of the living species are 

more delicate. The protoconch of the type (the only 

available specimen) is blunt and paucigyrate: 1% 

whorls. It shows a suggestion of a few axial riblets, 

but is worn. The primary spiral threads of C. felis 

Olsson (1964, p. 102, pi. 19, fig. 6), a middle to late 

Miocene, or early Pliocene, species from Ecuador, are

more widely spaced and fewer (12 on body whorl), 

except on the penult whorl.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, locality 138a. Middle part, eastern area, localities 139b, 

139e, 146, 147b, 147f, 147g, 155c, 157, 159d. Upper 

part, eastern area, localities 173, 175.

Carinodrillia (Carinodrillia) cf. C. elocata 

(Pilsbry and Johnson)

Plate 58, figures 13, 14

Moderately large, moderately slender, pillar moderately long, anal fasciole constricted. Protoconch and 

early post-protoconch whorls missing. Earliest preserved whorls partly corroded. Axial ribs wide, swollen; 6 on penult and body whorls. A sutural primary 

spiral thread and 4 others on penult whorl; 19 others 

on body whorl, one of which is replaced by 2 weaker 

threads. A strong secondary spiral thread at lower edge 

of anal fasciole. Submicroscopic secondary spiral 

threads on noncorroded whorls; those on late half of 

body whorl strengthened. Exaggerated growth threads 

here and there, especially on late half of body whorl. 

Last axial rib widened to form a hump. Outer lip 

broken back. Upper limb of anal sinus heavily bordered. Aperture moderately long.

Height (almost complete) 26.8 mm, diameter (including hump) 9.8 mm (figured specimen).

One specimen of this species is in a collection from 

the middle part of the Gatun formation. It is more 

slender than Carinodrillia elocata (Pilsbry, 1922, p. 

318, pi. 16, fig. 9; Gurabo formation, Dominican Eepublic) and has stronger secondary spiral sculpture.

It has been claimed that C. winchesterce Pilsbry 

(1922, p. 318, pi. 16, figs. 7, 8), also from the Dominican Eepublic, is an unusually slender, long-canaled 

form of C. elocata (Woodring, 1928, p. 155), but it 

doubtless is a distinct species, as Pilsbry thought.

Occurrence: Middle part of Gatun formation (middle Miocene), eastern area, locality 139c.

Carinodrillia (Oarinodrillia) species 

Plate 63, figure 5

Moderately large, moderately slender, pillar moderately long, anal fasciole strongly constricted. Protoconch and early post-protoconch whorls missing. Axial 

ribs moderately wide, swollen; 8 on penult and body 

whorls. Sutural primary spiral thread closely hugging 

suture. Four other primary spiral threads on penult 

whorl; 11 on body whorl. Submicroscopic secondary 

spiral threads on noncorroded parts; those on anal 

fasciole fainter than elsewhere. A strong secondary 

thread at lower edge of anal fasciole. Last axial rib
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 377

higher and wider than others, forming a narrow hump. 

Outer lip broken back. Upper limb of anal sinus 

heavily bordered. Aperture moderately long.

Height (almost complete) 27.6 mm, diameter (including hump) 10.3 mm.

An unnamed clavine turrid from the Chagres sandstone is represented by an imperfect specimen. The 

entire apertural face of the shell and part of the adapertural face are corroded. The relatively narrow 

axial ribs and strongly constricted, weakly sculptured 

anal fascicle are distinctive. This species evidently is 

new and has no known close allies.

Occurrence: Chagres sandstone (late Miocene), 

locality 208.

Genus Darbya Bartsch?

Bartsch, Smithsonian Misc. Colln., v. 91, no. 2, p. 22, 1934. 

Type (orthotype): Darbya lira Bartsch, living off north coast 

of Puerto Rico.

Submenus Buridrillia Olsson?

Olsson, Bull. Am. Paleontology, v. 27, no. 106, p. 51, 1942.

Type (orthotype): ClatUrodrillla panarica Olsson, Pliocene, 

Pacific coast of Central America straddling Costa RicaPanama boundary.

Though the protoconch of Buridrillia is unknown 

and Darbya is so far represented by only the type 

species, Powell (1966, p. 63) was justified in pointing 

out their close alliance and possible identity. As matters now stand, the species of Buridrillia are moderately large and inflated, whereas the single species of 

Darbya is small and slender. Not all species of Buridrillia have a columellar swelling and only one of the 

five specimens of Darbya lira shows that feature. This 

peculiar swelling is hardly comparable to the columellar plaits of the borsonines and does not belie the 

clavine features.

A worn corroded shell, dredged at a depth of 134 

fathoms near Cocos Island (USNM 106888), is indistinguishable from the type species of Buridrillia, so 

far as it goes. Cocos Island is off Coiba Island, in the 

Pacific coastal waters of Panama.

Darbya? (Buridrillia?) species 

Plate 63, figures 6, 7

Small, moderately slender, whorls strongly inflated, 

pillar of moderate length. Protoconch and early postprotoconch whorls missing. Anal fascicle constricted. 

Axial ribs wide and widely spaced, attenuated and 

bent on anal fascicle, 10 on penult whorl. Spiral sculpture consisting of narrow bands, separated by grooves, 

in which microscopic axial threads are conspicuous. 

Spiral bands closely spaced on anal fascicle. A hump 

on body whorl near aperture. Outer lip broken back

to hump. Aperture and siphonal canal of moderate 

length. As shown by growth lines, anal sinus moderately deep; border of upper limb thin. Siphonal fascicle sculptured with closely spaced spiral threads.

Height (incomplete) 15 mm, diameter 6 mm.

A second unnamed clavine species from the Chagres 

sandstone, like the preceding species, is represented by 

an imperfect shell. The outer shell layer and therefore 

the spiral sculpture are corroded on the apertural face 

of the shell, except on the lower part of the body whorl. 

The outline, moderately long siphonal canal and other 

apertural features, so far as they are preserved, suggest 

a small species of Buridrillia that lacks a columellar 

swelling. The fossil species of the subgenus so far 

described are found in formations of late Miocene or 

Pliocene age in Ecuador (Olsson, 1964, p. 98-100) and 

on the Pacific coast of Central America straddling the 

Costa Rica-Panama boundary (Olsson, 1942, p. 51-53). 

The spiral sculpture of the Chagres species is stronger 

than that of the described species.

Occurrence: Chagres sandstone (late Miocene), 

locality 208.

Genus Crassispira Swainson

Swainson, A treatise on malacology, p. 313, 1840.

Type (logotype, Herrmannsen, Indicis generum malacozoorum, 

v. 1, p. 318, 1847): Pleurotoma Notice Valenciennes in 

Kiener (cited by Swainson as Pleurotoma Bottce Auct), 

living, Gulf of California.

Subgenus Crassispira s.s.

Crassispira henekeni has a narrower aperture and 

less inflated siphonal fascicle than the type species, as 

well as fewer and wider axial ribs. In number and 

width of ribs C. cegis (Woodring, 1928, p. 151, pi. 4, 

fig. 12; Bow den formation, Jamaica) is more similar 

to the type species, but, like C. henekeni, has a narrower aperture.

Crassispira (Crassispira) henekeni leptalea Woodring, n. subsp. 

Plate 58, figure 19; plate 65, figure 5

Of medium size, slender, aperture long and narrow. 

Protoconch acute, conical, 3-whorled; end marked by 

faint arcuate axial riblet. Sculpture consisting of 

slightly protractive, swollen axial ribs (8 on body 

whorl), overridden by strong, narrow primary spiral 

threads (7 or 8 on penult whorl). Subsutural thread 

strong; anal fascicle below subsutural thread bearing 

faint secondary spiral threads. Outer lip broken back. 

Anal sinus wide, apex below subsutural thread. Siphonal fascicle scarcely inflated.

Height (practically complete) 30.6 mm, diameter 

9.7 mm (type). Height (almost complete) 34 mm, 

diameter 10.5 mm.
378 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Type: USNM 645777.

Type locality: 147h (USGS 6442, i/2 mile (750 meters) south [southeast] of Gatun, Canal Zone), middle 

part of Gatun formation.

This subspecies, which is smaller and more slender 

than the nominate subspecies, and has more sharply 

chiseled spiral sculpture, is rare. Only two specimens, 

both from the middle part of the Gatun formation, 

have been found. It is the first record of the species 

along the west border of the Miocene Caribbean Sea. 

The nominate subspecies (Sowerby, 1850, p. 50, pi. 10, 

fig. 6) occurs in the Cercado and Gurabo formations 

of the Dominican Republic, but is much more abundant 

in the Gurabo, and also in middle Miocene deposits in 

Trinidad, Venezuela, and Colombia. The blunt, cylindrical protoconch is not thought to belie the affinities 

of the Panama form.

Crassispira inoequistriata (Li) (Pilsbry, 1931, p. 

437, pi. 41, fig. 2), living in Panama Bay, is the most 

similar living species. No comparable species is known 

in the western Atlantic Ocean. The deep-water western 

Atlantic species Pleurotoma (Drillia} horrenda Watson (1886, p. 308, pi. 26, fig. 4) bears a superficial 

resemblance. Its aperture, however, is wide and its 

outer lip strongly flaring.

Occurrence: Middle part of Gatun formation, eastern area (middle Miocene), localities 147h, 157.

Subgenus Hindsielava Hertlein and Strong

Hertlein and Strong, Am. Mus. Nat. History Bull., v. 107, p.

227, 1955. 

Type (orthotype): Pleurotoma militaris Hinds in Reeve, living,

Baja California to Colombia.

The more slender outline, more numerous and narrower axial ribs, stronger spiral sculpture, narrower 

aperture, and less inflated siphonal fascicle distinguish 

Hindsiclava from Grassispira s.s.

The Oligocene species Pleurotoma abundans Conrad 

(1848, p. 115, pi. 11, fig. 25; Byram marl, Mississippi) 

is the earliest Hindsiclava now known. Conrad's illustration, however, leaves much to be desired.

Crassispira (Hindsiclava) censors censors (Sowerby) 

Plate 58, figures 1, 22; plate 65, figure 6

Pleurotoma censors Sowerby, Geol. Soc. London Quart. Jour., 

v. 6, p. 50, 1850 (Miocene, Dominican Republic). Guppy, 

Idem, v. 32, p. 527, pi. 28, fig. 7, 1876 (Miocene, Dominican Republic).

Drillia censors (Sowerby), Brown and Pilsbry, Acad. Nat. Sci. 

Phila. Proc., v. 63, p. 345, 1911 (Miocene, Canal Zone). 

Maury, Bull. Am Paleontology, v. 5, no. 29, p. 53, pi. 8, 

figs. 15, 16, 1917 (Miocene, Dominican Republic). Maury, 

N.Y. Acad. Sci., Scientific Survey of Porto Rico and Virgin Islands, v. 3, pt. 1, p. 71,1920 (Miocene, Puerto Rico). 

Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 73, p. 319, pi. 16,

fig. 3, 1922 (Miocene, Dominican Republic). Olsson, Bull. 

Am. Paleontology, v. 9, no. 39, p. 61, pi. 4, figs. 8, 10, 13, 

1922 (Miocene, Costa Rica, Canal Zone, Panama). Palmer, Idem, v. 10, no. 40, p. 11, pi. 2, figs. 7, 8, 1923 (Miocene, Costa Rica, Dominican Republic). Maury, Idem, 

v. 10, no. 42, p. 190, pi. 32, fig. 10, 1925 (Miocene, Trinidad). Maury, Brasil Servigo Geol. Mineral. Mon. 4, p. 

203, pi. 12, fig. 5, 1925 (Miocene, Dominican Republic).

Drillia censors (Guppy), Cossmann, Jour. Conchyliologie, v. 61, 

p. 20, pi. 2, figs. 8-14, 1913 (Miocene, Martinique, Dominican Republic, Canal Zone).

Turris (Crassispira) censors (Sowerby), Rutsch, Schweizer 

Palseont. Gesell. Abh., v. 55, p. 99, pi. 8, figs. 13-16, 1934 

(Miocene, Venezuela).

Crassispira censors (Sowerby), Pflug, Acta Humboldtiana, Geol. 

Palseontologica ser., no. 1, p. 67, pi. 19, figs. 4, 7, 10, 1961 

(Miocene, Dominican Republic).

Crassispira aff. censors (Sowerby), Jung, Bull. Am. Paleontology, v. 49, no. 223, p. 565, pi. 76, figs. 14, 15, 1965 (Miocene, Venezuela).

1'Crassispira? cf. Crassispira? consors (Guppy), Marks, Idem, 

v. 33, no. 139, p. 135, 1951 (Miocene, Ecuador).

Wrillia consors portoriccensis Hubbard, N.Y. Acad. Sci., Scientific Survey of Porto Rico and Virgin Islands, v. 3, pt. 2, 

p. 158, pi. 24, figs. 8, 9, 1920 (Miocene, Puerto Rico; not 

Drillia portoricosnsis Hubbard, Idem, p. 159, pi. 24, figs. 

11, 12).

Wrillia consors fittW&roofci Mansfield, U.S. Natl. Mus. Proc., v. 

66, p. 16, pi. 3, fig. 10, 1925 (Miocene, Trinidad).

Wrillia consors trinitatensis Mansfield, Idem, p. 17, pi. 3, figs. 

12, 13, 1925 (Miocene, Trinidad).

Turris (Drillia) militaris (Hinds), Gabb, Am. Philos. Soc. 

Trans., n. ser., v. 15, p. 207, 1873 (Miocene, Dominican 

Republic). Gabb, Acad. Nat. Sci. Phila. Jour., 2d ser., 

v. 8, p. 350, 1881 (Pliocene, Costa Rica).

Pleurotoma sp. aff. P. alesidota macilenta Dall, Toula, K. k. 

Geol. Reichsanstalt Jahrb., v. 61, p. 506, pi. 30, fig. 11, 

1911 (Miocene, Canal Zone).

Pleurotoma (Drillia) Dalli Toula, Idem, p. 506, pi. 30, fig. 12, 

1911 (Miocene, Canal Zone).

Not Pleurotoma dalli Verrill and Smith, Connecticut Acad. 

Trans., v. 5, p. 451, pi. 57, figs. 1, la, 1882.

Drillia macilenta rectaxis Pilsbry, Acad. Nat. Sci. Phila. Proc., 

v. 73, p. 319, pi. 16, figs. 1, 2, 1922 (Miocene, Dominican 

Republic).

Surcula Tw&soni Hanna, California Acad. Sci. Proc., 4th ser., 

v. 13, no. 10, p. 181, 1924 (substitute name for Pleurotoma dalli Toula).

Moderately large, slender, aperture long and narrow. Protoconch acute (apex almost invariably broken), 21/2- to 3-whorled, last half whorl slightly or 

decidedly bulging, end marked by appearance of first 

axial rib. First 3 to 5 axial ribs narrow, arcuate, extending practically from suture to suture. Later ribs 

wider, slightly protractive, cut off at anal fascicle. 

Ribs progressively narrower and more closely spaced. 

Sculpture of late whorls consisting of skewed rectangles formed by strong, slightly protractive axial ribs 

(22 to 30 on mature penult whorl) and narrower primary spiral threads (4 or 5 on mature penult whorl),
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 379

slightly swollen on crest of ribs. Microscopic spiral 

threads, crenulated by microscopic growth threads, between primary threads. Anal fasciole dominated by 

strong subsutural thread, flanked on lower side by 

secondary and microscopic spiral threads and somewhat exaggerated growth threads, and on upper side 

by microscopic spiral threads. Subsutural thread and 

space between it and suture slightly undulated by 

swellings corresponding in position to axial ribs. Anal 

sinus wide, moderately deep, apex on lower part of 

fasciole.

Height (not quite complete) 42.9 mm, diameter 12 

mm (larger figured specimen).

Type: British Museum (Natural History) G83972.

Type locality: Valley of Rio Yaque del Norte, Dominican Republic, presumably Gurabo formation.

This moderately large turrid is widespread in the 

Gatun formation. The largest number of specimens 

were found at localities 138c, 155c, and 175: 15, 20, 

and 11, respectively. The largest, an incomplete shell 

from locality 138c, would have a height of about 50 

mm, if it were complete, comparable to the height of 

a large shell from the upper part of the Gatun illustrated by Cossmann (1913, pi. 2, figs. 13, 14). The 

number and spacing of the axial ribs and the number 

and strength of the secondary spiral threads on the 

anal fasciole show minor variation. The subsutural 

thread of Toula's Pleurotoma aff. P. alesidota maciJenta is split and the lower part itself is doubled. The 

type of his Pleurotoma dalli is an immature shell 

(height 26.6 mm).

Crassispira consors and the nominate subspecies have 

been found in many areas in the Miocene and Pliocene 

Caribbean province. Sowerby's specimens presumably 

were collected from the Gurabo formation. A lectotype has been designated and illustrated recently by 

Pflug. Slender specimens in Gabb's Dominican Republic collection were named Drillia macilenta rectaxis by 

Pilsbry. C. consors consors reaches a height of about 

75 mm in the middle Miocene deposits of southeastern 

Costa Rica. The axial ribs on the last two whorls of 

that large specimen and another almost as large, illustrated by Olsson (1922, pi. 4, fig. 8; height 60 mm), 

are narrow and very closely spaced.

All of the some 100 specimens from the Gatun formation that are not worn or corroded show, between 

the primary spiral threads, microscopic spiral threads 

crimped by microscopic growth threads. With few 

known exceptions, shells from the type region and 

many other localities have microscopic growth threads, 

but lack microscopic spiral threads. A late Miocene 

specimen, collected on a cay off Bluefields Point at 

the west end of Valiente Peninsula (in the Bocas del

Toro area of northwestern Panama), has microscopic 

spiral threads, whereas they are absent on two of the 

same age collected on Swan Cay, off the west end of 

Bocas Island, 50 km northwest of Bluefields Point. 

The last two or three whorls of C. consors bullbrooki 

and C. consors trinitatensis, both from the middle 

Miocene part of the Brasso formation of Trinidad, 

bear one to three microscopic spiral threads, like those 

of the Panama fossils, or a little stronger. Both subspecies probably are to be suppressed. All the specimens are small, doubtless immature, and C. consors 

tnnitatensis was based on a stream-float collection. Elsewhere the middle Miocene part of the Brasso formation contains moderately large specimens that show 

faint microscopic threads (USGS 21234,21240). Should 

it be considered appropriate to name the Gatun form, 

C. consors hobsoni, which was used by Gardner (1926- 

47, p. 300, 1938) is available. The name Surcula hobsoni was proposed in a wholesale naming of junior 

homonyms.

The earliest Caribbean fossils of the C. consors lineage are of early Miocene age. Those of that age from 

Haiti, the Dominican Republic, and Venezuela are like 

small replicas of middle Miocene specimens (height 15 

to 25 mm). The Brazilian C. consors pennce (Maury), 

(1925a, p. 205, pi. 12, figs. 1, 2, 4, 6), however, is large 

(height 40 mm) and has more ribs than C. consors 

consors. The slightly arcuate ribs shown in Maury's 

illustrations presumably are due to inaccurate drawing. 

Pleurotoma (Drillia) alesidota magna Bose (1906, p. 

47, pi. 5, figs. 30, 31, 33, 45) may be treated as a middle Miocene Mexican subspecies of D. consors that has 

spiral bands instead of narrower threads.

The status of the lineage in the Miocene of Florida 

has not yet been carefully evaluated: Drillia eupora 

Dall (1915, p. 42, pi. 5, fig. 3), Crassispira prceconsors 

Gardner (1926-47, p. 299, pi. 39, fig. 3, 1938), C. calligonoides Gardner (Idem, p. 300, pi. 39, figs. 4, 9), 

C. Uountensis Mansfield (1935, p. 22, pi. 1, fig. 11), 

and C. antealesidota Mansfield (1930, p. 39, pi. 2, fig. 

10). At least some of the Florida fossils have a shorter, 

wider aperture and more inflated siphonal fasciole 

than C. consors. This tendency culminates in the unillustrated Pliocene species C. perspirata (Dall) (1890- 

1903, p. 31, 1890).

C. consors left descendants on both sides of Central 

America: C. alesidota (Dall) (1889, p. 84) and C. macilenta (Dall) (Idem, p. 85, pi. 36, fig. 1) in western 

Atlantic waters, and C. militaris (Hinds in Reeve) 

(1843, species 55, pi. 7, fig. 55), the type of Hindsiclava, 

in eastern Pacific waters. C. alesidota ranges from 

Cape Hatter as to southern Florida at depths of 35 to 

110 fathoms. It has a somewhat less constricted anal
380 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

fascicle than C. consors; the subsutural spiral thread 

is weaker; and that thread and the space between it 

and the suture bear stronger undulations. The protoconch is blunt and 2- to 21/4-whorled. The type of C. 

macilenta, dredged in the northeastern Gulf of Mexico 

(depth 111 fathoms) is the only specimen of that species in the collections of the U.S. National Museum. 

It is more slender than C. consors and C. alesidota, 

and has a less constricted anal fascicle than either of 

those species. The secondary threads on the anal fasciole and the swellings on the axial ribs are strong. The 

protoconch is like that of C. alesidota. A specimen of 

C. alesidota, dredged off Cape Hatteras, was identified 

by Dall as C. macilenta. It remains to be seen whether 

Hindsiclava actually has withdrawn from the Caribbean Sea since Pliocene time. C. militaris has strong 

swellings on the axial ribs and fairly strong undulations on the subsutural spiral thread. It occurs in the 

Esmeraldas formation, of late Miocene age, of Ecuador 

(Olsson, 1964, p. 97, pi. 17, figs. 3, 3a, 3b).

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 138a, 138c, 138d, 138e, 138f. Middle part, eastern area, localities 139c, 139e, 146, 147b, 147f, 147g, 

147h, 151, 155, 155a, 155b, 155c, 156, 157, 159d; western 

area, locality 161a. Upper part, eastern area, localities 

172, 173, 175, 176, I76a, l77b; western area, locality 

182a. Thomonde formation (early Miocene), Haiti. 

Baitoa formation (early Miocene), Dominican Eepublic. Aymamon limestone (early Miocene), Puerto Eico. 

La Eosa formation (early Miocene), Venezuela. Early 

Miocene part of Uscari shale, Costa Eica. Gurabo formation (middle Miocene), Dominican Eepublic. Middle Miocene deposits, Martinique, Trinidad, Venezuela, 

Costa Eica, Colombia, Darien area, Panama (USGS 

8477). Punta Gavilan formation (late Miocene), Venezuela. Limon formation (late Miocene), Costa Eica, 

and Bocas del Toro area, Panama. Deposits of late 

Miocene age, Tehuantepec are, Mexico. Pliocene deposits, Cumana area, Venezuela, Costa Rica.

Crassispira (Hindsiclava) pyrgoma Woodring, n. sp. 

Plate 62, figures 21, 27

Of medium size, slender, distinctly turreted, aperture long and narrow. Protoconch and early postprotoconch whorls missing. Sculpture consisting of 

slightly protractive axial ribs (15 or 16 on penult 

whorl) and low primary spiral bands (4 or 5 on penult whorl), swollen on crest of ribs. Secondary threads 

of varying width between primary bands, some almost 

as wide as the bands, others microscopic. Anal fascicle 

wide, strongly constricted. Subsutural thread strong,

undulated secondary spiral threads between it and 

base of fascicle. Anal sinus wide, shallow, apex on 

lower part of fascicle.

Height (incomplete) 35.5 mm, diameter 10.5 mm 

(type).

Type: USNM 645867. Paratype, USNM 645866.

Type locality: 208 (USGS 8437, Caribbean coast at 

mouth of Eio Indio, Panama), Chagres sandstone.

Though the type and paratype the only specimens 

 are incomplete and partly worn and corroded, this 

species is named, as the turreted outline, and wide and 

strongly constricted anal fascicle are distinctive. In 

general the sculpture resembles that of C. consors 

magna, mentioned under that species.

C. pyrgoma doubtless is an offshoot from the lineage of C. consors. A middle Miocene Mexican turrid, o '

from the Sayula district, Chiapas (USGS 5886), has 

a similar slender, turreted outline, a few less ribs (13 

on penult), and somewhat weaker primary spiral 

sculpture.

Occurrence: Chagres sandstone (late Miocene), 

locality 208.

Subgenus Crassispirella Bartsch and Rehder

Bartsch and Rehder, U.S. Natl. Mus. Proc., v. 87, p. 135, 1939.

Type (orthotype) : Crassispira (Crassispirella) rugitecta (Dall) 

[Turris (Crassispira) rugitecta Dall], living, Baja California.

Crassispirella is adopted as the name for species of 

Crassispira that have a short aperture. They generally 

are small or of medium size. The type species, however, is relatively large (height 30 mm) and its subsutural spiral thread is only slightly wider and higher 

than other threads on the anal fascicle.

The earliest species are found in formations of early 

Miocene age: the Tampa formation of Florida and the 

Thomonde formation of Haiti. The subgenus now lives 

in both eastern Atlantic and western Pacific waters, 

but is represented by far more species in the eastern 

Pacific Ocean than in the Atlantic.

Crassispira (Crassispirella) species

The La Boca formation yielded a small incomplete 

Crassispirella and a minute shell that evidently is a 

tip of the same species. The axial ribs are closely 

spaced (about 15 on last whorl). The subsutural thread 

is strong, undulated, and occupies most of the anal 

fascicle. The spiral sculpture below the anal fascicle 

is weak. The height, almost complete, is 8.8 mm and 

the diameter 3.6 mm.

Occurrence: La Boca formation (early Miocene}* 

locality 116a.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 381

Crassispira (Crassispirella) cymation Woodring, n. sp. 

Plate 58, figures 4, 5

Small, rapidly enlarging. Protoconch acute, 3- to 

3i^-whorled, a wide axial swelling preceding end, 

marked by an arcuate riblet. Sculpture consisting of 

slightly protractive, inverted V-shaped axial ribs, 

swollen at apex (10 on body whorl) and low, narrow 

primary spiral threads (8 or 9 on penult whorl), 

roughened by microscopic growth threads. Spiral 

threads subdued or disappearing on swollen apex of 

ribs; those on lower part of body whorl above siphonal 

fascicle widely spaced. Anal fascicle undulated; subsutural thread strong; fascicle below subsutural thread 

bearing subdued secondary spiral threads. Rib adjoining outer lip varicose. Anal sinus wide, upper 

border thickened on parietal wall. Stromboid notch 

on lower part of outer lip very shallow. Siphonal fasciole slightly inflated.

Height (not quite complete) 8 mm, diameter (incomplete) 3.3 mm (type). Height (incomplete) 7.5 

mm, diameter 4 mm (paratype).

Type: USNM 645781. Paratype, USNM 645782.

Type locality: 161c (USGS 8382, railroad cuts west 

of Gatun Dam, station B, Canal Zone), middle part 

of Gatun formation.

The undulated anal fascicle, including the subsutural 

thread, is a distinguishing feature of this small species, 

which is represented by three specimens, all found in 

the middle part of the Gatun formation in the western 

area. In sculptural plan it is not comparable to other 

small species from the Caribbean region: Orassispira 

maonisripantm (Maury) (1917, p. 56, pi. 9, fig. 5; 

Cercado formation, Dominican Republic), C. hispaniolce (Maury) (1917, p. 56, pi. 9, fig. 6; Gurabo formation, Dominican Republic), C. ritanida (Mansfield) 

(1925, p. 24, pi. 4, fig. 10; Matura formation, Trinidad), and C. annella Woodring (1928, p. 151, pi. 4, 

fig. 11; Bowden formation, Jamaica).

Occurrence: Middle part of Gatun formation (middle Miocene), western area, localities 161c, I70a.

Crassispira (Crassispirella) tyloessa Woodring, n. sp. 

Plate 58, figures 2, 3

Of medium size, subsutural spiral thread noded. 

Protoconch acute, about 2i/£>-whorled (apex worn), last 

whorl bulging below middle, apparently merging into 

first post-protoconch whorl. Axial ribs narrow, 20 on 

penult whorl. Subsutural spiral thread bearing nodes 

alined with axial ribs. Spiral threads below anal fasciole narrow, relatively widely spaced, slightly noding 

crest of ribs; 5 on penult whorl. Anal fascicle below 

subsutural thread bearing faint microscopic spiral

threads. A varicose hump on body whorl near outer 

lip. Axial ribs subdued and closely spaced on hump 

and between it and outer lip. Outer lip broken back. 

Growth lines showing a deep anal sinus and shallow 

stromboid notch. Upper border of anal sinus thickened. Siphonal fascicle slightly inflated, sculptured 

with spiral threads cruder than those elsewhere.

Height 13.4 mm, diameter (including hump) 5.2 mm 

(type). Height 15.3 mm, diameter 5.8 mm (paratype).

Type: USNM 645783; paratype USNM 645784.

Type locality: 139c (USGS 22018, east side of road 

leading from Transisthmian Highway to refinery site 

011 Payardi Island, Panama; about 100 meters southwest of refinery gate),middle part of Gatun formation.

The combination of noded subsutural spiral thread 

and relatively widely spaced other spiral threads, not 

observed on other species of Crassispirella, is distinctive. The paratype is slightly larger than the type, but 

its outer lip is broken back beyond the hump. No other 

specimens are in the collections.

It is strange that no larger species of Crassispirella, 

similar to Crassispira jamaicensis (Guppy) (Woodring, 

1928, p. 149, pi. 4, fig. 8) and its fossil and living 

allies, has so far been found in the Gatun formation.

Occurrence: Middle part of Gatun formation (middle Miocene), eastern area, locality 139c.

Genus Clathrodrillia Hall

Dall, U.S. Natl. Mus. Proc., v. 54, p. 317, 323, 1918.

Type (orthotype, p. 323): Pleurotoma gibbosa Reeve (Pleurotoma gibbosa Kiener = Murex gibbosa Born), living,

south border of Caribbean Sea.

A fragment of two incomplete whorls from the La 

Boca formation is listed as Clathrodrillia sp.

Clathrodrillia gatunensis (Toula) 

Plate 58, figures 8, 20, 21, 23-25; plate 65, figure 7

Pleurotoma (Drillia) gatunensis Toula, K. k. Geol. Reichsanstalt Jahrb., v. 58, p. 707, pi. 25, fig. 16, 1909 (Miocene, 

Canal Zone). Engerrand and TJrbina, Soc. Geol. Mexicana 

Bol., v. 6, pi. 59, fig. 27, 1910 (reproduction of Toula's 

illustration).

Pleurotoma (Drillia) gatunensis Toula, n. var., Toula, K. k. 

Geol. Reichsanstalt Jahrb., v. 61, p. 507, pi. 31, fig. 20, 

1911 (Miocene, Canal Zone).

Drillia, gatunensis (Toula), Brown and Pilsbry, Acad. Nat. Sci. 

Phila. Proc., v. 63, p. 344, 1911 (Miocene, Canal Zone). 

Pilsbry and Brown, Idem, v. 69, p. 32 (list), 1917 (Miocene, Colombia). Barrios, Colombia Servicio Geol. Nac., 

Bol. Geol., v. 6, nos. 1-3 (Informe 1082), p. 292, 1960 

(Miocene, Colombia).

Drillia venusta (Sowerby), Olsson, Bull. Am. Paleontology, 

v. 9, no. 39, p. 61, pi. 4, fig. 9, 1922 (Miocene, Costa Rica).

Drillia puertocolombiana Weisbord, Idem, v. 14, no. 54, p. 55, 

pi. 5, fig. 16, 1929 (Miocene, Colombia).
382 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Clathrodrillia puertocolomMana (Weisbord), Jung, Idem, v. 49, 

no. 223, p. 566, pi. 76, figs. 12, 13, 19, 1965 (Miocene, 

Venezuela).

Clathrodrillia onzola Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst, p. 96, pi. 15, 

figs. 6, 6a, 1964.

Moderately large, moderately slender, pillar strongly 

constricted, aperture relatively short and wide. Protoconch blunt, almost cylindrical, 1% to 4-whorled, generally 1% to 2; lower part of last half whorl faintly 

to distinctly angulated. Lower part of first post-protoconch whorl or two angulated; first whorl sculptured 

with 7 or 8 axial ribs, swollen on angulation, gradually pinched off near suture. Faint spiral sculpture 

appearing on second or third post-protoconch whorl; 

anal fascicle on about third. At that and later stages 

axial ribs slightly protractive, moderately narrow, 

separated by spaces greater than their width, disappearing on body whorl below periphery; 24 to 26 

on penult whorl. Spiral sculpture below anal fascicle 

consisting of narrow bands (4 to 7 on penult whorl), 

which widen on late whorls of a few specimens and 

are separated by narrow grooves. Secondary spiral 

threads between bands on a few specimens. Anal fasciole strongly constricted. Subsutural cord of variable 

strength and width. Wide subsutural cord strongly 

wrinkled by swellings corresponding to axial ribs 

and also swellings intermediate in position with reference to ribs. Narrow subsutural cord faintly wrinkled. On shells that have narrow subsutural cord, concave lower part of fascicle generally steeply sloping; 

rarely gently sloping, producing distinctly turreted 

outline. Concave lower part of fasciole bearing exaggerated growth threads, some as strong as wrinkles 

on subsutural cord. Faint spiral sculpture visible on 

entire fasciole of some shells, especially on early 

whorls. Pillar sculptured with widely spaced spiral 

cords and faint secondary spiral threads. A massive 

hump at varying distance from outer lip; ribs subdued and closely spaced between hump and outer lip. 

Earlier humps, as many as 10, but generally 1 to 5, on 

spire whorls of some shells. Outer lip flaring forward, 

bearing a deep, wide stromboid notch at top of pillar. 

Anal sinus deep, lower limb long, upper limb bordered by thick pad of callus on parietal wall. Siphonal notch wide and deep. Siphonal fasciole slightly 

inflated, bearing no spiral sculpture or weak spiral 

sculpture, generally limited by a strong, narrow 

thread, especially on mature shells.

Height (almost complete) 52.4 mm, diameter (including hump) 17.5 mm (largest figured specimen). 

Height (almost complete) 40.5 mm, diameter (including hump) 12.5 mm (figured specimen of intermediate size).

Type: Tech. Hochschule, Vienna.

Type locality: Presumably Gatun Locks excavation, 

Canal Zone, middle part of Gatun formation.

The 76 specimens of Clathrodrillia from the Gatun 

formation (lower and middle parts, and upper part in 

eastern area) closely related to C. gibbosa are identified as C. gatunensis. Otherwise three names, presumably subspecific names, would be needed and some 

specimens that show intermediate features would have 

to be placed doubtfully under one of the three names. 

The three main forms are as follows:

Forms of Clathrodrillia gatunensis in Gatun 

formation

1. Subsutural cord narrow, weakly wrinkled.

a. Concave part of anal fasciole below subsutural cord steeply sloping (pi. 58, figs. 

24, 25).

b. Concave part of anal fasciole below subsutural cord gently sloping (pi. 58, fig. 8).

2. Subsutural cord wide, strongly wrinkled (pi. 58, 

figs. 20, 21, 23).

Type la is represented in the lower part of the 

Gatun and in strata near the base of the middle part, 

although some specimens in that part are transitional 

toward type 2. Only one shell, collected at a middle 

Gatun locality in the western area, represents type Ib. 

It is associated with a specimen leaning toward type 

2. The remaining specimens from the middle part that 

are larger than a few whorls and those from the upper 

part, 65 percent of the total number, are unmistakably 

referred to type 2. That type agrees with Toula's type 

specimen: an incomplete shell of six whorls (height 

32.6 mm, diameter 12.3 mm). His Pleurotoma (Drillia) gatunensis n. var. is an immature shell (height 

10.1 mm, diameter 4.1 mm). It shows the 2-whorled, 

almost cylindrical protoconch. Type Ib closely duplicates the Ecuadoreaii fossils named C. onzola.

These Gatun fossils could be further subdivided on 

the basis of outline and spiral sculpture. One of five 

specimens from locality 139c is as slender as Orassispira consors. On a few, like that shown on plate 58, 

figures 20, 21, the spiral cords of late whorls are wide 

and separated by narrow grooves. That specimen is 

one of three that has a perfectly preserved outer lip.

As now identified, Clathrodrillia gatunensis occurs 

in deposits of middle Miocene age in Venezuela, Colombia, Costa Rica, the Canal Zone, the Darien and 

Chiriqui areas of Panama, and Ecuador, and at least 

a related form in the Miocene of Martinique (Cossmann, 1913, p. 22, pi. 2, fig. 7). If the indefinite type 

locality of C. puertocolombiana represents the Las 

Perdices shale, the age range extends back to early
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 383

Miocene. The Venezuelan shells identified as C. puertocolotnbiana have a few less ribs than Gatun shells 

(18 to 20 on penult whorl) and lack a strong thread 

at the margin of the siphonal fascicle. A middle Miocene specimen of C. puertocolombiana from Colombia 

(USGS 11241) has 25 ribs on the penult whorl and a 

strong thread at the margin of the fascicle.

C. venusta (Sowerby) (1850, p. 50, pi. 10, fig. 7), 

which was well illustrated by Sowerby, was the first 

Miocene species of a closely knit group of Clathrodrillia to be described. It occurs in the Gurabo formation 

of the Dominican Republic and is distinguished by 

the absence of a subsutural cord and by the attenuated 

continuation of the axial ribs across the entire anal 

fascicle. A lectotype was designated and illustrated 

by Pflug (1961, p. 65, pi. 19, fig. 9). C. gibbosa, the 

type of Clathrodrillia, survives in the Caribbean Sea. 

Its subsutural cord is slightly raised and slightly 

wrinkled. It is a rare species and for many years its 

habitat was unknown. Only two specimens, both originally in the Casey turrid collection, are in USNM 

collections. It is found as a Pliocene fossil in the Cabo 

Blanco and Cumana areas of Venezuela. C. mareana 

(Weisbord, 1962, p. 437, pi. 41, figs. 16, 17; Pliocene, 

Venezuela) is an immature C. gibbosa.

The earliest of this closely knit group of species in 

Florida, perhaps earlier than any in the Caribbean 

region, occurs in the early Miocene Chipola formation: 

C. aulakoessa (Gardner) (1926-47, p. 310, pi. 40, figs. 

2-4, 1938), a small, slender species that has a moderately constricted anal fascicle. The younger C. subvaricosa (Gardner) (Idem, p. 312, pi. 40, figs. 5, 6; 

Oak Grove sand member of Shoal River formation) 

also has a moderately constricted anal fascicle. The 

fascicle of the still younger C. empera (Gardner) 

(Idem, p. 312, pi. 40, figs. 7, 8; Shoal River formation 

proper), however, is strongly constricted. That species, as Gardner realized, is like a small version of 

type 2 of C, gatunensis, but, at least on intermediate 

whorls, attenuated wrinkles more consistently extend 

across the anal fascicle.

No matter whether the synonymy of C. gatunensis 

is modified, the past and present distribution of C. 

gibbosa and its close allies, like that of Muracypraea, 

Sconsia and Xancus, reverses the general trend. Oose 

allies formerly lived in western Atlantic and eastern 

Pacific waters, but the Caribbean C. gibbosa itself is 

the sole survivor.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, 

localities 138c, 138d, 138f. Middle part, eastern area, 

localities 139b, 139c, 139e, 139f, 147f, 147g, 151, 155, 

155a, 155b, 155c, 156, 157, 159a, 159b, 159d; western

area, locality 161a, Upper part, eastern area, localities 

175, 176. Deposits of early (?) Miocene age, Colombia, 

Ecuador. Deposits of middle Miocene age, Venezuela, 

Colombia, Costa Rica, Darien area, Panama (USGS 

8477), Chiriqui area, Panama (USGS 7955), Ecuador.

Clathrodrillia saavedrai Woodring, n. sp. 

Plate 59, figures 13, 14; plate 65, figure 8

Small, subsutural cord noded, anal fascicle constricted, pillar short. Protoconch blunt, ll^-whorled, 

lower part of last half whorl swollen. First post-protoconch whorl sculptured with 8 axial ribs, swollen on 

lower part, pinched off near suture. At later stages 

axial ribs narrow; 17 or 18 on penult whorl. Subsutural cord strong, bearing elongate retractive nodes, 

alined with axial ribs. A spiral thread marking lower 

edge of anal fascicle. Spiral sculpture below anal fascicle consisting of narrow bands (4 to 6 on penult 

whorl), separated by deep grooves. Axial ribs on late 

part of body whorl slightly noded by spiral bands. A 

hump on body whorl about 45° from outer lip. Axial 

ribs subdued and irregular in spacing between hump 

and outer lip. Anal sinus deep, upper limb heavily 

bordered. Stromboid notch wide and deep. Siphonal 

notch deep. Siphonal fascicle slightly inflated, bearing 

no spiral sculpture, limited by a fairly strong thread.

Height 16.5 mm, diameter 6.2 mm (type).

Type: USNM 645790.

Type locality 147b (USGS 6033c, Panama Railroad, about 3,500 feet (1,065 m) southeast of Gatun 

railroad station, Canal Zone), middle part of Gatun 

formation.

Three specimens of Clathrodrillia saavedrai have 

been found in the middle part of the Gatun formation. 

It suggests a miniature C. gatunensis, but is distinguished from immature specimens of that species by 

its noded subsutural cord, more delicate sculpture, 

and weaker thread limiting the siphonal fascicle. 

Though the outer lip of the type was well preserved, 

it was damaged in handling.

This species is named for Alvaro de Saavedra 

Ceron, who early in the 16th century suggested Central American routes to open the land from sea to 

sea (Howarth, 1966, p. 56-57).

Occurrence: Middle part of Gatun formation, eastern area (middle Miocene), localities 147b, 147h.

Clathrodrillia aff. C. saavedrai Woodring, n. sp. 

Plate 58, figures 6, 7

Resembling ClatJirodriUia saavedrai in general features of protoconch and post-protoconch whorls, but 

distinguished as follows. Smaller and more slender.
384 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Axial ribs narrower (16 on penult whorl), pinched 

on lower part of anal fasciole, but continuing across 

entire fasciole, including subsutural cord. Axial ribs 

011 subsutural cord slightly arcuate on intermediate 

and late whorls. Grooves between spiral bands wider 

than those of C. saavedrai. Siphoiial fasciole sculptured with 2 coarse spiral threads.

Height 10.4 mm, diameter 3.5 mm (figured specimen).

The middle part of the Gatun formation in the 

western area yielded the only specimen of this minute 

Clathrodrillia. Though it evidently represents a new 

species, the outer lip is damaged and the shell is further disfigured by two bore holes, which are large for 

the size of the shell.

Occurrence: Middle part of Gatun formation (middle Miocene), western area, locality 161c.

Clathrodrillia aff. C. lelandi (Olsson) 

Plate 59, figures 5, 6

Of medium size, slender, whorls slightly inflated, 

pillar short. Protoconch and first few post-protoconch 

whorls missing. Anal fasciole slightly constricted on 

earliest preserved whorls, distinctly constricted on 

later whorls. Axial ribs narrow, slightly protractive, 

slightly bent backward and pinched at lower edge of 

anal fasciole, continuing across subsutural cord marking anal fasciole, straight or slightly arcuate on fasciole; 21 on penult whorl. Spiral sculpture below anal 

fasciole consisting of low bands (9 or 10 on penult 

whorl), separated by narrow grooves, faint except on 

body whorl. Faint spiral grooves visible on anal fasciole adjoining anal sinus. A hump on body whorl 

opposite aperture. Axial ribs subdued and closely 

spaced between hump and outer lip. Aperture short. 

Stromboid notch on outer lip shallow. Anal sinus deep, 

upper limb heavily bordered. Siphonal notch deep, 

siphonal fasciole limited by a strong thread.

Height (incomplete) 21.8 mm, diameter 7.5 mm 

(figured specimen).

This slender species is represented by one somewhat 

worn specimen from the lower part of the Gatun 

formation. Owing to wear, the spiral sculpture is faint, 

except on the abapertural face of the body whorl near 

the outer lip. The whorls are less inflated, or shouldered, below the anal fasciole than those of the middle 

Miocene Ecuadoreaii Agladrillia lelandi (Olsson, 1964, 

p. 101, pi. 15, fig. 7) and the spiral bands are slightly 

wider.

It seems to be more appropriate to assign Agladrillia lelandi to Clathrodrillia.

Occurrence: Lower part of Gatun formation (middle Miocene), locality 136a.

Clathrodrillia cf. C. islalindae (Maury) 

Plate 58, figures 15, 16

Small, slender, whorls slightly inflated, pillar short. 

Protoconch and first few post-protoconch whorls missing. Anal fasciole slightly constricted, except on late 

half of body whorl. Axial ribs narrow, slightly protractive, continuing to suture without being pinched 

at lower edge of anal fasciole, arcuate on fasciole; 16 

to 18 on penult whorl. Spiral sculpture below anal 

fasciole consisting of low bands (6 on penult whorl), 

separated by narrow grooves. Faint spiral sculpture 

visible here and there on anal fasciole. A hump on 

body whorl near outer lip. Aperture short. Outer lip 

broken back to hump, except at and near anal fasciole. 

Anal sinus deep, upper limb heavily bordered. End 

of siphonal canal broken. Siphonal fasciole limited by 

a strong, though worn, thread.

Height (not quite complete) 15.8 mm, diameter (including hump) 5.5 mm (figured specimen).

Four more or less worn specimens, also from the 

lower part of the Gatun formation, are identified as 

another species of Clathrodrillia. The illustrated shell 

is the largest and the least worn. In general features 

this species is similar to C. aff. C. lelandi. It is, however, smaller, has less inflated whorls, and its ribs are 

uninterrupted and decidedly arcuate on the anal fasciole. The ribs of C. islalindce (Maury, 1917, p. 57, pi. 

9, fig. 7; Cercado and Gurabo formations, Dominican 

Republic) are similar, but are more protractive, fewer 

in number, and therefore more widely spaced. Moreover, the spiral sculpture of that species is stronger.

Occurrence: Lower part of Gatun formation (middle 

Miocene), localities 138d, 138f.

Clathrodrillia? species 

Plate 63, figures 1, 2

Of medium size, moderately slender, whorls moderately inflated, pillar short. Protoconch and early 

post-protoconch whorls missing. Anal fasciole molded 

against preceding whorl, progressively more strongly 

constricted. Axial ribs wide, bent and attenuated on 

anal fasciole, subdued on anal fasciole of body whorl 

and absent on last quarter of that whorl, 17 on penult 

whorl. Spiral sculpture faint, except on lower twothirds of body whorl, where it consists of narrow, 

closely spaced, crimped threads. A hump on body 

whorl about 60° from aperture. Aperture short. Outer 

lip and lower part of siphonal canal broken back. As 

shown by growth lines, anal sinus deep; upper limb 

heavily bordered. Stromboid notch wide and very 

shallow, as indicated by growth lines. Siphonal fasciole not bordered by a strong thread.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 385

Height (incomplete) 25.9 mm, diameter (including 

hump) 9.7 mm (figured specimen).

The Chagres sandstone yielded an incomplete, corroded clavine turrid of undetermined affinities. The 

anal fasciole, molded against the preceding whorl in 

the fashion of some species of Latirus, is a diagnostic 

feature. The outer shell layer is missing on the crest 

of most body-whorl ribs and on all parts of spire 

whorls, except on the lower part of the penult, where 

faint spiral sculpture is discernible.

Though the apertural features are imperfect, aside 

from the heavy border of the upper limb of the anal 

sinus, this species evidently is not a typical Clathrodrillia.

Occurrence: Chagres sandstone (late Miocene), 

locality 208.

Genus Agladrillia Woodring

Woodring, Carnegie Inst. Washington Pub. 385, p. 157, 1928. 

Type (orthotype): Agladrillia callothyra Woodring, Miocene, 

Jamaica.

Subgenus Agladrillia s.s.

The earliest species of Agladrillia s.s. now known 

occur in the late early Miocene Thomonde formation 

of Haiti. The subgenus survives in the eastern Pacific 

Ocean, but has not been recognized in the western 

Atlantic.

The four species in the Gatun formation, all of 

which are small, have a hump on the body whorl 90° 

or less from the outer lip. Between the hump and the 

outer lip the axial ribs are suppressed, irregularly and 

generally closely spaced, and generally noded by the 

spiral bands, or are absent. As shown by the few 

shells that have an intact outer lip and by growth 

lines on other shells, the anal sinus is deep and the 

stromboid notch shallow, although distinct. The siphonal notch is moderately deep, and the siphonal fasciole 

is slightly inflated and sculptured with relatively 

coarse spiral threads. These nondiagnostic features are 

not repeated in the descriptions of the species.

Agladrillia (Agladrillia) characta Woodring, n. sp. 

Plate 59, figures 3, 4; plate 64, figure 1

Small, moderately inflated, anal fasciole slightly 

constricted, subsutural cord generally faint, rarely 

distinct. Protoconch acute, 1%- to 3-whorled (generally 1% to 2i£), lower part of last whorl swollen. 

Axial ribs pinched off near suture on first, or first two, 

post-protoconch whorls. On remaining whorls axial 

ribs extending across the faint, or distinct, subsutural 

cord, arcuate on the cord; 11 to 15 ribs (generally 12 

to 14) on penult whorl. Spiral sculpture below subsutural cord consisting of bands separated by relatively deep and relatively wide grooves; 5 to 7 bands 

on penult whorl, not including 1 to 3 minor bands on 

or near anal fasciole.

Height 12.8 mm, diameter (including hump) 4.3 

mm (type).

Type: USNM 645794.

Type locality: 147g (TJSGS 5899, highest fossilbearing beds, Quebrancha Hills, % mile (1.2 km) from 

Gatun, Canal Zone), middle part of Gatun formation.

This species is the largest of the four in the Gatun 

formation. The relatively large size and more uniform 

greater inflation distinguish it from the others.

A. characta, represented by 22 specimens, occurs in 

the lower and middle parts of the Gatun formation. 

Three additional specimens from the lower part are 

doubtfully referred to it. They have a blunt protoconch 

of 1% whorls and the lower part of the last whorl 

is subcarinate. The spiral sculpture of two agrees with 

that of A. characta, but the spiral bands of the third 

are narrow. More material is needed to resolve the 

identification of these specimens. They probably represent still another species.

Agladrillia mim.ys Olsson (1964, p. 100, pi. 16, figs. 

6, 6a; Esmeraldas formation, Ecuador) is the most 

similar described fossil species. Its ribs are subdued 

near the suture and its spiral sculpture is weaker than 

that of A. cliamcta. In spiral sculpture the Gatun species resembles A. callothyra (Woodring, 1928, p. 158, 

pi. 5, fig. 7), the type species, but that species is larger 

(height 20 mm) and has a more constricted anal fasciole, on which the axial ribs are subdued. Aside from 

its larger size and fewer, wider ribs, A. callothyra is 

similar to Clathurella panamella Dall (1908, p. 288, 

pi. 14, fig. 1). In Dall's illustration the ribs on the anal 

fasciole are too strong. Powell (1966, p. 92) suspected 

that Dall's species is to be assigned to Agladrillia. It 

has been dredged in the Gulf of California at depths 

of 47 to 153 fathoms.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, locality 138a 

(identification doubtful), 138c, 138d (identification 

doubtful). Middle part, eastern area, localities 139b, 

146, 147b, 147g, 151, 159d.

Agladrillia (Agladrillia) enneacyma enneacyma 

(Brown and Pilsbry)

Plate 59, figures 10, 11, 15, 16; plate 64, figure 4

Very small, slender to moderately inflated, anal fasciole slightly constricted, or not constricted. Subsutural 

cord faint on last whorl or two, or absent on all whorls. 

Protoconch acute, 3- to 3i£-whorled, lower part of 

last whorl swollen, rarely subcarinate. Axial ribs extending to suture, or almost to suture, on first post-
386 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

protoconch whorl. On remaining whorls ribs extending across entire whorl of shells that have no subsutural cord, but gradually bent on anal fasciole, or on 

noncoiistricted part of whorl corresponding to anal 

fasciole, forming elongate nodes on anal fasciole; 9 

to 12 ribs on penult whorl. Spiral sculpture consisting 

of narrow bands separated by shallow grooves; 5 or 

6 bands on penult whorl, not including 1 to 3 narrow 

bands at lower edge of anal fasciole, or on corresponding part of whorls that lack a constricted fasciole.

Height 7.7 mm, diameter (including hump) 2.5 nun 

(figured slender specimen). Height 7.2 mm, diameter 

(including hump) 2.8 mm (figured inflated specimen).

Type: Acad. Nat. Sci. Phila. 3848.

Type locality: Gatim lower locks excavation, Canal 

Zone, middle part of Gatun formation.

The little glistening shells of Agladrillia enneacyma 

enneacyma were abundant in the middle part of the 

Gatun formation about a kilometer southeast of Gatun 

railroad station during the construction period, when 

fresh cuts were accessible on the present line of the 

Panama Railroad. A hundred shells, all of which have 

the protoconch preserved, were collected at locality 

147b, where many other small species were found to 

be abundant, and 45 at locality 147g. Shells that have 

an intact outer lip, however, are rare. The upper part 

of the Gatun yielded only one specimen and its spiral 

sculpture consists of narrow threads instead of bands.

This minute Agladrillia is variable in outline and 

in the presence or absence of a subsutural cord. Aside 

from the single upper Gatun shell, the sculpture is 

basically uniform, although the spiral grooves on 

many well-preserved shells are faint and the crest of 

the axial ribs of the type is exceptionally wide. The 

illustrations on plate 59, representing two specimens 

selected from the large 147b collection, show some of 

the range in variation. The contrast between the unpinched early axial ribs of A. enneacyma enneacyma 

and the pinched ribs of A. characta is shown by figures 

1 and 4 on plate 64.

A larger similar species occurs in the Thomonde 

formation of Haiti and a smaller species in the Cerado 

and Gurabo formations of the Dominican Republic. 

The early axial ribs of both are pinched.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

localities 139b, 139c, 146, 147b, 147f, 147g, 147h, 153a, 

155c, 159d. Upper part, eastern area, locality I73a.

Agladrillia (Agladrillia) enneacyma (Brown and Pilsbry), 

subsp.

Four specimens from the lower part of the Gatun 

formation are distinguished from the nominate subspecies of Agladrillia enneacyma by the presence of a 

subsutural cord on all except the first few post-protoconch whorls and by relatively deep grooves between 

the spiral bands, like those of A. characta. These shells 

have the protoconch and unpinched early axial ribs of 

the nominate subspecies. Two (height 8.7 and 9.5 mm) 

are larger than any shells of the nominate subspecies, 

The sample of this form is inadequate.

Occurrence: Lower part of Gatun formation (middle Miocene), localities 137, 138, 138c.

Agladrillia (Agladrillia) phengoides Woodring, n. sp. 

Plate 59, figures 1, 2; plate 64, figure 3

Very small, slender, anal fasciole constructed, subsutural cord narrow, moderately strong. Protoconch 

blunt, iy2 - to 1%-whorled, lower part of about last 

half whorl subcarinate. Axial ribs pinched off near 

suture on first few post-protoconch whorls. Subsutural 

cord appearing on fourth or fifth post-protoconch 

whorl. On later whorls axial ribs pinched at lower 

edge of subsutural cord, forming small, retractives 

nodes on the cord; 9 or 10 ribs on penult whorl. Spiral 

sculpture below subsutural cord consisting of narrow 

bands separated by shallow grooves; 5 to 7 bands on 

penult whorl, not including 1 or 2 minor bands present 

on some shells near edge of subsutural cord. Grooves 

near subsutural cord distinct, indistinct, or even absent.

Height 6.8 mm, diameter (including hump) 2.2 mm 

(type).

Type: USNM 645799.

Type locality: 147b (USGS 6033c, Panama Railroad, about 3,500 feet (1,065 meters) southeast of 

Gatun railroad station, Canal Zone), middle part of 

Gatun formation.

The slender outline, blunt, paucispiral protoconch, 

and moderately strong, noded subsutural cord on intermediate and late whorls are characteristic features of 

this little glistening species. Unlike Agladrillia enneacyma enneacyma, it is not variable in basic features, 

but the spiral grooves near the subsutural cord may 

be distinct, indistinct, or absent.

A. phengoides was found in the middle part of the 

Gatun formation. The largest number of specimens 

(46) was collected at the type locality, where it is 

associated with A. characta and A. enneacyma enneacyma. It is, in fact, associated with A. enneacyma 

enneacyma at the six localities where it occurs.

The more slender outline, more strongly constricted 

anal fasciole, and more bulging whorls distinguish A. 

phengoides from A. estrellana (Olsson) (1922, p. 69, 

pi. 10, figs. 31, 32), a middle Miocene Costa Rican 

species.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 387

Occurrence: Middle part of Gatun formation (middle Miocene), eastern area, localities 146, 147b, 147f, 

147g, 147h, 159d.

Agladrillia (Agladrillia) acaria ectypha Woodring, n. subsp. 

Plate 59, figures 12, 17; plate 64, figure 8

Very small, moderately slender, anal fasciole not 

constricted, subsutural cord strong. Protoconch blunt, 

1- to 2-whorled, lower part of last half whorl swollen. 

Axial ribs pinched off near suture on first, or first two, 

post-protoconch whorls. Subsutural cord appearing on 

second or third post-protoconch whorl, strong on later 

whorls. Axial ribs wide, swollen, pinched at edge of 

subsutural cord, forming spirally elongate nodes on 

the cord; 9 to 11 ribs on penult whorl. Spiral sculpture 

below subsutural cord consisting of bands separated 

by relatively deep and relatively wide grooves; 4 to 

6 bands on penult whorl, not including 1 or 2 narrow 

bands near edge of subsutural cord.

Height 6.5 mm, diameter 2.3 mm (type).

Type: USNM 645801.

Type locality: 138 (TJSGS 16909, north and south 

sides of Transisthmian Highway, 1.6 kilometers northeast of Canal Zone boundary, Panama), lower part of 

Gatim formation.

Agladrillia acaria ectypha is represented by 60 specimens, 16 of which were collected at locality 138c. All 

the localities are in the lower part of the Gatun formation and in strata near the base of the middle part.

This Agladrillia is distinguished by its wide, swollen 

axial ribs and strong subsutural cord, which appears 

at an early stage on the second or third post-protoconch whorl. The spiral sculpture is like that of A. 

characta on a diminutive scale.

The axial ribs of the Gatun subspecies are wider and 

more swollen than those of the nominate subspecies 

(Mansfield, 1925, p. 18, pi. 1, fig. 2), which occurs in 

the early middle Miocene part of the Brasso formation 

of Trinidad (float from upper part of Globorotalia 

fohsi zone s.l.), and in the late middle Miocene San 

Jose calcareous silt member of the Manzanilla formation. A comparable species from the early Miocene Thomonde formation of Haiti (USGS 9945, 9946) has 

smaller nodes on the subsutural cord and weaker spiral 

sculpture.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 137, 

138, 138a, 138c, 138d, 138e. Middle part, eastern area, 

localities 139b, 139c, 139d, 139e. Middle Miocene deposits, Darien area (USGS 8477).

Subgenus Eumetadrillia Woodring

Woodring, Carnegie Inst. Washington Pub. 385, p. 159, 1928. 

Type (orthotype): Agladrillia (Eumetadrillia) serra Woodring, 

Miocene, Jamaica.

The strong, crosscutting, exaggerated growth threads 

of Eumetadrillia are a conspicuous feature on wellpreserved shells. They are not visible on the illustration of the type species (Woodring, 1928, pi. 5, fig. 9), 

which sho\vs a somewhat worn shell. In different species the protoconch is blunt and paucispiral, or acute 

and multispiral. A subsutural cord is present or absent, and the stromboid notch on the outer lip is shallow or deep.

Eumetadrillia has been found in the Miocene of the 

Dominican Republic, Jamaica, Panama, and Florida. 

It is not known to be living in the western Atlantic or 

the eastern Pacific oceans. A Magellanic species, Pleurotoma (Surcula] fuegiensis Smith (1888, p. 300), has 

been assigned to Eumetadrillia (Powell, 1951, p. 173, 

fig. N, 118). The protoconch has been illustrated, but 

not the shell itself. It would be remarkable for this 

Miocene tropical subgenus to survive in Magellanic 

waters.

Agladrillia (Eumetadrillia) isthmica (Brown and Pilsbry) 

Plate 60, figures 11, 18, 19; plate 64, figure 10

Drillia istJimica Brown and Pilsbry, Acad. Nat. Sci. Phila. Proc., 

v. 63, p. 344, pi. 23, figs. 10, 11, 1911 (Miocene, Canal 

Zone).

Of medium size, moderately slender, anal fasciole 

not constricted, obscured by strong subsutural cord, 

pillar constricted in alinement with stromboid notch 

on outer lip. Protoconch acute, 3- to 3^-whorled (generally 31/2). Subsutural cord appearing on first or second post-protoconch whorl (generally second), undulated by axial ribs of preceding whorl, bearing irregularly spaced, retractive wrinkles. On intermediate and 

late whorls an obscure spiral cord of about same width 

as subsutural cord, for most part represented by a 

groove in interrib spaces, generally adjoining subsutural cord, faintly overriding axial ribs on early part 

of body whorl up to hump. Otherwise spiral sculpture 

absent, except thread limiting siphonal fasciole. On 

intermediate and late whorls axial ribs wide, swollen, 

wide-crested, separated by V-shaped interspaces; 7 to 

10 (generally 8) on penult whorl. Ribs narrower and 

more closely spaced on early part of body whorl up to 

hump. Exaggerated growth threads crosscutting ribs. 

A hump about 90° from outer lip. Axial ribs absent 

between hump and outer lip, except near lip, where 

they are narrow and closely spaced. Ribs continuing 

on pillar as subdued irregular wrinkles and exagger-
388 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

ated growth threads, rarely as regular narrow riblets. 

Anal sinus wide and deep. Stromboid notch narrow 

and deep. Siphonal noM.i deep. Siphonal fascicle not 

inflated, bearing subdued growth threads, limited by 

a narrow spiral thread.

Height 19.9 mm, diameter (incomplete) 6.8 mm 

(larger figured specimen). Height 13.9 mm, diameter 

5.6 mm (smaller figured specimen). Height 21.2 mm, 

diameter (incomplete) 6.4 mm (largest specimen).

Type: Acad. Nat. Sci. Phila. 1696.

Type locality: Gatun Locks excavation, Canal Zone, 

middle part of Gatun formation.

Twenty-three specimens of Agladrillia isthmica from 

the three parts of the Gatun formation in the eastern 

area, are available, not more than six at any of the 

eight localities. All except one (pi. 60, fig. 11) are more 

or less damaged, chiefly through breakage of the outer 

lip far back. A few show no trace of the faint cord 

adjoining the subsutural cord. On the other shells it 

is represented by a short groove in the interrib spaces, 

except on the early part of the body whorl, where it 

faintly overrides the narrow ribs. The constriction of 

the pillar, in alinement with the stromboid notch, in 

Oyma.tosyrinx (p. 374) and some species of Eumetadrillia is correlated with a deep notch.

A. isthmica is closely related to A. bocasensis (Olsson) (1922, p. 73; pi. 5, fig. 5; late Miocene, Bocas del 

Toro area), the only other described species marked by 

a subsutural cord. A. bocasensis has a blunt, paucispiral protoconch and weaker subsutural cord, and its 

ribs have a narrower base.

Occurrence: Lower, middle, and upper parts of 

Gatun formation, eastern area (middle Miocene). Lower part, localities 138, 138c, 138d, 1381 Middle part, 

localities 139c, 155b, 155c. Upper part, eastern area, 

Reality 175.

Agladrillia (Eumetadrillia) acidna Woodring, n. sp.

Of medium size, slender, anal fascicle constricted, 

subsutural cord weak. Protoconch acute, 3-whorled. 

Subsutural cord appearing on fourth post-protoconch 

whorl, weak and slightly undulated on all whorls, not 

obscuring anal fascicle. On intermediate and late 

whorls axial ribs moderately wide, swollen, narrowcrested, separated by U-shaped interspaces; 10 on 

penult whorl. Ribs continued as narrow, low wrinkles 

on lower part of anal fascicle, greatly attenuated or 

absent on subsutural cord. Bibs of uniform spacing on 

body whorl up to hump, absent between hump and 

outer lip, replaced by a wrinkle near lip. Somewhat 

exaggerated growth threads crosscutting ribs. Faint

microscopic spiral lineation discernible. A hump on 

body whorl about 90° from outer lip. Anal sinus wide 

and deep. Stromboid notch moderately deep and siphonal notch deep, as shown by growth lines. Siphonal 

fascicle not inflated, marked by growth threads, limited 

by a low, narrow spiral thread.

Height 18 mm, diameter 5.9 mm (type).

Type: USNM 645806.

Type locality: 172 (USGS 6035, Mindi Hill cut, near 

bottom of canal, Canal Zone), upper part of Gatun 

formation.

Though the only specimen of this species, collected 

at a locality in the upper part of the Gatun formation, 

has a damaged outer lip and siphonal canal and is disfigured by two bore holes, it shows distinctive features. 

It is a little more slender than Agladrillia isthmica, 

and has a constricted anal fascicle and weaker subsutural cord; its axial ribs are narrower and have a 

narrower crest and base. The acute protoconch indicates that it is not closely allied to A. bocasensis. In 

addition, the subsutural cord of A. acidna is even weaker than that of A. bocasensis, and the rib crests are 

narrower. The crosscutting growth threads of A. acidna 

are not as strong as those of the other two species mentioned. The faint microscopic spiral lineation is discernible only because the glistening surface of the 

shell is wholly intact.

Occurrence: Upper part of Gatun formation (middle Miocene), eastern area, locality 172.

Genus Syntomodrillia Woodring

Woodring, Carnegie Inst. Washington Pub. 385, p. 160, 1928. 

Type (orthotype): Drillla lissotropis (Dall) [Pleurotoma (Mangilia) lissotropis Dall], liA^ing, Gulf of Mexico and Florida

to West Indies.

Though the minute genus /Syntomodrillia has not 

been found in the faunas under consideration, a misrepresentation concerning it should be rectified.

Bartsch (1934, p. 25) gratuitously confused the designation of the type species. He claimed that Pleurotoma (Mangilia] lissotropis Dall (1881, p. 58) is not 

the same species as Drillia lissot'/'opis (Dall) (1889, 

p. 91, pi. 11, figs. 3, 4). (As a matter of fact, when the 

name Syntomodrillia was proposed, I neglected to cite 

Dall's original combination.) He further remarked 

that "Woodring, in selecting a specimen for his genotype" chose a specimen that has microscopic spiral 

threads between the ribs of late whorls. The type of a 

genus is, of course, a species, not a specimen. Bartsch's 

statement was based on the description of the genus, 

which was not a description of the type species, much 

less a description of a specimen of the type species. It
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 389

would have been better, however, to include in the 

description of the genus a statement that faint microscopic spiral sculpture may be present or absent on 

late whorls.

The immature specimen (height 4.5 mm; Dall, 1889, 

pi. 11, fig. 3) that is the type of Pleurotoma (Mangilia) 

lissotropis has not been found in the collections of the 

TT.S. National Museum or of the Museum of Comparative Zoology. In his 1881 description Dall mentioned 

110 spiral sculpture above the pillar, but, according to 

his 1889 description "the spaces between [the ribs] are 

indifferently perfectly smooth, or finely spirally striate, 

especially toward the anterior end of the shell"; that 

is, on late whorls. That statement was found to be 

appropriate when Syntomodrillia was named, and still 

is appropriate. Bartsch (1934, p. 27, pi. 8, figs. 5, 7, 9) 

selected from a lot of seven, identified by Dall as Drillia 

lissotropis (USNM 87477, off Barbados, 103 fathoms), 

a specimen that has faint microscopic spiral threads 

on late whorls, too strongly retouched on his illustrations, as the type of his new species /Syntomodrillia 

woodringi, and cited that species as the type of Syntomodrillia. (The height is 8.6 mm, not 18.2 mm). S. 

looodringi is a synonym of S. lissotropis.

Powell, as summarized in his latest publication (1966, 

p. 84), assigned to Syntomodrillia species in the Tertiary deposits of New Zealand and Australia and 

treated it as a subgenus of Splendrillia. His size range 

for Syntomodrillia (height 4.5 to 18 mm) was based 

on the erroneous dimension of S. woodringi. No American species has a height greater than 10 mm. The 

earliest American species is of early Miocene age (Thomonde formation of Haiti).

Genus Microdrillia Casey

Casey, Acad. Nat. Sci. Phila. Proc., v. 55, p. 276, 1903.

Type (logotype, Cossmann, Essais de paleoconchologie comparee, pt. 7, p. 223, 1906): Pleurotoma cossmanni Meyer,

1887, Eocene, Miss.

Meyer's name, of course, is not a primary homonym 

of Purpura cossmanni de Raincourt (1884, p. 345, pi. 

12, fig. 11) ; nor is it a secondary homonym. Cossmann 

(1889, p. 252; p. 256 of separate publication), thinking 

that de Raincourt's and Meyer's species are congeneric, 

proposed the substitute name Oligotoma meyeri for 

Meyer's species. That name, in the combination P. 

[leurotoma] meyeri, used by Cossmann in 1906, is generally cited for the type of Microdrillia: Purpura cossmanni., however, is not a Microdrillia.

This ancient, widespread genus widespread in Tertiary seas and in modern warm seas occurs in the 

Gatuii formation.

Microdrillia trina Mansfield 

Plate 59, figures 7-9; plate 64, figure 5

Microdrillia trina Mansfield, U.S. Natl. Mus. Proc., v. 66, art. 

22, p. 28, pi. 4, fig. 5, 1925 (Miocene, Trinidad).

Moderately small, whorls generally carinate at periphery, exceptionally flat-sided. Protoconch acute, 4- 

to 514-whorled (generally 5), last to last 214 whorls 

(generally 11/4 to li/£) bearing arcuate axial riblets, 

ending just below middle of whorl, or extending practically to base of whorl. A gradually strengthened 

sutural spiral thread on sculptured part of protoconch. 

First post-protoconch whorl sculptured with a sutural 

spiral thread (a continuation of protoconch thread), a 

peripheral spiral thread, and generally a lower spiral 

thread. On some shells appearance of lower thread 

delayed until second whorl. Remaining spire whorls 

generally bearing three spiral threads, exceptionally 

four. Position and relative strength of threads variable. 

Peripheral or lower thread, or both, generally stronger 

than sutural thread, modifying whorl profile, except 

on flat-sided shells, which have subequal threads. Regularly spaced submicroscopic axial threads on postprotocoiich whorls, arcuate on anal fasciole, protractive 

below fasciole. Anal sinus between sutural and next 

spiral thread, apex semicircular. Interior of outer lip 

bearing 4 widely spaced lirse. Two elongate denticles 

present or absent on columellar lip. Siphonal notch 

shallow. Siphonal fasciole slightly inflated, limited by 

a narrow spiral thread, sculptured with a few coarser 

threads.

Height 6.7 mm, diameter 2 mm (largest specimen, 

flat-sided). Height 5.6 mm, diameter 2 mm (larger 

figured carinate specimen).

Type: USNM 352624.

Type locality: USGS 9212, about one mile south of 

Brasso, Trinidad, flood wash from stream, Brasso formation.

Microdrillia trina was found in the three parts of 

the Gatun formation in the eastern area. It is rare in 

the lower and upper parts, and locally abundant in the 

middle part. One hundred and sixty-five of the 225 

specimens were collected at locality 147b, where so 

many small species are abundant. Th<* outer lip of all 

is more or less defective and only one (locality 177b) 

has an intact siphonal canal.

This is a variable species variable in size and profile, 

and in relative strength of the spiral threads. The prevailing profile and sculpture is shown on plate 59, figure 9; subequal peripheral and lower spiral threads on 

plate 59, figure 8. Ten shells from the three parts of 

the Gatun are flat-sided and have subequal spiral 

threads. They reach a relatively large size: height up
390 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

to 6.7 mm. Although no complete shell of normal profile is that large, a few incomplete ones indicate a comparable size. Plate 59, figure 7, represents one of the 

flat-sided shells. One specimen (locality 147g) has two 

spiral threads below the peripheral thread on the penult and antepenult whorls, and another (locality 174) 

shows the same feature on the last half of the penult. 

This condition is determined, of course, by the low 

height of whorl overlap.

When M. trina was named it was based on a float 

specimen from the early middle Miocene part of the 

Brasso formation (upper part of Globorotalia fohsi 

zone s.l.). An outcrop collection from the Globorotalia 

fohsi barisanensis zone on Mayo River (USGS 21234, 

deposited by H. G. Kugler) contains 25 specimens of 

the Microdrillia. They show practically the same range 

of variation as the more numerous Gatun specimens. 

This species ranges upward into the San Jose calcareous silt member of the Manzanilla formation (USGS 

21233).

M, trina occurs also in the subsurface La Rosa formation of the Maracaibo basin, Venezuela. In fact, 

Hoffmeister (1938, p. 106) named a zone the Microdrillia zone. According to his faunal lists (Hoffmeister, 

1938, p. 106, 107-109), both that zone and the underlying Oadulus zone are of middle Miocene age. Fossiliferous outcrop strata at El Mene de Saladillo near Los 

Quiroz, Hodson's (in Hodson and Hodson, 1931, p. 5) 

localities 6 [and 1140], which have been assigned to 

the La Rosa formation, are late early Miocene.

M. trina is closely related to the living M. comatotropis (Dall) (1881, p. 58; 1889, p. 116, pi. 11, fig. 12), 

which ranges from Georgia to Yucatan Strait and the 

West Indies at depths of 100 to 640 fathoms. It has a 

more evenly tapering protoconch than the Miocene 

species, a weaker sutural spiral thread, and lacks lirse 

on the interior of the outer lip. The type evidently is 

lost. Dall thought that Pleurotoma (Mangelia] tiara 

Watson (1886, p. 347, pi. 21, fig. 7) is a synonym. If 

so, the geographic range is to be extended to Bermuda 

and the depth range to 1,075 fathoms.

So far Microdrillia has been found only in middle 

Miocene deposits in the Caribbean region. The youngest species in southern United States, M. liebetika 

Gardner (1926-1947, p. 356, pi. 42, figs. 15, 16, 1938), 

occurs in the late early Miocene Chipola formation of 

Florida. Aside from its smaller protoconch and unsculptured siphonal fasciole, it is similar to the middle 

Miocene Jamaican M. tersa (Woodring, 1928, p. 197, 

pi. 8, fig. 14).

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene), eastern area. 

Lower part, localities 138, 138c. Middle part, localities 139b, 139c, 146, 147b, 147f, 147g, 147h, 153a, 155, 

155c, 159d. Upper part, eastern area, localities 174, 

I77b, 177c. Early middle Miocene part, of Brasso formation, Trinidad. San Jose calcareous silt member of 

Manzanilla formation (late middle Miocene), Trinidad. Subsurface La Rosa formation (middle Miocene), 

Maracaibo basin, Venezuela.

Subfamily MANGELIINAE 

Genus Lepicythara Olsson

Olsson, Neogene mollusks from northwestern Ecuador, p. 110,

1964. 

Type (orthotype): Cythara terminula Dall, Pliocene, Fla.

A genus of relatively large, biconic endemic American niangelines, of traditional "Cythara" aspect, has 

long needed a name, which was supplied recently by 

Olsson. The species of this genus have an age range 

of early Miocene to Pliocene. The axial riblets on the 

last whorl of the protoconch of the type species are so 

faint that good preservation is needed to show them.

Lepicythara heptagona (Gabb) 

Plate 60, figure 4; plate 64, figure 11

MangcUa heptaaona Gabb, Am. Philos. Soc. Trans., n. ser., v.

15, p. 211, 1873 (Miocene, Dominican Republic). 

Cythara heptaaona (Gabb), Brown and Pilsbry, Acad. Nat.

Sci. Phila. Proc., v. 63, p. 345, 1911 (Miocene, Canal

Zone). Pilsbry, Idem, v. 73, p. 322, pi. 17, fig. 9, 1922

(Miocene, Dominican Republic). 

Cythara heptagona? (Gabb), Weisbord, Bull. Am. Paleontology,

v. 14, no. 54, p. 55, pi. 5, figs. 13, 14, 1929 (Miocene,

Colombia). 

Cythara ccrcadica Maury, Bull. Am. Paleontology, v. 5, no. 29,

p. 61, pi. 9, fig. 15, 1917 (Miocene, Dominican Republic). 

fCythara cf. cercadica Maury, Oinomikado, Geol. Soc. Japan

Jour., v. 46, p. 624, 1939 (Miocene, Colombia). 

Cythara terminula costaricensis Olsson, Bull. Am. Paleontology,

v. 9, no. 39, p. 77, pi. 5, figs. 21, 22, 1922 (Miocene, Costa

Rica). 

Cythara sp., Mansfield, U.S. Natl. Mus. Proc., v. 66, art. 22, p.

26, 1925 (Miocene, Trinidad).

Relatively large, biconic, spire moderately high. 

Protoconch a wide-based cone of 3!/£> whorls, tip acute, 

last whorl strongly bulging near base and bearing arcuate axial riblets, all except the last 4 or 5 of which are 

faint and closely spaced. End of protoconch marked 

by appearance of heavier subcarinate axial ribs. First 

post-protoconch whorl progressively more distinctly 

subcarinate near base, sculptured with 6 to 8 axial ribs. 

Later whorls sculptured with widely spaced, narrowcrested axial ribs, alined from whorl to whorl, or not 

alined; 7 or 8 011 penult whorl, 7 to 9 on body whorl. 

Spiral sculpture limited to submicroscopic, irregularly 

spaced, shallow grooves and microscopic lineation. 

Faint microscopic axial threads visible in widest
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 391

grooves. Rib adjoining outer lip varicose. Anal sinus 

adjoining suture, shallow. Aperture long, narrow. 

Siphonal notch shallow. Siphonal fascicle barely inflated.

Height 11.5 mm, diameter 5.7 mm (figured specimen). Height 12.7 mm, diameter (incomplete) 5.2 mm 

(largest specimen).

Type: Acad. Nat. Sci. Phila. 2915.

Type locality: Dominican Republic, Miocene.

This relatively large mangeline is rare, although it 

was collected in the three parts of the Gatun formation. Only three of the six specimens are complete, 

mature shells and one of those is worn. The protoconch 

is preserved on three. As a matter of fact, though 

Lepicythara heptagona is widely distributed, it has not 

yet been found to be abundant anywhere.

Weisbord's Colombian specimen is poorly preserved, 

but two well-preserved specimens are in a north Colombian collection (USGS 7873, about half a kilometer 

east of Usiacuri). Though Mansfield's middle Miocene 

Trinidad shell is imperfect and immature, it shows 

the protoconch and basic post-protoconch features. 

Oinomikado's record of a comparable form in southwestern Colombia is undocumented, but the species 

itself occurs in Darien (USGS, 8477).

Details of the pattern of the finely engraved spiral 

grooves (or striae) differ from place to place. The pattern is essentially the same in the Canal Zone, Colombia, and Trinidad. The grooves of shells from both 

the Cercado and Gurabo formations of the Dominican 

Republic and from Costa Rica are in general more 

uniformly spaced than at other localities.

The Lepicythara in the late Miocene Savaneta glauconite sandstone member of the Springvale formation 

of Trinidad Rutsch's "Oythara" (Brachycytharaf] 

cf. termimda Dall (Rutsch, 1942, p. 169, pi. 3, figs. 10, 

11) has a lower spire and heavier axial ribs than L. 

heptagona. On the contrary, the ribs of L. camaronensis (Olsson, 1964, p. 110, pi. 20, fig. 3; late Miocene or 

early Pliocene, Ecuador), are weak.

L. terminula (Dall) (1890-1903, p. 38, pi. 2, fig. 5, 

1890), from the Pliocene Caloosahatchie formation of 

Florida, is the type species and the youngest Lepicythara. It occurs also in deposits of late Miocene age in 

peninsular Florida, where it is represented in Druid 

Wilson's collection from Acline. Pilsbry thought that 

this species is the same as L. heptagona. Though a close 

relationship is unmistakable, the protoconch stage of 

four or five widely spaced axial riblets is skipped on 

the Florida species; the subcarinate outline is continued on the second post-protoconch whorl; and the 

axial ribs of spire whorls are attenuated near the 

suture. The Florida species also reaches a slightly

larger size: 17 mm, as compared with a maximum of 

15 mm for L. heptagona.

A distinctive species is found in late Miocene deposits in western Florida: Brachycythara turrita 

Mansfield (1930, p. 43, pi. 3, fig. 8). The last % protoconch whorl is subcarinate and bears fairly strong, 

closely spaced axial riblets; the lower part of the first 

three post-protoconch whorls also are subcarinate and 

swollen; and the axial ribs on those and later whorls 

are attenuated near the suture.

The earliest species showing alliance to L. heptagona are of early Miocene age: "Cythara1 '1 basilissa 

Gardner (1926-47, p. 344, pi. 42, figs. 23, 24, 1938; 

Chipola formation, Fla.) and an undescribed species 

from the Thomonde formation of Haiti (USGS 9945, 

9946).

Occurrence: Lower, middle and upper parts of 

Gatun formation (middle Miocene). Lower part, locality 138a. Middle part, eastern area, localities 155, 157; 

western area, locality 161c. Upper part, western area, 

locality 185. Middle Miocene deposits, Costa Rica, 

Colombia, Darien, Panama. Early middle Miocene 

part of Brasso formation, Trinidad. Cercado and Gurabo formations (middle Miocene), Dominican Republic.

Genus Ithycythara Woodring

Woodring, Carnegie Inst. Washington Pub. 385, p. 168, 1928. 

Type (orthotype): Mangilia [Mangelia] psila Bush, living,

North Carolina to Florida, eastern Gulf of Mexico, and

West Indies.

The earliest species of Ithycythara are found in deposits of early Miocene age in Puerto Rico, Haiti, and 

Florida; in Haiti and Florida in association with the 

earliest species of Lepicythara. The genus survives in 

western Atlantic and eastern Pacific waters.

Ithycythara defuniak Gardner 

Plate 60, figure 13; plate 64, figure 12

Ithycythara defuniak Gardner, U.S. Geol. Survey Prof. Paper 

142, p. 333, pi. 41, figs. 31, 32, 1937 (1938) (Miocene, 

Florida).

Very small, slender, biconic. Protoconch a widebased cone of 3 to 3~y2 whorls, tip acute, last half whorl 

bulging near base, bearing weak to moderately strong 

arcuate axial riblets; 1 or 2 faint spiral threads present 

or absent at base of whorl near end of protoconch. End 

marked by appearance of heavier axial ribs. Post-protoconch whorls sculptured with 6 narrow axial ribs, 

somewhat swollen near base on spire whorls, alined 

from whorl to whorl, except on first. A faint suggestion of microscopic, spiral lineation on body whorl. 

Rib adjoining outer lip varicose. Anal sinus shallow. 

A small denticle on interior of outer lip at lower end

368-278 O - 70 - 7
392 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

of sinus. Other weaker denticles present or absent. 

Siphonal canal not notched, siphonal fasciole barely 

inflated.

Height 4 mm, diameter 1.5 mm (figured specimen).

Type: USNM 352137.

Type locality: USGS 7264, Alaqua Creek, Walton 

County, Florida, Shoal Kiver formation.

Four small shells from widely separated localities in 

the Gatim formation are identified as Ithycythara defuniak. One consists of the protoconch and three postprotoconch whorls, but the others have an additional 

whorl and their varicose rib adjoining the outer lip 

indicates they are mature. If so, they represent a small 

race of the Florida form, which has five whorls, making a difference of two millimeters in the height. The 

type is the only Florida specimen now available. /. defunidk may be a synonym of Mangilia tarn, Maury 

(1910, p. 13, pi. 3, fig. 7), from the Oak Grove sand 

member of the Shoal Kiver formation, but the type 

and other specimens handled by Maury the only 

available material is poorly preserved or immature.

Unlike most species of Ithycythara, including the 

type species, the whorls of /. defuniak are not even 

slightly angulated.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, locality 138c. 

Middle part, eastern area, locality 147b; western area, 

locality 161c. Shoal Kiver formation (middle Miocene), Florida.

Ithycythara cf. I. elongata (Gabb) 

Plate 60, figure 14

Moderately small, slender, biconic. Last half protoconch whorl preserved; strongly bulging near base 

and bearing closely spaced, narrow, arcuate axial riblets. First two post-protoconch whorls somewhat angulated. Post-protoconch whorls sculptured with 6 narrow axial ribs, alined from whorl to whorl. Spiral 

sculpture consisting of faint, microscopic, closely 

spaced threads of varying width. Rib adjoining outer 

lip varicose. Anal sinus shallow. A denticle on interior of outer lip at base of sinus. Siphonal canal not 

notched, siphonal fasciole barely inflated.

Height (not quite complete) 6.2 mm, diameter 2.2 

mm (figured specimen).

The description is based on a specimen from the 

lower part of the Gatun formation at locality 138c, 

where it is associated with Ithycythara defuniak,. It is 

larger and more slender than that species, and has 

distinct spiral sculpture. It is smaller and more slender 

than /. elongata (Gabb) (Pilsbry, 1922, p. 323, pi. 18, 

fig. 6), which occurs in the Cercado formation of the

Dominican Republic and also in the late Pliocene Moin 

formation of Costa Rica. Moreover, the preserved part 

of the protoconch of the Gatun species bulges more 

strongly, and its axial riblets are narrower and more 

closely spaced. None of the few specimens of /. elongata in the USNM collections shows spiral sculpture, 

but that may be a matter of preservation. Spire whorls 

and the upper part of the body whorl of the type is 

faintly sculptured.

A species comparable in sculpture to the Gatun species, but smaller and more slender, is found in middle 

Miocene deposits in Costa Rica (USGS 5288g).

Occurrence: Lower part of Gatun formation (middle Miocene), locality 138c.

Genus Cytharella Monterosato?

Monterosato, Soc. Malac. Italiana Bull., v. 1, p. 73, 1875 (Cyliarella by error).

Type (logotype, Woodring, Carnegie Inst. Washington Pub. 385, 

p. 168, 1925): Murex costatus Donovan, living, northeastern Atlantic Ocean and Mediterranean Sea.

A few years after the name was proposed Monterosato (1884, p. 128) corrected the spelling, when he 

cited his genus as a synonym of Mangelia.

Cytharella? cf. C. compsacosta (Gardner) 

Plate 60, figure 12

Very small, slender, periphery of whorls rounded. 

Protoconch missing, except about last half whorl, corroded. First post-protoconch whorl sculptured with 

about 10 axial ribs, swollen on lower part of whorl, 

attenuated near suture; spiral sculpture obscure. Remaining whorls sculptured with widely spaced, swollen axial ribs, somewhat attenuated near suture; 6 on 

penult and body whorls. Spiral sculpture consisting 

of faint, closely spaced, microscopically roughened 

threads. Outer lip broken back. Anal sinus shallow, 

according to growth lines. Aperture moderately long, 

narrow. Siphonal canal not notched, siphonal fasciole 

barely inflated, its spiral sculpture weaker than that 

elsewhere on body whorl.

Height (not quite complete) 4.2 mm, diameter 1.7 

mm (figured specimen).

Cytharella? cf. C. compsacosta, like Ithycythara cf. 

/. elongata, is represented by an almost complete shell 

collected at locality 138c. Whether it is mature is indeterminable. In general features it resembles Ithycythara compsacosta (Gardner, 1926-47, p. 334, pi. 41, 

figs. 33, 34, 1938; Chipola formation, Florida), but has 

more swollen axial ribs and distinct spiral sculpture.

Cytharella seems to be an appropriate name for 

small mangelines of traditional "Oythara" aspect that
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 393

have a narrow, long, or moderately long, aperture; 

strong, nonalined axial ribs, generally widely spaced, 

except on the earliest post-protoconch whorls; and 

faint spiral sculpture. These species include Ithycythara compsacosta, as well as four other Chipola species described in the same publication ("Mangelia" 

cryptopleura Gardner, "Cythara" chariessa Gardner, 

"6X ." barbadoides Gardner, and "6V isabellce Maury), 

Mangelia elevata Gabb (Pilsbry, 1922, p. 323, pi. 18, 

fig. 3; Miocene, Dominican Kepublic), Cytharella 

limata Olsson, 1922, p. 77, pi. 5, fig. 20; Miocene, Costa 

Rica), Brachycythara galce Fargo (in Olsson and Harbison, 1953, p. 388, pi. 20, figs. 5, 5a; Pliocene, 

Florida), and Cytliara cymella Dall (1889, p. 101, pi. 

12, fig. 4; living, Florida, Gulf of Mexico, eastern 

Caribbean Sea). The type species of Cytharella has 

spiral threads between the arcuate axial riblets on the 

last quarter whorl of the protocoiich. The Pliocene 

Florida species is the only one of the American species 

mentioned that shows this feature. The others have a 

protocoiich like that of Ithycythara; nevertheless the 

basic protocoiich pattern is similar in all. Some specimens of Ithycythara, as noted in the description of 

/. defwniak) have faint protocoiich spiral threads. 

The protocoiich of the Gatun species doubtfully assigned to Cytharella is unknown.

Occurrence: Lower part of Gatun formation (middle Miocene), locality 138c.

Genus Kurtziella Dall

Dall, Biol. Soc. Washington Proc., v. 31, p. 137, 1918. 

Type (orthotype): Plcnrotoma ccrinum Kurtz and Stimpson, 

living, Massachusetts to Florida.

A considerable number of generic names have been 

proposed for mangelines characterized by delicately 

frosted spiral sculpture. Kurtzia Bartsch (1944, p. 63; 

type (orthotype) : Mangilia arteaga Dall and Bartsch, 

living, Vancouver Island) and Granotums Fargo (in 

Olsson and Harbison, 1953, p. 394; type (orthotype) : 

Granoturris padolina Fargo, Pliocene, Florida) may 

be treated as synonyms of Kurtziella. The siphonal 

canal of all the species is not notched and the siphonal 

fascicle is not inflated or barely inflated.

Submenus Kurtziella s.s.

The reticulately sculptured, minutely noded protocoiich is a readily recognized feature of Kurtziella s.s. 

Though the subgenus has been found in the Miocene 

of the Dominican Republic, Costa Rica, and Panama, 

and in the Pliocene of Costa Rica and Venezuela, the 

Venezuelan occurrence is the only one heretofore recorded.

Kurtziella (Kurtziella) pagella Woodring, n. sp. 

Plate 60, figure 5; plate 64, figure 6

Very small, slender, spire whorls subangulated. Protocoiich a wide-based cone of 3 to 3^ whorls, tip acute 

(generally broken), last whorl (or little more or less) 

generally subcarinate near base, bearing gradually 

strengthened reticulate, minutely noded sculpture. End 

marked by appearance of heavier axial ribs. Post-protoconch whorls sculptured with widely spaced, narrow 

axial ribs (10 or 11 on first whorl, 8 on body whorl), 

and primary and secondary minutely frosted spiral 

threads. Three or four primary threads on and below 

periphery of spire whorls, peripheral thread slightly 

stronger than others. Closely spaced secondary threads 

between suture and periphery and between primary 

threads. Anal sinus wide and moderately deep, as 

shown by growth lines.

Height 4 nun, diameter 1.8 mm (type). Height 4.9 

mm, diameter 1.7 mm (exceptionally large specimen).

Type: USNM 645816.

Type locality: 147b (USGS 6033c, Panama Railroad, about 3,500 feet (1,065 meters) southeast of 

Gatun railroad station, Canal Zone), middle part of 

Gatun formation.

This minute Kurtziella was collected at 12 localities 

in the lower and middle parts of the Gatun formation, 

but, like many other small species, is abundant only 

at the type locality, which yielded 90 specimens. It occurs also in middle Miocene deposits in Costa Rica 

(USGS 5882h). The single Costa Rican shell has an 

intact outer lip, whereas the lip is broken on all of the 

some 130 Panama specimens.

The combination of small, slender outline and widely 

spaced axial ribs distinguish K. pagella from other 

species. A similar, but more inflated, species is found 

in the Cercado formation of the Dominican Republic 

(USGS 8525).

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 137, 

138a, 138c, 138d, 138e. Middle part, eastern area, localities 139b, 139c, 146, 147b, 147g, 147h, 153a.

Kurtziella (Kurtziella) stenotella Woodring, n. sp. 

Plate 60, figure 6; plate 64, figure 9

Very small, moderately inflated, spire whorls subangulated. Protocoiich like that of Kurtziella pagella^ 

last to last iy2 whorls sculptured. Axial ribs narrow 

on first post-protoconch whorl, moderately wide on 

succeeding whorls, widely spaced; 11 or 12 (generally 

12) on first whorl, 7 to 9 (generally 8) on body whorl. 

Spiral sculpture on and below periphery and above 

pillar consisting of closely spaced flat bands; 4 to 6
394 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

on penult whorl; a fine thread between some bands. 

Spiral threads between periphery and suture and a 

wider thread adjoining suture. Frosting greatly subdued. Growth lines indicating a wide, moderately deep 

anal sinus.

Height 4.5 mm, diameter 1.9 mm (type).

Type: USNM 645818.

Type locality: 147b (USGS 6033c, Panama Railroad, about 3,500 feet (1,065 meters) southeast of Gatun 

railroad station, Canal Zone), middle part of Gatun 

formation.

Kurtziella stenotella is of about the same size as K. 

pagella and has the same kind of protoconch, but its 

post-protoconch sculpture is notably different. It is associated with K. pagella at the type locality and at 

three other localities in the middle part of the Gatun 

formation. Like K. pagella, it is abundant only at the 

type locality, where 30 of the 45 specimens were collected. One specimen is in an upper Gatun collection.

In general sculptural plan K. stenotella is similar 

to the living Panamic species "Mangelia" cymatias 

Pilsbry and Lowe (1932, p. 56, pi. 3, fig. 10). The axial 

ribs of that species, however, are narrower and are 

angulated at the periphery.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene), eastern area. Middle part, 

localities 146, 147b, 147f, 147g, 147h, 155. Upper part, 

eastern area, locality I77b.

Subgenus Cryoturris Woodring

Woodring, Carnegie Inst. Washington Pub. 385, p. 178, 1928. 

Type (orthotype): Cryoturris cngonia Woodring, Miocene, Jamaica.

Cryoturris, which was proposed at the generic level, 

is similar to Kurtziella s.s., but lacks spiral sculpture 

on the protoconch. Saccharoturris (Woodring, 1928, p. 

182; type (orthotype) : Mangilia consentanea Guppy, 

Miocene, Jamaica), characterized by a noded carina 

on the last two protoconch whorls and by strongly 

shouldered post-protoconch whorls, also may be treated 

as a subgenus of Kurtziella.

The earliest species of Cryoturris now known occur 

in the Chipola formation of Florida and the Thomonde 

formation of Haiti, both of late early Miocene age. 

The Oligocene species described as Cryoturris? hittsJ)orougJiensis (Mansfield, 1937, p. 92, pi. 2, figs. 7, 10) 

evidently is a turriculine turrid.

Kurtziella (Cryoturris) habra Woodring, n. sp. 

Plate 60, figure 15

Small, slender, early whorls angulated, later whorls 

bulging, anal fascicle constricted. Protoconch acute, 

31/^-whorled, last half whorl bulging and bearing

slightly arcuate riblets, at first closely spaced, later 

more widely spaced. End not sharply defined, marked 

by strengthening of axial ribs and appearance of faint 

spiral threads below periphery. First post-protoconch 

whorl sculptured with about 15 narrow, angulated 

axial ribs and faint spiral threads. At later stages axial 

ribs progressively less angulated, extending onto lower 

part of anal fascicle in attenuated form; 11 on body 

whorl. Spiral sculpture consisting of not well differentiated, minutely frosted, primary and secondary 

threads; 5 primary threads on penult whorl. Peripheral primary thread swollen on crest of ribs, others 

somewhat swollen. A secondary thread on anal fasciole 

almost as strong as primary threads. Anal fasciole 

constricted. Anal sinus wide, of moderate depth.

Height 5.2 mm, diameter 1.9 mm (type).

Type: USNM 645820.

Type locality: 138a (Stanford University 2656, 

Transisthmian Highway, latitude 9° 21' N., plus 5,000 

feet (1,525 meters), longitude 79° 50' W., plus 1,000 

feet (300 meters), Panama; same as USGS 16909), 

lower part of Gatun formation.

Kurtziella habra is based on a specimen from the 

lower part of the Gatun formation. Its constricted anal 

fasciole sets it apart from other species of Cryoturris. 

Aside from that feature, it is similar to Turns (Bela) 

dominicensis Gabb (Pilsbry, 1922, p. 322, pi. 35, fig. 

1), although the primary spiral threads of the Gatun 

fossil are stronger and its secondary spiral threads 

are weaker. "Turns" dominicensis is found in the Cercado formation of the Dominican Republic.

Occurrence: Lower part of Gatun formation (middle 

Miocene), locality 138a.

Kurtziella (Cryoturris) species 

Plate 60, figure 8

Very small, slender, whorls angulated. Protoconch 

acute (tip broken after description was written), 3- 

whorled, last % whorl carinate near base and bearing 

slightly arcuate axial riblets extending from carina to 

suture. End marked by abrupt appearance of axial and 

spiral sculpture. Post-protoconch whorls sculptured 

with axial ribs (10 on first whorl and 9 on last complete whorl) and closely spaced, coarsely frosted spiral 

threads. Peripheral spiral thread stronger than others. 

Those between periphery and suture of essentially uniform rank, those below periphery less uniform. Outer 

lip broken back, anal sinus not outlined.

Height 2.5 mm, diameter 1.2 mm (figured specimen).

This unnamed species, like the preceding, is represented by one specimen from the lower part of the 

Gatun formation. It doubtless is immature. It is un-
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 395

like any known species of Cryoturris; in fact, the 

coarsely frosted spiral sculpture suggests species of 

Kurtziella s.s.

Occurrence: Lower part of Gatun formation (middle 

Miocene), locality 138d.

Genus Nannodiella Ball

Dall, U.S. Natl. Mus. Proc., v. 56, p. 59, 1919. 

Type (orthotype): Nannodiella nana (Dall) [Philberta (Nannodiella) nana Dall], living, Gulf of California.

The extraordinarily large anal sinus of Nannodiella 

is unique among American mangeline turrids. The 

aperture is armed with denticles of varying strength 

or unarmed. The genus has been found in the lower 

Miocene of Florida; the middle Miocene of Trinidad, 

the Dominican Republic, Jamaica, the Caribbean coastal area of Panama, and Florida; the upper Miocene 

of the Bocas del Toro area, Panama, Florida, and 

Ecuador; and the Pliocene of Ecuador and Florida; 

and is living in western Atlantic and eastern Pacific 

waters, and also, according to Powell (1966, p. 115) 

in the western Pacific Ocean. Of the three species, 

other than the type species, assigned, or doubtfully 

assigned, by Dall (1919, p. 60-61) to Nannodiella, only 

Philbertia (N annodiell'O) fratertmlis is acceptable as 

Nannodiella.

Nannodiella rintriada (Mansfield) 

Plate 60, figure 10; plate 64, figure 7

Glyphostoma amicta rintriada Mansfield, U.S. Natl. Mus. Proc., v. 

66, art. 22, p. 27, pi. 4, figs. 2, 3, 1925 (Miocene, Trinidad).

Very small, slender, anal fascicle constricted. Protoconch 4-whorled, tip acute and minute, last whorl 

to last 11/2 whorls carinate below middle of whorl. End 

marked by appearance of sculpture. First post-protoconch whorl carinate, succeeding whorls progressively 

less carinate. Post-protoconch whorls sculptured with 

10 to 13 (generally 10 to 12) narrow axial ribs, pinched 

on anal fascicle, but continuing to suture. Primary 

spiral sculpture consisting of 2 subequal threads below 

anal fascicle on spire whorls (exceptionally 3 on penult 

whorl), 3 on body whorl, overriding ribs and slightly 

swollen on their crests. Anal fascicle bearing 4 to 6 

very narrow threads; wider, more closely spaced 

threads on lower part of body whorl. Body whorl swollen adjoining outer lip. Anal sinus extraordinarily 

large for size of shell and aperture, apex and upper 

limb heavily bordered. Aperture short. Outer and 

columellar lips bearing moderately strong denticles. 

Siphonal canal very short, barely emarginate; siphonal fascicle barely inflated.

Height 4.5 mm, diameter 1.9 mm (figured specimen).

Type material: Lectotype, herewith designated, 

larger of 2 syntypes, USNM 352642.

Type locality: USGS 8302, a mile south of Brasso 

railway station, Trinidad, flood-wash from stream 

bank, Brasso formation.

The middle part of the Gatun formation yielded 32 

specimens all except seven from the rich locality 147b 

 of this small mangeline. The axial ribs on the single 

specimen from the upper part of the formation are 

more subdued than on the other shells.

Nannodiella rintriada is more slender than N. amicta 

(Guppy) (Woodring, 1928, p. 195, pi. 8, fig. 12), a 

middle Miocene Jamaican species; its primary spiral 

threads are of subequal strength; and its apertural 

denticles are stronger. The type material of N. rintriada, like that of 30 other species and subspecies 

named by Mansfield, was collected from float along 

Caparo Eiver above Brasso railway station: a convenient collecting locality, but a poor type locality. 

Eight specimens are in an outcrop collection from 

Mayo River (USGS 21234, Globorotalia fohsi robusta 

zone) and seven in an outcrop collection from Cascade River (USGS 19856, upper part of Globorotalia 

fohsi zone s.l.), both deposited by H. G. Kugler. The 

shells from locality 21234 have heavier apertural denticles than Gatun shells, whereas those from locality 

19856 have weaker denticles than those from the Gatun 

formation.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

localities 139c, 147b, 147g, 155, 155c; western area, 

locality 161. Upper part, eastern area, locality 163. 

Early middle Miocene part of Brasso formation, Trinidad.

Nannodiella cf. N. melanitica (Dall)

Two specimens found at locality 139b represent a 

second species of Nannodiella, although N. rintriada 

occurs in another collection (139c) from the same 

locality in strata near the base of the middle part of 

the Gatun formation. It is smaller (height 3.5 mm) 

and more slender than N. rintriada and its sculpture 

is more subdued. The axial ribs are narrower and 

more numerous (14 or 16 on penult whorl), and the 

primary spiral threads are narrower. Both shells are 

somewhat chalky and cannot be cleaned properly for 

photographing.

This unnamed species has stronger axial and spiral 

sculpture than Mangilia melanitica Dall (in Dall and 

Simpson, 1901, p. 390, pi. 58, fig. 10), which ranges 

from Cape Hatteras to the eastern Gulf of Mexico 

and Puerto Rico.

Occurrence: Middle part of Gatun formation (middle Miocene), eastern area, locality 139b.
396 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Genus Euclathurella Woodring 

Submenus Euclathurella s.s.

Woodring, Carnegie Inst. Washington Pub. 385, p. 187, 1928. 

Type (orthotype): Clathurella vendryesiana Ball, Miocene, Jamaica.

When Euclathurella was named and compared with 

Paraclathurella (Boettger, 1895, p. 56; type (orthotype) : Pleurotoma gracilenta Reeve, living, western 

Pacific Ocean) on the basis of Keeve's unsatisfactory 

illustration of the type species, it was not realized that 

Hedley had published a satisfactory illustration (Hedley, 1909, p. 456, pi. 44, fig. 91), recently reproduced 

by Powell (1966, p. 108, pi. 17, fig. 1). That illustration 

shows, as was suspected, that Euclathurella s.s. closely 

resembles Paraclathurella in general facies. To be sure, 

differences in post-protocoiich sculpture are apparent, 

but they are not greater than may be expected in species of the same genus. The protoconch plan, however, 

is so different that it is unlikely they are closely related. As described by Powell, the last protoconch 

whorl of Paraclathurella is sculptured with four or 

five granulose spiral threads, whereas the last half 

whorl of Euclathurella s.s. is sculptured with faint, 

closely spaced, arcuate axial riblets. The treatment of 

Kurtzietta and other genera shows that in the present 

account protoconch details are not given undue weight 

in distinguishing turrid genera. Nevertheless, protoconch plan is considered to be significant.

So far Euclathurella s.s. is represented only by two 

middle Miocene Caribbean species: an undescribed 

species from the Cercado formation of the Dominican 

Kepublic and the following species. Euclathurella f 

Uveoakensis Mansfield (1937, p. 92, pi. 2, figs. 2, 6), of 

late Oligocene age, probably is a turriculine.

Euclathurella (Euclathurella) vendryesiana (Ball) 

Plate 60, figures 2, 3

Clathurella vendryesiana Ball, in Guppy and Ball, U.S. Natl.

Mus. Proc., v. 19, p. 306, pi. 27, fig. 1, 1896 (Miocene,

Jamaica). Maury, Bull. Am. Paleontology, v. 5, no. 29,

p. 62, pi. 9, fig. 18, 1917 (Miocene, Dominican Republic). 

Euclathurella vendryesiana (Ball), Woodring, Carnegie Inst.

Washington Pub. 385, p. 189, pi. 8, fig. 1, 1928 (Miocene,

Jamaica). 

Pleurotoma (Genota) gertrudis Toula, K. k. Geol. Reichsanstalt

Jahrb., v. 58, p. 708, pi. 25, fig. 17, 1909 (Miocene, Canal

Zone). 

Pleurotoma gertrudis Toula, Brown and Pilsbry, Acad. Nat. Sci.

Phila. Proc., v. 63, p. 345, 1911 (Miocene, Canal Zone;

Toula's record).

No specimens of this species are in the Gatun collections now being studied. The type of Pleurotoma 

(Genota}gertrudis (height 12 mm, incomplete diameter

3.8 mm), now at the Technische Hochschule in Vienna, 

is illustrated. The worn protoconch is 214-whorled. 

The early post-protoconch whorls are sculptured with 

fewer axial ribs than those of specimens from Bowden, Jamaica: 10 on the antepenult whorl (the last 

whorl to bear wide axial ribs), as compared with 14 

to 16. The outer lip is broken far back.

A shell from the Cercado formation has nine antepenult axial ribs and the ribs continue in subdued 

form almost to the end of the penult whorl. The anal 

fascicle of that specimen, and also of Maury's from 

the Gurabo formation, is slightly less constricted than 

011 Bowden shells.

Type: USNM 107086.

Type locality: Bowden, Jamaica, Bowden formation.

Occurrence: Middle part of Gatun formation (middle Miocene), eastern area, Gatun Locks excavation. 

Bowden formation (middle Miocene), Jamaica. Cercado and Gurabo formations (middle Miocene), 

Dominican Kepublic.

Submenus Miraclathurella Woodring

Woodring, Carnegie Inst. Washington Pub. 385, p. 189, 1928. 

Type (orthotype): Miraclathurella vittata Woodring, Miocene, 

Jamaica.

I agree with Powell (1966, p. 109) that it is better 

to assign Miraclathuretta to subgeneric rank under 

Euclathurella than to retain the generic rank originally proposed.

The sculptured part of the protoconch of Miraclathuretta consists of less than a quarter whorl to an entire whorl. It is bulging to distinctly carinate below 

the middle of the whorl and bears a few strong, widely 

spaced, strongly arcuate axial riblets. In post-protoconch axial sculpture Miraclathurella is more similar 

to Paraclathurella than to Euclathurella s.s. The 

sutural cord of Miraclathurella, which recalls the cord 

of many clavine genera, is a readily recognized feature.

The stromboid notch, mentioned in the original description of Euclathurella and Miraclathurella, is not 

as conspicuous as the notch of Clathrodrillia and other 

clavine genera. It is an indistinct to distinct, narrow 

to wide indentation, or is absent.

Unlike Euclathurella s.s., Miraclathurella is widely 

distributed in the middle Miocene of Trinidad, the 

Dominican Kepublic, Jamaica, the Caribbean coastal 

area of Panama, southeastern Costa Rica, and Darien; 

and the upper Miocene of Trinidad, the Bocas del Toro 

area of Panama, the Limon Peninsula of Costa Rica, 

and probably the Tehuantepec area of Mexico. It is 

still living in the western Atlantic and eastern Pacific
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 397

Oceans. It is represented in the western Atlantic by 

Mangilia aguadillana Dall and Simpson (1901, p. 389, 

pi. 57, fig. 22) and an undescribed species dredged off 

southern Florida (USNM 412149), and in the eastern 

Pacific, by a species from the Gulf of California described in manuscript by Bartsch (USNM 212366, 

348242).

Euclathurella (Miraclathurella) eucharis Woodring1, n. sp. 

Plate 60, figure 21; plate 66, figure 8

Relatively large, slender, anal fascicle constricted, 

bordered by sutural cord. Protoconch acute (tip generally broken), 3- to 4-whorled. About last i/2 whorl 

bulging below middle, last 14, or less, sculptured with a 

few (as few as 2) strong, widely spaced, strongly 

arcuate axial riblets. End marked by abrupt appearance of axial ribs, a sutural spiral thread, and 2 other 

spiral threads overriding ribs; 7 to 10 ribs (generally 

8 or 9) on first post-protoconch whorl. Axial ribs on 

post-protoconch whorls wide, suppressed on lower part 

of anal fascicle; 9 or 10 (generally 10) on antepenult 

and penult whorls; reduced, narrower, and more 

numerous on body whorl. Sutural spiral cord strong. 

Spiral threads below anal fascicle narrow, overriding 

axial ribs, slightly swollen on crest of ribs on body 

and penult whorls; 4 to 6 (generally 4) on penult 

whorl; an additional weaker thread on lower part of 

anal fascicle of some specimens. Microscopic, faintly 

crimped spiral threads on anal fascicle and on upper 

part of body whorl below anal fascicle. Aperture long, 

narrow. Outer lip varicose; anal notch wide, deep, 

apex semicircular, upper limb heavily bordered; 

stromboid notch distinct, narrow.

Height 9 mm, diameter 3.6 mm (type). Height 11.2 

mm, diameter 4 mm (largest specimen).

Type: USNM 645825.

Type locality 138c (USGS 21956, about 100 meters 

north of Transisthmian Highway and about 75 meters 

west of road to refinery site on Payardi Island, 

Panama; immediately east of Cativa and 100 meters 

north of locality 138), lower part of Gatun formation.

Euclathurella eucharis occurs in the lower and middle parts of the Gatun formation and in the upper 

part in the western area. Eleven of the 55 specimens 

were found at the type locality and 12 at locality 147b. 

The single incomplete specimen from the upper part 

is more inflated than the others. Four specimens are 

in a Darien collection (USGS 8477).

This species is closely related to E. entemna (Woodring, 1928, p. 190, pi. 8, figs. 5, 6), from the Bowden 

formation of Jamaica. It is slightly smaller than the 

Jamaican species and has narrower spiral threads,

which are not as strongly swollen on the crests of the 

axial ribs, and less distinct microscopic spiral threads. 

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 137a, 138, 138a, 138c, 138d. Middle part, eastern 

area, localities 139b, 139c, 146, 147b, 147f, 147g, 147h, 

159d. Upper part, western area, locality 183. Middle 

Miocene deposits, Darien, Panama.

Genus Dolostoma Woodring1, n. gen.

Type: Dolostoma anorhepes Woodring, n. sp., Gatun formation, 

Canal Zone and Panama, Miocene.

Relatively large (height 13 to 15 mm), moderately 

inflated, anal fascicle wide or moderately wide, slightly 

concave. Protoconch a wide-based cone, tip acute, 3- 

to 3^-whorled, last whorl, or last % whorl, bulging 

near base and bearing protractive, arcuate axial riblets. At first riblets closely spaced, at later stage more 

widely spaced and spiral threads appearing between 

them. End marked by a strong axial rib. Post-protoconch spire whorls sculptured with axial ribs, overridden below anal fascicle by narrow, closely spaced 

spiral threads. Ribs pinched on anal fascicle, but extending to suture, arcuate on fascicle, except on first 

few whorls. Ribs of body whorl like those of spire 

whorls, or subdued, more numerous, and closely spaced. 

Spiral sculpture on anal fascicle limited to closely 

spaced microscopic threads, faintly crimped under high 

magnification, or consisting of threads almost as wide 

as those below fascicle. Microscopic threads present or 

absent between those of primary rank on body whorl 

between anal fascicle and pillar, and on corresponding 

part of preceding two whorls. Outer lip strongly varicose, anal sinus wide and deep, apex semicircular. Aperture long, narrow. Siphonal canal slightly notched, 

siphonal fascicle slightly inflated.

The protoconch suggests that of Kurtziella s.s., but 

the sculptured part is not subcarinate, and the weaker 

spiral threads do not override the axial riblets. The 

mature facies is entirely different. Though Dolostoma 

resembles Lioglyphostoma (p. 399) in mature facies, 

the protoconch and sculptured plan are not similar. So 

far Dolostoma has been recognized in the Gatun formation, in strata of middle Miocene age in Darien 

(USGS 8477), and in the San Jose calcareous silt member of the Manzanilla formation of Trinidad, also of 

middle Miocene age.

In preliminary sorting immature specimens of Dolostoma anorhepes, like that shown on pi. 60, fig. 9, and 

mature shells of Kurtiziella stenotella may be confused. 

They occur together in the middle part of the Gatun 

formation. On closer examination, however, they are 

seen to differ in every diagnostic feature.
398 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Dolostoma anorhepes Woodring, n. sp. 

Plate 60, figures 9, 20; plate 66, figure 2

Outline, protoconch, sculptured plan, outer lip, and 

aperture as described under the genus. Spire whorls 

moderately bulging, anal fasciole wide. Axial ribs on 

body whorl generally of about same width and spacing 

as on spire whorls, exceptionally, as on type, narrower 

and secondary ribs intercalated between them on lower 

part of whorl; 9 or 10 ribs on first post-protoconch 

whorl, 9 to 11 on penult whorl. Penult whorl bearing 

4 to 6 primary spiral threads. On body whorl spiral 

threads on pillar more closely spaced and coarser than 

elsewhere. Spiral sculpture on anal fasciole limited to 

closely spaced faint microscopic threads. Similar 

threads between primary threads visible on last three 

whorls of well-preserved mature shells. Anal fasciole 

appearing at beginning of first post-protoconch whorl, 

or on a later part of that whorl (as late as half a 

whorl), in that event preceding part rounded and 

sculptured with closely spaced spiral threads like those 

below fasciole at later stage. A faint swelling on interior of outer lip below anal sinus.

Height (almost complete) 15.5 mm, diameter 6 mm 

(type).

Type: USNM 645822.

Type locality: 138 (USGS 16909, North and south 

sides of Transisthmian Highway, 1.6 kilometers northeast of Canal Zone boundary, Panama), lower part of 

Gatun formation.

Dolostoma, anorhepes occurs in the lower and middle parts of the Gatun formation. Locality 147b, mentioned as a rich locality for many of the mangeline 

turrids, yielded 30 of the 52 specimens. Perfect preservation is needed to show the microscopic spiral sculpture. It is shown by the type, but not to good advantage. A mature specimen (locality 139c) has an intact 

siphonal canal, which is broken on the type. A varicose outer lip is preserved on two immature shells, 

even at a height of five millimeters. The shells from 

the lower part of the Gatun and from strata near the 

base of the middle part are more inflated than those 

from higher horizons.

This species is found also in the middle Miocene 

San Jose calcareous silt member of the Manzanilla 

formation of Trinidad (USGS 21745 and four other 

localities). All the specimens are slender, but not more 

slender than some from the middle part of the Gatun.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 137, 

138, 138b, 138c, 138d. Middle part, eastern area, localities 139c, 146, 147b, 147g, 147h, 153a, 159d. San Jose

calcareous silt member of Manzanilla formation (middle Miocene), Trinidad.

Dolostoma dinota Woodring1, n. sp. 

Plate 61, figure 4

Moderately large, moderately inflated, spire whorls 

strongly bulging, anal fasciole moderately wide. Anal 

fasciole not distinguishable on first three post-protoconch whorls, flat on fourth, slightly concave on fifth, 

more distinctly concave on sixth (body whorl). Tip of 

protoconch broken, remainder exfoliated. Post-protoconch spire whorls sculptured below anal fasciole with 

narrow axial ribs (12 on antepenult whorl, 16 on 

penult), overridden by flat spiral threads (5 or 6 on 

penult whorl), slightly swollen on crest of ribs. Kibs 

subdued, narrower, more numerous, and closely spaced 

on body whorl. On spire whorls ribs slightly narrower 

and slightly arcuate on anal fasciole; on body whorl 

subdued, more strongly arcuate, and interspersed with 

exaggerated growth threads. Spiral threads on anal 

fasciole a little narrower than those below fasciole; 5 

or 6 on penult whorl. On body whorl between anal 

fasciole and pillar microscopic growth threads visible 

between spiral threads. Spiral threads on pillar more 

closely spaced than those elsewhere on body whorl. A 

strong swelling on interior of outer lip below anal 

sinus.

Height (practically complete) 12.9 mm, diameter 

5.3 mm (type).

Type: USNM 645829.

Type locality: 175 (USGS 8410, cuts on north [west] 

side of French Canal [East Diversion], Mount Hope, 

Canal Zone), upper part of Gatun formation.

The type and the body whorl of a specimen of about 

the same size as the type, both found in the upper 

part of the Gatun formation near Mount Hope, are the 

only representatives of Dolostoma dinota.

The spire whorls of this species bulge more strongly 

than those of D. anorhepes] the anal fasciole appears 

at a later stage and is sculptured with spiral threads 

almost as strong as those below the fasciole; and the 

axial ribs are more numerous and, except on the 

earliest whorls, are narrower.

Occurrence: Upper part of Gatun formation (middle Miocene), eastern area, locality 175.

Genus Glyphostoma Gabb

Gabb, Acad. Nat. Sci. Philadelphia Proc., v. 24, p. 270, 1872

(1873). 

Type (monotype) : Glyphostoma dentifera Gabb [dentifcrum],

Miocene, Dominican Republic.

Some 70 late Tertiary and living species of Glypho- 

^ all American, have been described and three
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 399

more are now added. As suggested by Powell (1966, 

p. 115), non-American species are suspect. The genus 

includes the largest American mangeline turrids, reaching a maximum height of about 45 millimeters. All the 

species have one feature in common. Although the 

protoconch is blunt and paucigyrate or acute and polygyrate, the last half whorl to last whorl and a half 

(exceptionally two and a half), depending on the total 

number of whorls, are carinate. To be sure, they are 

weakly carinate or merely bulging in one subgenus: 

LioglypJiostoma. The other essential features are gradational. The apertural armature runs from strong 

through weak or faint to complete absence. The axial 

sculpture on the anal fascicle ranges from strong 

puckers through weak puckers or exaggerated growth 

threads to ordinary growth threads. Owing to gradation, the main subdivisions are regarded as subgenera. 

They may be arranged as follows:

Subgenera of Glyphostoma 

Aperture armed

Strong to weak puckers on anal fascicle. _ Glyphostoma s.s. 

Exaggerated or ordinary growth threads

on anal fascicle. 

On late whorls spiral sculpture limited

to pillar_________________________ Euglyphostoma.

On late whorls spiral sculpture not limited to pillar.____________________ Rhiglyphostoma.

Aperture unarmed

Pillar slightly or moderately constricted. Lioglyphostoma. 

Pillar strongly constricted ______________ Glyphostomops.

The earliest species occur in the Thomonde formation of Haiti (TTSGS 9907), the outcrop La Rosa 

formation of Venezuela (TJSGS 18007), and the 

Chipola formation of Florida, all of late early Miocene age. Even at that time Glyphostoma s.s., Rhiglyphostoma, and Lioglyphostoma are represented.

Glyphostoma s.s. may be divided arbitrarily into a 

group of large species (height 24 to about 45 mm) 

and a group of small species (height 9.5 to 23 mm). 

Twelve of the 15 large species are found in the middle 

and late Miocene Caribbean province, one in the early 

Miocene of Florida, one in the middle Miocene of 

Florida, and the other is living in the Caribbean 

region: G. epicasta Bartsch (1934, p. 14, pi. 4, figs. 4, 

7, 9). Ten of the large fossil species, including the 

type species and also the first to appear (G. locTdini], 

have been described recently by Olsson (1964, p. 105- 

109). The 17 small species are more evenly distributed 

in the Miocene of the Caribbean region, the Miocene 

and Pliocene of Florida, and in western Atlantic and 

eastern Pacific waters at low latitudes. One species 

occurs in the Pliocene of North Carolina the northernmost record for the subgenus.

Euglyphostoma is a minor, but morphologically distinctive, new subgenus embracing a Gatun species and 

two living eastern Pacific species.

The new subgenus Rhiglyphostoma includes a Gatun 

species, a not closely related species from the northern 

part of the late Miocene Caribbean province Pleurotoma (Glyphostoma} mexicana Toula (1911, p. 483, 

pi. 29, fig. 16) 20 overnamed Miocene and Pliocene 

species from Florida, two Miocene species from North 

Carolina, and one each from the Miocene of Maryland 

and New Jersey. Like Gymia (p. 223), Rhiglyphostoma 

reached New Jersey in middle Miocene time, but, unlike Cymia, did not bypass the Chesapeake embayment. As pointed out by Pilsbry and Harbison (1933, 

p. 113), the New Jersey record is the northernmost for 

the genus. A species, G. conradiana (Gabb), occurs in 

the Pliocene and Pleistocene of California. Living 

species are found in the western Atlantic and eastern 

Pacific Oceans, extending to higher latitudes in the 

eastern Pacific than Glyphostoma s.s.

Lioglyphostoma (Woodring, 1928, p. 193; type (orthotype) : Lioglyphostoma adematum Woodring, Miocene, Jamaica) includes five fossil species: one in the 

late early Miocene of Florida and four (two undescribed) in the middle and late Miocene of the Caribbean region. It is now represented at the south end of 

the Florida Keys and in the Caribbean Sea by a small 

species misidentified by Dall (1889, p. Ill) as Mangilia lavelleana (d'Orbigny) and in Galapagos waters 

by Glyphostoma sirena Dall (1919, p. 53, pi. 17, fig. 3). 

Other fossil and living species assigned to Lioglyphostoma are to be referred to Glyphostoma s.s. or Rhiglyphostoma, or even to other genera.

Glyphostomops (Bartsch, 1934, p. 17; type (orthotype) : Glyphostoma (Glyphostomops) hendersoni 

Bartsch, living, southern Florida) is another minor 

subgenus embracing two, possibly three, living western Atlantic species and a doubtful Pliocene species 

from Florida.

Subgenus Glyphostoma s.s.

Glyphostoma (Glyphostoma) dentiferum Gabb

Plate 60, figures 22, 25; plate 66, figure 3

Glyphostoma dentifera Gabb, Acad. Nat. Sci. Phila. Proc., v. 24, 

p. 971 [271], pi. 11, fig. 4, 1872 (1873) (Miocene, Dominican Republic). Gabb, Am. Philos. Soc. Trans., vol. 15, p. 

210, 1873 (Miocene, Dominican Republic). Maury, Bull. 

Am. Paleontology, v. 5, no. 29, p. 61, pi. 9, fig. 16, 1917 

(Miocene, Dominican Republic).

Glyphostoma, dentiferum Gabb, Pilsbry and Brown, Acad. Nat. 

Sci. Phila. Proc., v. 64, p. 501 (list), 1913 (Miocene, Canal 

Zone). Pilsbry, Idem, v. 73, p. 324, pi. 17, fig. 15, 1922 

(Miocene, Dominican Republic). Jung, Bull. Am. Paleontology, v, 49, no. 223, p. 571, pi. 77, figs. 9, 10, 1965 

(Miocene, Venezuela). Olsson, Neogene mollusks from 

northwestern Ecuador, p. 105, pi. 17, fig. 1, 1964 (Miocene, 

Dominican Republic).
400 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Clathurella (Glyphostoma) dcntifera (Hinds), Cossman, Jour. 

Conchyliologie, v. 61, p. 31, pi. 2, figs. 15-17, not figs. 18- 

20, 1913 (Miocene, Dominican Republic).

Not Glyphostoma dcntifera Gabb, Olsson, Bull. Am. Paleontology, v. 9, no. 39, p. 76, 1922 (Miocene, Costa Rica; = G. 

guppyi Woodring).

Large, moderately slender, spire not strongly turreted. Protoconch blunt, l*/^- to 2-whorled, last half 

whorl carinate. End marked by gradual appearance of 

sculpture. First post-protoconch whorl sculptured with 

7 wide axial ribs, angulated in alinement with protoconch carina, overridden by a peripheral spiral thread 

and another below periphery. Sculpture changing on 

body whorl, or on penult, or even on antepenult, depending principally on size of shell. Before changing, 

axial ribs wide and swollen, gradually disappearing 

near anal fascicle, 8 to 11 on last whorl preceding 

change. Ribs overridden by narrow spiral cords, swollen on ribs, 5 or 6 cords on penult whorl, not including 

a narrower thread adjoining anal fascicle. Change in 

sculpture due to narrowing of ribs, introduction of a 

narrow rib between them, and downward bifurcation 

of main ribs, all the ribs eventually of about same 

width, as many as 25 on mature body whorl. At same 

stage spiral cords somewhat suppressed between ribs, 

forming spirally elongate nodes on ribs. Spiral cords 

reduced in width on pillar. Anal fascicle constricted, 

bearing strong, arcuate, axial puckers and faint spiral 

threads. Puckers appearing on fourth or fifth post-protoconch whorl. Outer lip strongly varicose. Anal sinus 

narrow, deep, everted at semicircular apex. Apertural 

armature strong on both lips. Siphonal canal slightly 

notched, according to growth lines. Siphonal fascicle 

slightly inflated, sculptured with smooth or slightly 

noded spiral cords.

Height (incomplete) 38.2 mm (estimated height 43 

mm), diameter (including outer lip) 15.2 mm, (not 

including outer lip) 11.5 mm (larger figured specimen).

Type: Acad. Nat. Sci. Phila. 2910.

Type locality: Dominican Republic, presumably Cercado or Gurabo formation.

Seventeen specimens of Glyphostoma dentiferum 

were found in the lower and middle parts of the Gatun 

formation, but only one (pi. 60, fig. 22) is a mature 

shell. Much of the face of its outer lip is damaged and 

the end of the siphonal canal is missing. A more complete immature shell is shown on plate 60, figure 25. 

An incomplete, doubtfully identified specimen has a 

varicose outer lip and complete apertural armature at 

an estimated height of seven millimeters.

These Gatun fossils are consistently more slender 

than the type. As noted by Pilsbry (1922, p. 324), 

however, Gabb's nine specimens [now eight] at the

Academy of Natural Sciences of Philadelphia show 

some variation in outline. Gabb collected all the Dominican Republic shells in American museums. That 

at Cornell University, illustrated by Maury, is immature, and so is the one illustrated by Cossmann. The 

incomplete shell from the Angostura formation of 

Ecuador, illustrated by Olsson (1964, pi. 17, fig. 8) 

as Glyphostoma sp., closely resembles Gatun specimens, 

so far as it goes.

G. woodringi Olsson (1964, p. 106, pi. 18, figs. 1-lc), 

which occurs in the Cercado formation of the Dominican Republic, is more slender and more delicately 

sculptured than G. dentiferum. Lioglyphostoma woodringi Fargo (in Olsson and Harbison, 1953, p. 405, 

pi. 21, figs. 7-7b; Caloosahatchee formation, Florida) 

is a small species of Rhiglyphostoma that has weak, 

or faint, apertural armature. If the view that Rhiglyphostoma is to be assigned to subgeneric rank under 

Glyphostoma is adopted, Olsson's species needs a new 

name.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138c, 

138e (immature, identification doubtful). Middle part, 

eastern area, localities 139c, 155, 155a, 155c, 156, 157. 

Miocene, Dominican Republic. Cantaure formation 

(middle Miocene), Venezuela.

Glyphostoma (Glyphostoma) pyrgota Woodring, n. sp. 

Plate 60, figures 23, 24

Large, very slender, strongly turreted. Protoconch 

and early post-protoconch whorls missing. Sculpture 

of late whorls like that of Glyphostoma dentiferum. 

Eleven wide, swollen axial ribs on last whorl preceding 

change in sculpture. Eight similar ribs on upper part 

of early two-thirds of mature body whorl; 20 narrow 

ribs on lower part of mature body whorl. Eight spiral 

cords on penult whorl, lowermost two noded, uppermost two noded and narrower than others. Microscopic, irregularly trending, axial vermiculation visible on well-preserved whorls. Anal fascicle strongly 

constricted, sculptured with strong, arcuate, axial 

puckers. Outer lip strongly varicose, edge serrated by 

short projections opposite spaces between spiral cords. 

Anal sinus like that of G. dentiferum. End of siphonal 

canal missing.

Height (incomplete, 2 whorls) 37.3 mm (estimated 

height 45 mm), diameter (including outer lip) 15 mm, 

(not including outer lip) 7.4 mm (type).

Type: USNM 645833.

Type locality: 157 (USGS 16926, westernmost cut 

on Panama Railroad cutoff south of Fort Davis, 1.2 

miles (1.9 kilometers) northeast of Gatun railroad 

station, Canal Zone), middle part of Gatun formation.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 401

Although this handsome species is represented by 

only two incomplete specimens, it is named, as the 

slender, strongly turreted outline and microscopic, irregularly trending, axial vermiculation are unique in 

the genus. Both specimens occur in the middle part of 

the Gatun formation in association with Glyphostoma 

dentiferum. The type is the larger of the two. The 

other, which is somewhat worn and does not show the 

vermiculation, has an estimated height of 36 millimeters. The estimated height of the type indicates that 

it is the largest known Glyphostoma.

Occurrence: Middle part of Gatun formation (middle Miocene), eastern area, localities 155c, 157.

Subgenus Euglyphostoma Woodring, n. subgen.

Type: Glyphostoma partefilosa Ball, living, Gulf of California 

and west coast of Baja California.

Of medium size (height 11 to 14 mm), moderately 

slender, anal fascicle moderately to strongly constricted. Protoconch acute, 3- to 31^-whorled, last 

whorl to last 1^4 whorls carinate. Axial ribs widebased and swollen, crest continuing in subdued form 

across anal fascicle, or absent on fascicle. Spiral sculpture above pillar limited to first 2 to 4 post-protoconch 

whorls. Pillar sculptured with spiral cords, decreasing 

in width downward, those above siphonal fascicle 

slightly undulated by subdued continuation of axial 

ribs. Outer lip varicose, its edge slightly serrate. Anal 

fascicle bearing subdued, arcuate, exaggerated growth 

threads. Anal sinus wide for size of shell, deep, everted 

at semicircular apex. Apertural armature moderately 

strong. Siphonal canal moderately notched. Siphonal 

fascicle slightly inflated, sculptured with narrow spiral 

cords. Kadula consisting of a pair of long, slender, 

needlelike, nonbarbed marginals.

The absence of spiral sculpture on late whorls, except on the pillar, is the most distinctive feature of 

Euglyphostoma. The protoconch, apertural features, 

and radula are like those of Glyphostoma s.s. Fresh 

dredged shells are vitreous. The three lots of the type 

species in the USNM collections represent a depth 

range of 10 to 44 fathoms,

I am indebted to J. P. Morrison, of the U. S. National Museum, for the opportunity to examine his 

drawing of the radula of the type species. The radula 

was extracted from the animal in a worn shell dredged 

at a depth of 44 fathoms off Cedros Island, Baja California (USNM 97029).

Glyphostoma (Euglyphostoma) olssoni Woodring, n. sp. 

Plate 60, figure 7

Size, outline, sculptural plan, and apertural features 

as described under the subgenus. Anal fascicle strongly

constricted. Protoconch acute, 3-whorled, last whorl to 

last 114 whorls carinate. End marked by thin varix 

or appearance of first axial rib. First post-protoconch 

whorl sculptured with 8 or 9 axial ribs and 2 weak 

spiral cords: one at periphery and one below periphery. 

Cords suppressed between ribs, disappearing on third 

or fourth whorl. Nine or 10 ribs on penult whorl. 

Several axial puckers on anal fascicle adjoining anal 

sinus, somewhat exaggerated, arcuate growth threads 

elsewhere on fascicle.

Height 13.6 mm, diameter (including outer lip) 5.7 

mm, (not including outer lip) 4.7 mm (type).

Type: USNM 645834.

Type locality: 176 (USGS 8358, Koad bordering 

French Canal [East Diversion], near Mount Hope, 

Canal Zone), upper part of Gatun formation.

The upper part of the Gatun formation yielded three 

specimens of this species. One, an immature shell of 

4^/2 whorls (height 6 mm), shows to good advantage 

the sculpture of early whorls. Even at that stage the 

outer lip is varicose and the apertural armature is 

fully formed. The wide, swollen, smooth axial ribs of 

all except the first few whorls recall those of Eumetadrttlia (p. 387).

Glyphostoma olssoni is more inflated than >G. partefilosa Dall (1919, p. 53, pi. 17, fig. 4), the type species 

of Euglyphostoma, and has a more constricted anal 

fascicle and heavier axial ribs. In outline and constriction of anal fascicle the fossil species is more 

similar to Clavatula Candida Hinds (1844-45, p. 20, 

pi. 6, fig. 18, 1844), another living Panamic species. 

Hinds' illustration, however, shows narrower axial ribs.

This species, the only fossil species of Euglyphostoma now known, is named for A. A. Olsson as a 

tribute to his long and exceptionally productive career 

as a paleontologist and zoologist. He collected the type 

50 years ago.

Occurrence: Upper part of Gatun formation (middle Miocene), eastern area, localities 173, 176, 177c.

Subgenus Rhiglyphostoma Woodring, n. subgen. 

Type: Glyphostoma xeston Gardner, Miocene, Florida.

Very small to of medium sibe (height 5.5 to 17 mm), 

slender or moderately slender, pillar slightly to decidedly constricted, anal fascicle slightly to strongly 

constricted. Protoconch acute and polygyrate (as in 

type species), or blunt and paucigyrate, last half 

whorl to last whorl carinate. Sculpture consisting of 

moderately wide axial ribs, overridden by spiral 

threads. Anal fascicle bearing exaggerated, or ordinary, arcuate growth threads, or sculptured with fine 

spiral threads. Outer lip varicose, or exceptionally nonvaricose and flaring. Anal sinus narrow, deep, semi-
402 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

circular apex everted. Apertural armature heavy for 

size of shell to weak, or faint. Exceptionally absent on 

some specimens of species that normally have weak, or 

faint, armature. Siphonal canal moderately notched. 

Siphonal fasciole slightly inflated, sculptured with 

closely spaced spiral threads.

As is evident from the description, Rhiglyphostoma 

is interpreted to embrace species of diverse features. It 

includes the smallest species of the genus and also 

some of moderate size. The change from the analfasciole puckers of Glyphostoma s.s. to exaggerated 

growth threads is gradational. In fact, the exaggerated growth threads of some species of Rhiglyphostoma are about as strong as the weak puckers of some 

species of Glyphostoma s.s. Though the range of 

variation in strength of apertural armature is pronounced, RhiglypJiostoma includes all the species of 

the genus that have faint armature. It may be so faint 

that it is barely discernible. The name G-lyphostoma 

zoster mansfieldi (Gardner, 1943 (1944)-48, p. 273, 

pi. 37, fig. 17, 1948; Duplin formation, North Carolina) was proposed for specimens of G. zoster that 

have barely perceptible swellings on the lower part of 

the columellar lip and no armature on the outer lip, 

which is nonvaricose and flaring. The same aberrant 

features are shown by Fargo's Lioglyphostoma woodringi subspecies (Fargo in Olsson and Harbison, 1953, 

p. 406, pi. 21, figs. 6, 6a; Caloosahatchee marl, Florida). 

Normal specimens of both species have weak armature 

on both lips and a varicose outer lip.

Glyphostoma (Rhiglyphostoma) allodapum Woodring, n. sp. 

Plate 60, figure 1; plate 66, figure 6

Very small, moderately slender, pillar and anal 

fasciole slightly constricted. Protoconch acute, 2^- to 

2%-whorled, last whorl carinate. First post-protoconch 

whorl sculptured with 8 narrow axial ribs, a weak 

peripheral spiral thread, and a weak spiral thread 

below periphery, both slightly swollen on ribs. Axial 

ribs of succeeding whorls progressively wider, 8 on 

each whorl. Spiral threads weak throughout, 4 on 

penult whorl. Anal fasciole bearing only ordinary 

growth threads, generally obscure. Outer lip varicose. 

Anal sinus narrow, deep, semi-circular apex everted. 

Apertural armature heavy for size of shell. Siphonal 

canal moderately notched. Siphonal fasciole slightly 

inflated, sculptured with closely spaced spiral threads.

Height (incomplete) 7 mm (estimated height 8 mm), 

diameter (including outer lip) 3.3 mm (type).

Type: USNM 645835; paratype 645836.

Type locality: 138d (USGS 22016, About 100 meters 

north of Transisthmian Highway and about 75 meters

west of road to refinery site on Payardi Island, Panama ; immediately east of Cativa and 100 meters north 

of locality 138), lower part of Gatun formation.

The type of this small Gatun species evidently is 

mature. The features of the early whorls, missing on 

the type, are shown by the immature paratype. It is 

from the type locality and consists of the protoconch 

and four post-protoconch whorls. The early whorls are 

shown also by another minute shell collected at locality 

147b.

Though the three specimens constitute a meager 

sample, Glyphostoma allodapum is not closely related 

to any known species. In general plan it resembles 

<#. typhon Gardner (1926-47, p. 350, pi. 42, figs. 11, 

12, 1938), from the Chipola formation of Florida. 

That species is smaller (height 5.5 mm, the smallest 

species of the genus), much more slender, and its anal 

fasciole is more constricted.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, locality 138d. 

Middle part, eastern area, locality 147b.

Subfamily DAPHNELLINAE 

Genus Daphnella Hinds

Hinds, The zoology of the voyage of H.M.S. Sulphur, v. 2, 

Mollusca, p. 25, 1844.

Type (logotype, Herrmannsen, Indicis generum malacozoorum, 

vol. 1, p. 370, 1847): Pleurotoma lymneiformis Kiener, 

living, Florida, Bahamas, and West Indies.

The representation of daphnelline turrids in the 

Tertiary Caribbean province is meager. In fact, the 

two unnamed middle Miocene species of Daphnella 

from Bowden, Jamaica, described as Daphnella? sp. 

and "Daphnella" sp. (Woodring, 1928, p. 198-199, pi. 8, 

figs. 15, 16) are the only unequivocal representatives 

heretofore recorded. In addition to the Bowden and 

Gatun specimens, the Cercado formation of the Dominican Republic yielded an immature shell of Daphnella. Though Eocene and Pliocene turrids from southeastern United States have been referred to Daphnella, 

no daphnellines are known in the Tertiary deposits of 

that region.

In outline and aperture the Caribbean fossils are 

notably different from Pleurotoma lymneiformis 

Kiener (1839-40, p. 62, pi. 22, fig. 3), the type species 

of Daphnella. That species is ovate in outline and the 

base of its aperture is wide and flaring. On the contrary, the pillar of the fossils is constricted and the 

aperture ends in a short siphonal canal. Some species, 

however, such as Daphnella bartschi Ball (1919, pi. 

19, figs. 4, 5), are intermediate between D. lymneiformis and these fossil and related living species.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 403

Daphnella pagera Woodring, n. sp. 

Plate 61, figure 2

Of medium size, slender, pillar constricted. Early 

protoconch whorls missing, last whorl of small diameter, bearing diagonally cancellate sculpture. First postprotoconch whorl sculptured with 9 narrow axial ribs, 

overridden by 2 peripheral spiral threads; other narrower threads gradually added. On succeeding whorls 

axial ribs moderately strong to end of penult whorl, 

disappearing on body whorl; 13 on penult whorl. 

Three widely spaced primary spiral threads on penult 

whorl, upper two stronger than other, and many secondary threads, all strongly frosted by fine axial 

threads. Secondary spiral threads and slightly arcuate, protractive axial threads on anal fascicle. Outer 

lip broken back. As shown by axial threads, anal sinus 

narrow, shaped like a reclining L. Aperture relatively 

narrow, siphonal canal short, narrow, not notched, 

according to growth threads. Siphonal fascicle slightly 

inflated, sculptured with slightly frosted, closely spaced 

spiral threads.

Height (almost complete) 12.3 mm, diameter 4.7 mm 

(type).

Type: USNM 135580.

Type locality: 173a (USGS 2689, [French Canal], 

10.5 kilometers from Colon, Canal Zone), upper part 

of Gatun formation.

The type, the only specimen, was collected from the 

upper part of the Gatun formation by E. T. Hill in 

1895.

Daphnella pagera is closely related to an undescribed 

species dredged by Henderson off the coast of southern 

Florida at depths of 16 and 25 fathoms (USNM 

318809, 318806). The fossil is smaller and its primary 

spiral threads are stronger. In outline and sculpture 

both species closely mimic Asperdaphne versivestita 

(Hedley) (1912, p. 148, pi. 43, fig. 33), but the protoconch of that living Australian species is spirally 

sculptured.

Occurrence: Upper part of Gatun formation (middle Miocene), eastern area, locality 173a.

Daphnella species

The lower part of the Gatun formation at locality 

138a yielded a minute, incomplete very slender DapJinella, consisting of the protoconch and four post-protoconch whorls. The protoconch is an acute, wide-based 

cone of 3% whorls, the last 2,y2 of which are diagonally cancellate. The first post-protoconch whorl is 

sculptured with nine axial ribs, overridden by two 

spiral threads. The last whorl bears 12 axial ribs and 

four spiral threads. The anal fascicle is scupltured

with fine spiral threads. The height, minus the aperture, is 3.5 millimeters and the diameter 1.7 millimeters.

The affinities of this species are undetermined.

Occurrence: Lower part of Gatun formation (middle Miocene), locality 138a.

Daphnella? species

A second minute, incomplete unnamed daphhelline 

was collected at the same locality as the preceding 

species, but at a later date. It consists of the protoconch and two post-protoconch whorls. The protoconch 

is an acute, wide-based cone of 4^ whorls, the last 

half whorl of which is subcarinate. Much of the entire 

protoconch is so worn that the sculpture is effaced. It 

can be seen, however, that on the last half whorl the 

part below the subdued carina bears diagonally reticulate sculpture and the part above the carina bears arcuate riblets. Traces of the same kind of sculpture are 

visible on the next to last whorl. The post-protoconch 

whorls are somewhat turreted. They are sculptured 

with 10 strong, narrow, high axial ribs, overridden by 

three (later four) strong, narrow spiral threads. The 

ribs continue in subdued form across the gently sloping anal fascicle. The height, minus the aperture, is 

2.7 millimeters and the diameter 1.5 millimeters.

This daphnelline may represent a new subgenus of 

Daphnella, or a new genus. Its protoconch and cancellarid-like post-protoconch sculpture are unique, at 

least among American daphnellines.

Occurrence: Lower part of Gatun formation (middle Miocene), locality 138.

Family TEREBRIDAE 

Genus Terebra Bruguiere

Bruguiere, Encyclopedic methodique, Histoire naturelle des 

vers, v. 1, p. XV, 1789 (genus without species).

Type (monotype, Lamarck, Soc. Histoire Nat. Paris Mem., p. 71, 

1799): Buccinum subulatum Linne, living, western Pacific Ocean.

The American late Tertiary and living large terebrids are difficult to classify. Every feature that has 

been used to group them into subgenera shows a practically continuous spectrum. Nevertheless, a grouping 

into subgenera is desirable, even though agreement 

concerning boundaries is not to be expected.

In the present account the subgenera are arranged 

in order of increasing loss of sculpture.

Subgenus Oreoterebra Olsson

Olsson, Some Tertiary mollusks from south Florida and the 

Caribbean, p. 15, Paleontological Research Inst, 1967.

Type (orthotype): Terebra mauryw Olsson (:= T. haitensis 

Dall), Miocene, Dominican Republic.
404 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

The type species (Olsson, 1967, p. 15, pi. 4, figs. 8, 

8a) is a synonym of Terebra haitensis Dall (1895 

(1896), p. 35; 1890-1903, pi. 59, fig. 30, 1903), which, 

as shown by topotypes of both, is variable in the 

strength or absence of secondary spiral sculpture. It 

is fairly common in the Gurabo formation, but has 

not been found in the Cercado formation.

Oreotere'bra embraces slender species, including the 

most slender of the large American terebrids. Typically the sculpture of Oreoterebra is strong throughout, 

but is subdued on the late whorls of some species. As 

interpreted in the present report, the sculpture is tripartate or bipartate and the columella is uniplicate or 

biplicate.

The age range of Oreotere'bra is early Miocene to 

the present time. A tripartate, uniplicate species T. 

ftoridana Dall (1889, p. 63; 1902, p. 503, pi. 29, fig. 9; 

Abbott, 1954a, p. 38, pi. 2, fig. 7) is living in the western Atlantic Ocean. A tripartate, biplicate species, described in manuscript by Bartsch,'ranges from Guaymas, Mexico, to Panama.

Terebra (Oreoterebra) dicheres Woodring1, n. sp. 

Plate 50, figure 3

Moderately large, very slender, sculpture bipartate. 

Sutural band wide, separated from remainder of whorl 

by deep, narrow groove. Axial sculpture consisting of 

narrow, closely spaced riblets, retractive on sutural 

band, slightly arcuate on remainder of whorl. Siphonal 

canal not preserved. Columella weakly uniplicate.

Height (body whorl, minus siphonal canal, and three 

preceding whorls) 35 mm, (estimated restored height 

55 mm), diameter 11.5 mm (type).

Type: USNM 645870.

Type locality: 115a (TJSGS 6515, West side of Culebra [Gaillard] Cut, about one-third mile (500 meters) 

north [northwest] of Paraiso, Canal Zone), La Boca 

formation.

This species is represented by two incomplete specimens from the La Boca formation. The shell material 

is replaced by calcite. The sculptural plan is similar 

to that of the much smaller, weakly biplicate Terebra 

odopoia Gardner (1926-47, p. 280, pi. 38, fig. 2, 1938), 

a species from the Chipola and Shoal River formations 

of Florida. The La Boca fossils show no faint secondary spiral sculpture, like that of the Florida fossils, 

but the absence may be a matter of preservation. The 

sutural band of a middle Miocene Colombian species 

(USGS 7857, Tenerife area, Magdalena) is wider than 

that of T. dicheres. The Limon formation at Limon, 

Costa Rica, contains an undescribed, exceptionally 

slender, bipartate, biplicate Oreoterebra, the axial ribs 

of which are practically vertical across the entire whorl.

T. dicheres is the oldest species of Oreotere'bra now 

known. It is considered to be a little older than the 

Chipola species and the forms from the Pirabas formation of Brazil (Maury, 1925a, p. 195,197, 397, pi. 10, 

figs. 5, 9, 10, 11, 13, 18, 19), which probably represent 

one species.

Occurrence: La Boca formation (early Miocene), 

locality 115a.

Terebra (Oreoterebra) subsulcifera subsulcifera 

Brown and Pilsbry

Plate 61, figures 3, 7, 8, 9

Terebra subsulcifera Brown and Pilsbry, Acad. Nat. Sci. Phila.

Proc., v. 63, p. 339, pi. 22, fig. 7, 1911 (Miocene, Canal

Zone). 

Terebra (Myurella) subsulcifera Brown and Pilsbry, Cossmann,

Jour. Conchyliologie, v. 61, p. 14, pi. 1, fig. 25, 1913

(Miocene, Canal Zone). 

Terebra (Myurelllna) subsulcifera Brown and Pilsbry, Olsson,

Neogene mollusks from northwestern Ecuador, Paleontological Research Inst, p. 77, pi. 10, fig. 2, not fig. 2a, 1964

(Miocene, Ecuador). 

fTerebra sp. ind., Toula, K. k. Geol. Reichsanstalt Jahrb., vol.

58, p. 707, pi. 25, fig. 15, 1909 (Miocene, Canal Zone). 

Not Terebra subsulcifera Brown and Pilsbry, Olsson, Bull. Am.

Paleontology, v. 9, no. 39, p. 35, 1922 (Miocene, Canal

Zone). 

Terebra bipartita Sowerby, Olsson, Idem, p. 35, pi. 1, fig. 2,

not fig. 1, 1922 (Miocene, Costa Rica).

Moderately large, very slender, sculpture tripartate. 

Sutural band moderately wide, separated by a groove 

of variable width from a narrow band of variable 

strength; both bands making up about half of spirewhorl height, or a little more. Axial riblets narrow, 

retractive on spiral bands, vertical and straight on remainder of whorl, except on late whorls, where they 

are slightly arcuate. Riblets fading off into low lamellae 

on lower part of body whorl. Weak secondary spiral 

sculpture on some specimens. Siphonal notch deep. 

Siphonal fasciole limited by a narrow thread, bearing 

arcuate axial lamellae. Columella strongly biplicate.

Height (almost complete) 68.5 mm, diameter 12.8 

mm (largest figured specimen). Estimated restored 

maximum height 80 mm, diameter 15 mm.

Type: Acad. Nat. Sci. Phila. 1679.

Type locality: Gatun Locks excavation, middle part 

of Gatun formation.

The nominate subspecies of Terebra subsulcifera 

ranges through the Gatun formation in the Canal 

Zone. Most of the 45 specimens consist of a few whorls, 

generally late whorls that have a damaged body whorl. 

The most nearly complete shell is shown on plate 61, 

figures 8, 9.

Variation affects the degree of slenderness, the 

coarseness of sculpture, the width and depth of the
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 405

groove between the spiral bands, the strength of the 

lower spiral band, especially on the body whorl, the 

spacing of the axial riblets, and the stage at which they 

become slightly arcuate. A coarsely sculptured form is 

shown on plate 61, figure 3. A fragment of seven early 

whorls from locality 155b has exceptionally coarse 

sculpture for that growth stage. The axial riblets of 

the largest lower Gatun shell (12 whorls, height 51 

mm, diameter 12 mm, locality 138c) are nonarcuate up 

to the beginning of the body whorl. Cossmann's 

illustration shows a very narrow band below the 

sutural band and fairly distinct secondary spiral 

sculpture. Toula's poorly preserved specimen is missing from his collection.

T. subsulcifera subsulcifera occurs in the southwestern part of the present Caribbean area and at 

nearby eastern Pacific localities. The coaresly sculptured form occurs in the middle Miocene Angostura 

formation of Ecuador. The living Panamic species 

already mentioned closely resembles the nominate subspecies. The four specimens in the TJSNM collections 

are badly worn. The one in best condition (TJSNM 

123083, Gulf of Panama, 322 fathoms) suggests that 

the sculpture is subdued on the last three whorls.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 138, 138a, 138c, 138d, 138e. Middle part, eastern 

area, localities 139c, 139f, 151, 155, 155a, 155b, 159, 

159d. Upper part, eastern area, localities 175,176,176a, 

177, I77b, I77c. Angostura formation (middle Miocene), Ecuador. Middle Miocene deposits, Costa Rica, 

Colombia (USGS 7852, 10927), Darien (USGS 

8479) and Chiriqui (USGS 7955) areas, Panama.

Terebra (Oreoterebra) subsulcifera cembra Olsson 

Plate 61, figure 5

Terebra (Paratercbra) cembra Olsson, Neogene mollusks from 

northwestern Ecuador, Paleontological Research Inst., 

p. 76, pi. 10, fig. 3, 1964 (Miocene, Ecuador).

Similar to Terebra subsulcifera subsulcifera, but 

sculpture bipartate. Width of sutural band about half 

of spire-whorl height. Axial sculpture fairly coarse. 

Upper end of axial riblets on lower half of earliest 

preserved whorls slightly swollen to form subdued 

nodes. Nodes not connected across space between riblets to form narrow band, like that of nominate subspecies.

Height (almost complete) 48 mm, diameter 10.8 mm 

(figured specimen). Estimated restored maximum 

height 65 mm, diameter 12.5 mm.

Type: USNM 643915.

Type locality: Cueva de Angostura, Rio Santiago, 

Esmeraldas, Ecuador, Angostura formation.

Fourteen specimens from the lower part of the 

Gatun formation are referred to this bipartate subspecies of T. subsulcifera. The axial sculpture is consistently fairly coarse. The earliest preserved whorls 

are almost tripartate. The upper end of the axial ribs 

on the lower half of early whorls of the type are not 

slightly noded, like those of Gatun shells at the same 

stage. That is, the type is strictly bipartate throughout.

A 3i^-whorled, uniplicate shell (height 31 mm), 

more coarsely sculptured than the others, collected at 

locality 138e, is doubtfully referred to T. subsulcifera 

cetnbra. It is more likely to be an aberrant form of that 

subspecies than a different species or subspecies.

T. subsulcifera cembra is known only in Panama 

and Ecuador. The two subspecies occur together at 

localities 138c and 138d (actually two collections of 

different date from the same locality).

Occurrence: Lower part of Gatun formation (middle Miocene), localities 136, 138c, 138d, 138e (identification doubtful), 138f. Angostura formation (middle Miocene), Ecuador.

Terebra (Oreoterebra) isaacpetiti Maury 

Plate 61, figures 12, 13

Terebra Petiti Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 31, 

pi. 4, fig. 4, 1917 (Miocene, Dominican Republic). Not 

T. Petitii Kiener.

Terebra isaacpetiti Maury, Idem, v. 10, no. 42, p. 184 1925 

(Miocene, Dominican Republic). Barrios, Colombia Servicio Geol. Nac., Bol. Geol., v. 6, nos. 1-3 (Informe 1082), 

p. 296, 1960 (Miocene, Colombia).

Terebra isaacpetitif Maury, Weisbord, Bull. Am. Paleontology, 

v. 14, no. 54, p. 52, pi. 6, fig. 3, 1929 (Miocene, Colombia).

Terebra (Paraterebra) isaacpetiti Maury, Perrilliat Montoya, 

Mexico Univ. Nac. Inst. Geologia Paleontologia Mexicana, no. 8, p. 31, pi. 4, figs. 15, 16, 1960 (Miocene, 

Mexico).

Terebra (Oreoterebra) isaacpctiti Maury, Olsson, Some Tertiary 

mollusks from south Florida and the Caribbean, Paleontological Research Inst., p. 16, pi. 3, figs. 4, 4a, 1967 

(Miocene, Dominican Republic).

Large, very slender or slender, tripartate or weakly 

tripartate, finely sculptured. Sutural band moderately 

wide, separated by a deep or shallow groove from a 

well defined or poorly defined narrow band, both bands 

making up a little more than half of spire-whorl 

height. Lower band strong throughout, or subdued 

and even absent on late whorls. Axial riblets narrow, 

closely spaced, retractive on spiral bands, vertical or 

slightly protractive, and straight on remainder of 

whorl, slightly arcuate on some late whorls, disappearing on lower part of body whorl. Weak or faint 

secondary spiral sculpture on some specimens. Columella uniplicate; behind the fold a swelling on some 

specimens.
406 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Height (almost complete) 99.5 mm, diameter 17.5 

mm (larger figured specimen).

Type: Cornell University 36891.

Type locality: Los Quemados, Rio Gurabo, Dominican Republic, Gurabo formation.

This large, tripartite or weakly tripartate, uniplicate, finely sculptured species, which reaches a height 

of 120 mm elsewhere, occurs in the middle and upper 

parts of the Gatun formation. It is represented by 15 

specimens. The largest (pi. 61, fig. 12), a shell from 

the upper part of the Gatun in the western area, is 

worn. The sculpture is shown to better advantage by 

the middle Gatun shell shown on plate 61, figure 13. 

The degree of slenderness and the strength of the 

lower spiral band or its absence are variable. It is 

absent on late whorls of one specimen (locality 155). 

The occurrence of this species in Gatun collections was 

recognized by A. A. Olsson.

Terebra isaacpetiti evidently is rare in the Gurabo 

formation. Only one specimen, of seven whorls, is in 

the USNM collections. Colombian shells are somewhat 

more coarsely sculptured. Those from Tehuantepec 

have slightly arcuate riblets on the lower part of intermediate and late whorls. Some have distinct spiral 

sculpture.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

localities 139c, 147h, 155, 155b, 155c, 157. Upper part, 

eastern area, localities 175, I77a, I77d; western area, 

locality 182, 182a. Gurabo formation (middle Miocene), Dominican Republic. Tubara formation (middle Miocene), Colombia. Agueguexquite formation 

(middle Miocene), Tehuantepec, Mexico.

Subgenus Paraterebra Woodring

Woodring, Carnegie Inst. Washington Pub. 385, p. 135, 1928.

Type (orthotype): Terebra tcxana Dall (= T. flammea Lama.?ck=Epitonium feldmanni ~R6<ling=Buccinuni taurinum Lightfoot), living, Florida, Bahamas, and Gulf of 

Mexico to northern Brazil.

For several decades only two specimens of Terebra 

tewana were recognized: the type, a worn shell collected on Matagorda Island, Texas, and a battered 

kitchen-midden shell from northern Brazil. The first 

dredged specimens came from Barbados (J. B. Henderson in Nutting, 1919, p. 89, pi. 40, fig. 5) and Puerto 

Plata, Dominican Republic (Clench, 1938, p. 114, pi. 

9, figs. 1, 2), identified as T. texana and T. flammea, 

respectively. It was soon realized that T. texana is T. 

flammea (Clench, 1939). Later Buccinwn taurinum 

was found to be the earliest name for this species 

(Render and Abbott, 1951a, p. 66). In the meantime 

it has been dredged at localities off both coasts of

Florida, in the northern Gulf of Mexico, and the 

Caribbean Sea. The depth range of USNM specimens 

is 6 to 20 fathoms.

Paraterebra is distinguished from Oreoterebra by its 

less slender outline and the suppression or loss of 

sculpture on late whorls. In both features, however, 

these subgenera more or less intergrade. The age range 

of Paraterebra is the same as that of Oreoterebra.

Terebra (Paraterebra) species

Of medium size, slender. Early and intermediate 

whorls missing. Upper two-thirds of last three whorls 

inflated. Entire whorl bearing exaggerated growth 

lines, retractive on inflated part, vertical to slightly 

protractive on remaining part. Faint traces of spiral 

sculpture on inflated part. Columella strongly uniplicate.

Height (3 whorls, minus siphonal canal) 20 mm, 

diameter 16.5 mm.

Unlike the condition of most of the aragonite-shelled 

fossils from the Culebra formation, the shell material 

j of the single, incomplete specimen of this species is 

preserved. The inflated upper two thirds of the almost 

smooth whorls indicate that they are mature whorls 

of a small Paraterebra allied to a middle Miocene 

j Venezuelan species identified as Terebra incequdlis 

Sowerby (Jung, 1965, p. 581, pi. 79, figs, 1-3, 9). The 

Venezuelan species, however, evidently is not Sowerby's species (Sowerby, 1850, p. 47). The illustrations 

indicate that it is not similar to the specimen designated the lectotype of T. incequalis (Jung, 1965, pi. 78, 

I fig. 14). It is more like what was selected as the lectotype of T. sulcifera Sowerby (Jimg, 1965, pi. 78, fig. 

13). Some of the labels in the Heneken collection at 

the British Museum (Natural History) are suspect. 

Anyone considering designation of lectotypes for 

species other than those illustrated by Sowerby should 

read Sherborn's remarks in his manuscript register of 

the Heneken collection, mentioned on page 347.

Occurrence: Culebra formation (early Miocene), 

locality lllb.

Terebra (Paraterebra) aff. T. taurina (Lightfoot) 

Plate 61, figures 1, 14

Large, moderately slender, tripartate, finely sculptured. Early whorls missing. Intermediate whorls tripartate, late whorls weakly tripartate. Sutural band 

moderately wide, separated by a relatively shallow 

groove from a poorly defined narrow band set off from 

remainder of whorl by a very shallow groove that almost disappears, or entirely disappears, on late whorls. 

Both bands making up a little more than half of spirewhorl height. Axial riblets narrow and closely spaced,
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 407

or very narrow and very closely spaced, retractive on 

spiral bands, practically vertical and straight on remainder of intermediate whorls, slightly arcuate on 

body whorl, or last three whorls, fading off into 

growth lines on lower part of body whorl. Siphonal 

notch deep. Siphonal fascicle limited by a narrow 

thread, bearing arcuate lamellae and growth lines. 

Columella weakly uniplicate at mature stage, strongly 

uniplicate at earlier stage.

Height (incomplete, 31/2 whorls) 43 mm (estimated 

restored height 95 mm), diameter 16.5 mm (finely 

sculptured figured specimen).

Four incomplete specimens from the middle and 

upper parts of the Gatun formation closely resemble 

Terebra taurina. The degree of slenderness is uniform, 

but one upper Gatun shell (pi. 61, fig. 1) is not as 

finely sculptured as the others, one of which is illustrated (pi. 61, fig. 14). The aperture of the illustrated 

finely sculptured specimen is well preserved, except 

that the outer lip is slightly broken.

So far as these incomplete fossils go, they may 

represent a small subspecies of T. taurina that shows 

a greater range of variation in width and spacing of 

the axial riblets than T. taurina itself, which reaches 

a height of 125 to 145 mm. As shown under occurrence, 

comparable specimens are in collections from middle 

Miocene deposits in southeastern Costa Rica and 

Colombia, and from upper Miocene in Costa Rica, 

Panama, and Venezuela. All of these fossils are incomplete. None exceeds an estimated restored height of 

100 mm. The numbers under occurrence are USGS 

locality numbers.

A fine large species (height 110 mm) that occurs in 

the Globorotalia fohsi barisanensis zone of the Brasso 

formation in Trinidad (USGS 21234, Mayo River) has 

stronger sculpture on late whorls.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

locality 155. Upper part, eastern area, locality 1Y1; 

western area, locality 179. Middle Miocene deposits 

Costa Rica (5882i, 5883e), Colombia (10156, 11325, 

11321, 11348, 11356, 8637, 10102). Limon formation 

(late Miocene), Costa Rica (21036), Bocas del Toro 

area, Panama (8318). Savaneta glauconitic sandstone 

member of Springvale formation (late Miocene), 

Trinidad.

Submenus Panaterebra Olsson

Olsson, Some Tertiary mollusks from south Florida and the 

Caribbean, p. 14, Paleontological Research Inst, 1967.

Type (orthotype): Terebra rolusta Hinds, living, Gulf of California to northern Peru.

The type species loses its sculpture on intermediate 

whorls, but otherwise is similar to Paraterebra. Panaterebra is another subgenus that formerly lived in the 

western Atlantic part of the Tertiary Caribbean province, but now is extinct there and lives in the eastern 

Pacific Ocean.

Terebra (Panaterebra) cucurrupiensis Oinomikado 

Plate 61, figures 10, 11, 15, 16

Terebra (Paraterebra) cucurrupiensis Oinomikado, Geol. Soc.

Japan Jour., v. 46, p. 626, pi. 29, fig. 1, 1939 (Miocene,

Colombia). 

f Terebra (Paraterebra) cf. T. cucurrupiensis Oinomikado,

Marks, Bull. Am. Paleontology, v. 33, no. 139, p. 123, 1951

(Miocene, Ecuador).

Large, early whorls slender, intermediate whorls 

more inflated, late whorls still more inflated, generally 

producing a slightly concave profile. Early whorls 

tripartate. Sutural band moderately wide, separated 

by a groove from a narrow band, both bands making 

up half of spire-whorl height or more than half. Axial 

riblets of variable width and spacing, retractive on 

spiral bands, protractive or vertical on remainder of 

whorl. Spiral bands coalescing, widening, and suppressed at variable stage on intermediate whorls, disappearing on late whorls, but leaving a narrow, slightly depressed area at base of late spire whorls, representing wider part of whorl below bands at earlier 

stage. Axial riblets fading off into growth lines on 

intermediate and late whorls. Faint secondary spiral 

sculpture on some specimens. Siphonal notch and 

fascicle like that of preceding species. Columella 

strongly uniplicate.

Height (incomplete) 75.5 mm (estimated restored 

height 115 mm), diameter 22 mm (largest figured specimen). Height (almost complete) 65.5 mm, diameter 

15.5 mm (figured slender specimen).

Type: Geological Survey of Japan.

Type locality: Rio San Juan, about 6 kilometers 

above Rio Cucurrupi, Choco Procince, Colombia, middle Miocene deposits.

Terebra cucurrupiensis is the most distinctive large 

terebrid in the Gatun formation, distinctive on account 

of the early loss of sculpture and the slightly concave 

profile shown by most of the mature specimens. It is 

especially characteristic of the lower part of the Gatun, 

which yielded 64 of the 73 specimens, but ranges up 

into the middle part.

As partly shown by the illustrations, it is a variable 

species: variable in outline, in the stage at which the 

sculpture disappears, and in width and spacing of the 

axial riblets. Only one specimen (locality 138c, height 

62.5 mm, estimated restored height 85 mm, diameter 

27.5 mm) is exceptionally inflated.

368-278 O - 70 - 8
408 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Oinomikado realized that T. cucurrupiensis is closely 

related to T. galli Dall (1895 (1896), p. 34; 1890- 

1903, pi. 59, fig. 31, 1903), which occurs in the Gurabo 

formation of the Dominican Republic. The type 

(height 68.5 mm) is the only specimen in USNM collections. It is smaller than T. cucurrupiensis, its early 

whorls increase in diameter at a less rapid rate, and 

its combined spiral bands are narrower.

T. loroisii Guerin-Meneville (1854, p. 218, pi. 4, 

fig. 5; not T. loroisi Deshaves, 1859), known only from 

the Pacific coast of Panama, is the only living species 

comparable to T. cucunwpiensis. The living species is 

larger (height up to 150 mm), loses its scuplture at 

an earlier stage, and the combined spiral bands are 

narrower. A few years ago T. loroisii was described as 

a new species: T. dumbauldi Hanna and Hertlein 

(1961, p. 77, pi. 6, fig. 2, pi. 7, figs. 2-5). Hanna and 

Hertlein (p. 72) accepted Reeve's opinion that T. 

loroisii is T. rofousta. Though Guerin-Meneville named 

his species without locality data, his illustration is 

unmistakable, as H. A. Render realized when he identified the species in the USNM collections. Those collections contain six lots, four from Panama, presumably near the city, one from Venado Beach in the Canal 

Zone, and one from the mouth of Rio Maje, in Darien 

near the Gulf of San Miguel. The three earliest lots 

are labelled in Dall's writing Terebra cratera Dall a 

name that was not validated, perhaps because Dall was 

suspicious that such a large, striking species had already been named.

The distribution of T. cucurrupiensis illustrates the 

effects of a branch of the Miocene North Equatorial 

Current sweeping through the Atrato Trough.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 136a, 

137a, 138, 138a, 138c, 138d, 138f, 138g. Middle part, 

eastern area, localities 139c, 139e, 140. Middle Miocene, 

Darien area, Panama (ITSGS 8477), southwestern 

Columbia, Daule formation (middle Miocene), Ecuador (identification doubtful). Early middle Miocene 

part of Brasso formation, Trinidad (USGS 18260).

Genus Strioterebrum Sacco

Sacco, I molluschi del terreni terziarii del Piemonte e della

Liguria, pt. 10, p. 33, 1891. 

Type (orthotype): Terebra bastcroti Nyst, Miocene, western

Europe.

Twenty-six names have been proposed or used for 

species of Strioterebrum from the Tertiary Caribbean 

province. Many of them, however, are difficult to 

recognize and they doubtless are overnamed.

The genus has a meager representation in the Bohio, 

Caimito, Culebra, and La Boca formations and Chagres

sandstone, but is abundant in the Gatun formation. 

Though some 1,100 specimens are in collections from 

the Gatun, three species (8. spiriferum, S. wolfgangi, 

and S. indocayapum) account for 93 per cent of them. 

The number of specimens from the three parts of the 

Gatun is shown in the following table.

Specimens of Strioterebrum from Gatun formation

Species

S. gausapatum (Brown and Pilsbry)______ ___________________

S. indocayapum Olsson

S. oresignum hadrum Woodring, n.

Total____________ _ _ _

Lower 

part

392

4

18

221

39

3

1

678

Middle 

part

45

2

12

125

21

1

206

Upper 

part

127

123

1

251

Total

564

129

30

347

21

39

4

1

1, 135

In addition to the described species the Culebra, La 

Boca, and Gatun formations and Chagres sandstone 

contain unidentified species listed as /Strioterebrum sp.

Strioterebrum listrotum Woodring, n. sp. 

Plate 48, figure 22

Very small, very slender, whorls practically flat. 

Protoconch acute, 3-whorled. Sutural band narrow, 

appearing on third post-protoconch whorl, indistinct 

throughout, owing to very narrow, very shallow groove 

setting it off. Groove hardly indenting axial ribs. Axial 

riblets narrow, closely spaced, slightly protractive. 

Faint traces of spiral sculpture between crests of axial 

riblets below sutural band. Siphonal notch deep. Siphonal fascicle limited by a narrow thread. Columella 

not exposed.

Height 10 mm, diameter 2.3 mm (type).

Type: USNM 645869.

Type locality: 42d (USGS 18837, Barro Colorado 

Island, northern part of island, stream heading west of 

Miller Trail near Miller 17, about 100 meters above 

mouth, Canal Zone), Bohio formation.

The upper part of the Bohio formation of Barro 

Colorado Island yielded the type, the only specimen, 

of this small, very slender, Turbonilla-\Tb& species the 

first Oligocene unequivocal Strioterebrum from the 

Tertiary Caribbean province. It resembles the diminutive S. isclinum (Woodring, 1928, p. 142, pi. 3, fig. 18, 

pi. 4, fig. 1; Bowden formation, Jamaica) which has 

similar obscure spiral sculpture and is even more 

slender.

Occurrence: Upper part of Bohio formation (late 

Oligocene), locality 42d.
GASTROPODS: EULIMIDAE, MARGESTELLIDAE TO HELMINTHOGLYPTIDAE 409

Strioterebrum species

Four specimens of a few whorls each, found in the 

moderately deep-water facies of the Caimito formation on Barro Colorado Island and in a somewhat 

shallower facies on Pato Horqueto Island, represent 

a species that is larger and more coarsely sculptured 

than Strioterebrum listrotum and evidently has a 

wider apical angle. This small sample shows a considerable range of variation in width and spacing of 

the axial riblets.

In general features this inadequate material is similar to larger Miocene species.

Occurrence: Caimito formation, Gatuii Lake area 

(late Oligocene), localities 54h, 54k, 55a.

Strioterebrum cf. S. clethra (Maury) 

Plate 50, figure 4

Of medium size, slender, whorls practically flat. Protoconch not preserved. Sutural band narrow, set off 

by wide, shallow depression. Axial riblets narrow, 

widely spaced, vertical on earliest preserved whorls, 

slightly arcuate on late whorls. Narrow spiral bands 

between crests of riblets below sutural band; on late 

spire whorls uppermost band doubled, other three 

widely spaced. Similar bands in subdued form on 

sutural band of late whorls. Siphonal canal missing. 

Columella not exposed.

Height (5 whorls less siphonal canal) 25.8 mm 

(estimated restored height 40 mm), diameter 9 mm 

(figured specimen).

An incomplete specimen of five late whorls and another of five early whorls from the La Boca formation 

at locality 115b are referred to an unnamed species. 

The smaller specimen shows that the apical angle is 

exceptionally narrow. The depression setting off the 

sutural band is as shallow as the depressions between 

the minor spiral bands below the sutural band. According to Maury's illustration, Strioterebrum clethra 

(Maury 1925a, p. 199, pi. 10, fig. 3), an early Miocene 

Brazilian species, has a narrow apical angle and coarse 

spiral sculpture, but the whorls are slightly bulging; 

the sutural band is stronger and narrower; and the 

axial riblets are narrower. Maury evidently derived 

the trivial name from a Greek noun.

Occurrence: La Boca formation (early Miocene), 

locality 115b.

Strioterebrum spiriferum (Ball) 

Plate 62, figures 25, 31, 32, 36; plate 66, figure 4

Terebra dislocata Say (part), Gabb, Am. Philos. Soc. Trans., 

n. ser., v. 15, p. 225, 1873 (Miocene, Dominican Republic).

Terebra (Acus) bipartita spirifera Dall, U.S. Natl. Mus. Proc., 

v. 18, p. 38, 1895 (1896) (Miocene, Dominican Republic, 

[Canal Zone]).

Terebra (Oxymeris) Mpartita spirifera Dall, Dall, Wagner Free 

Inst. Sci. Trans., v. 3, p. 1,632, pi. 59, fig. 13, 1903 (Oligocene, Dominican Republic [Canal Zone]).

Tcrebra (Strioterebrum) spirifera Dall, Gardner, U.S. Geol. 

Survey Prof. Paper 142, p. 281, pi. 38, figs. 6 (Dall's illustration), 7 (lectotype), 1938 [Miocene, Canal Zone; not 

Shoal River formation and Oak Grove sand member; = . 

Strioterebrum waltonense (Gardner)].

Tercbra spirifera Dall, Barrios, Colombia Servicio Geol. Nac., 

Bol. Geol., v. 6, nos. 1-3 (Informe 1082), p. 298, 1960 

(Miocene, Colombia).

Not Tcrebra spirifera Dall, Maury, Bull. Am. Paleontology, v. 

5, no. 29, p. 24, pi. 3, figs. 15, 16, 1917 [Miocene, Dominican Republic; = Strioterebrum Icevifasciolum (Maury) ?].

Not Terebra spirifera Dall, Olsson, Idem, v. 9, no. 39, p. 38, 

pi. 1, fig. 13, 1922 (Miocene, Costa Rica, Panama).

Not Terebra (Strioterebrum) spirifera Dall, Perrilliat Montoya, 

Mexico Univ. Nac. Inst. Geologia, Paleontologia Mexicana, no. 14, p. 31, pi. 5, figs. 6, 7,1963 (Miocene, Mexico).

Tercbra (Oxymeria) gatunensis Toula, K. k. Geol. Reichsanstalt Jahrb., v. 58, p. 705, pi. 25, fig. 14, 1909 (Miocene, 

Canal Zone).

Tcrebra gatunensis Toula, Brown and Pilsbry, Acad. Nat. Sci. 

Phila. Proc., v. 63, p. 339, pi. 22, fig. 2, 1911 (Miocene, 

Canal Zone). Maury, Bull. Am. Paleontology, v. 5, no. 29, 

p. 31, pi. 4, fig. 5, 1917 (Miocene, Dominican Republic). 

Pilsbry, Acad. Nat. Sci. Phila. Proc., vol. 73, p. 316, 1922 

(Miocene, Dominican Republic). Olsson Bull. Am. Paleontology, v. 9, no. 39, p. 36, pi. 1, figs. 4-6, 1922 (Miocene, 

Canal Zone). Weisbord, Idem, v. 14, no. 54, p. 51, pi. 6, 

fig. 4, 1929 (Miocene, Colombia). Anderson, Calif. Acad. 

Sci. Proc., 4th ser., v. 18, no. 4, p. 106, 1929 (Miocene, 

Colombia). Barrios, Colombia Servicio Geol. Nac., Bol. 

Geol., v. 6, nos. 1-3 (Informe 1082), p. 297, 1960 (Miocene, Colombia).

Tercbra (Myurella) gatunensis Toula, Cossmann, Jour. Conchyliologie, v. 61, p. 13, pi. 1, figs. 26-29, 1913 (Miocene, 

Canal Zone, Martinique).

Tcrebra (Strioterebrum') gatunensis Toula, Oinomikado, Geol. 

Soc. Japan Jour., v. 46, p. 626, pi. 29, fig. 7, 1939 (Miocene, Colombia).

Strioterebrum gatunense (Toula), Olsson, Neogene mollusks 

from northwestern Ecuador, Paleontological Research 

Inst., p. 77, pi. 10, fig. 5 (S. telembiense in explanation 

of plate), 1964 (Miocene, Ecuador).

Strioterebrum cf. gatunense (Toula), Jung, Bull. Am. Paleontology, v. 55, no. 247, p. 558, pi. 60, fig. 3, 1969 (Miocene, 

Trinidad).

Not Terebra gatunensis Maury (error for Toula), Li, Geol. 

Soc. China Bull., v. 9, p. 273, pi. 8, fig. 65, 1930 (Miocene, 

Panama Bay; = Strioterebrum asperum (Hinds), fide 

Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 83, p. 433, 1931, 

living, Panama Bay).

Terebra (Strioterebrum) gatunensis Tcugleri Rutsch, Schweizer. 

Palaeont. Gesell. Abh., v. 54, p. 106, pi. 8 figs. 18, 19, pi. 9, 

figs. 12, 13, 1934 (Miocene, Venezuela).

Not Terebra (Strioterebrum) gatunensis kugleri Rutsch, Weisbord, Bull. Am. Paleontology, v. 42, no. 193, p. 428, pi. 40,
410 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

figs. 12, 13, pi. 45, figs. 24, 25, 1962 [Pliocene, Venezuela; 

= 8trlotereT)rum dislocatum (Say)].

Terebra wolfgangi Toula, Brown and Pilsbry, Acad. Nat. Sci. 

Phila. Proc., v. 63, p. 340, pi. 22, figs. 1, 3-6, 1911 (Miocene, Canal Zone). Maury, Bull. Am. Paleontology, v. 5, 

no. 29, p. 33, pi. 4, fig. 6, 1917 (Miocene, Dominican 

Republic).

Striotereljrum colombianum Olsson, Some Tertiary mollusks 

from south Florida and the Caribbean, Paleontological 

Research Inst, p. 18, pi. 4, figs. 1, la, 1967 (Miocene, 

Colombia).

Large, moderately slender, late whorls flat or slightly 

bulging. Protoconch blunt, iy2 - to 13^-whorled. Sutural 

band appearing on second or third post-protoconch whorl. Sutural band narrow, set off by shallow 

groove, not modifying whorl profile, or slightly 

modifying profile. Axial riblets narrow, closely spaced, 

slightly retractive on sutural band of late whorls, 

slightly arcuate on remainder of whorl; entire pattern 

forming an asymmetric arc on mature whorls. Spiral 

bands below sutural band narrow, slightly swollen on 

crest of riblets. Similar bands absent on sutural band, 

or present in subdued form. Siphonal notch deep, according to growth lamellae on fascicle. Siphonal fasciole limited by a low, narrow thread. Columella uniplicate; behind the fold a swelling of variable strength 

and width. Both fold and swelling indistinct at aperture of some shells, including the largest.

Height (almost complete) 30.1 mm, diameter 7.8 

mm (lectotype). Height (incomplete) 58.5 mm (estimated restored height 72 mm), diameter 14.5 mm 

(large, flat-whoiied figured specimen).

Lectotype: USNM 113654.

Type locality: Recorded as Ponton, Dominican Republic, but that record is rejected in favor of Mount 

Hope, Canal Zone, upper part of Gatun formation.

It is a minor misfortune to lose Toula's name Tere- 

~bra gatunensis, which has been used widely in Caribbean paleontological publications. Many years ago, 

however, Gardner reached the conclusion that it is to 

be suppressed as a junior synonym of T. Mpartita spirifera. As a matter of fact, her decision was more objective than she realized. She had no reason to doubt that 

the type lot of T, ~bipartita spin/era is from Ponton, 

Dominican Republic, as alleged by Dall. Actually there 

is no reasonable doubt that, like the type lot of Phos 

metuloides (p. 266), it is from Mount Hope, Canal 

Zone. Dall identified the Gatun specimens from locality 159 as his form. The type lot of T. ~bipartita spirifera consists of 16 specimens, all incomplete and immature. They can be duplicated by immature shells from 

Mount Hope. The lectotype (pi. 62, fig. 25), selected 

and illustrated by Gardner, is the most complete shell. 

It agrees with the dimension cited by Dall for his 

illustrated specimen, although in his drawing the aperture is restored and the axial riblets are too closely 

spaced, like those of other specimens in the type lot.

This large species, represented by some 560 specimens, is the most abundant Stricterebrum in the Gatun 

formation, especially at locality 138e (185 specimens), 

and ranges through the formation. The typical flatwhorled, finely sculptured form, which includes all the 

mature shells from the middle part of the Gatun, is 

shown on plate 62, figure 32, an illustration of a topotype of Terebra gatunensis. Only 7 percent of almost 

400 specimens collected from the lower part of the 

Gatun resemble the typical form. The others, like that 

illustrated on plate 62, figure 31, are not as flat-whorled 

or as finely sculptured. Some intermediates in the lower 

part (pi. 62, fig. 36), however, are apparent.

The flat, or almost flat, late whorls, sculptural pattern, blunt, cylindrical, paucigyrate protoconch, and 

uniplicate columella are distinctive of Strioterebrum 

spiriferum. The species ranges from early to late Miocene. In the western Atlantic part of the Miocene 

Caribbean province its distribution extends from Trinidad to Panama and includes the Dominican Republic; 

in the eastern Pacific part from Darien to Ecuador. 

Terebra gatunensis kugleri was based chiefly on its 

large size, but in the type region S. spiriferum reaches 

a size as large as Rutch's specimens from Punta Gavilan. S. guana'banum (Olsson) (1942, p. 46, pi. 11, fig. 

4; 1964, p. 78, pi. 10, figs. 1, la, Ib) is a moderately 

deep-water ally, possibly a subspecies, that occurs in 

the late Miocene or early Pliocene Esmeraldas formation of Ecuador and the Pliocene Charco Azul formation of southwestern Panama.

A living species collected at Farfan, at the Pacific 

entrance to the Panama Canal, labelled Terebra radula 

Hinds (USNM 589792), is more similar to S. spiriferum than any living western Atlantic species, but 

its axial riblets are wider. According to Hind's illustration (1845, p. 174, pi. 44, fig. 95), the eastern Pacific 

species is not T. radula.

Occurrence: Lower, middle and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 136, 136a, 137 (fragment, identification doubtful), 

138, 138a, 138b, 138c, 138d, 138e, 138f, 138g, 139 (mold, 

identification doubtful). Middle part, eastern area, 

localities 139c, 141,147g, 155,155b, 155c, 157, 159, 159d, 

160 (preservation poor, identification doubtful); western area, localities 161a, 161d, 162a, 170 (last three 

localities fragments only, identification doubtful). 

Upper part, eastern area, localities 172, 173, 175, 176, 

I76a, I77a, I77b, l77c, l77d, I77e, Dall's collection alleged to be from Ponton, Dominican Republic; western 

area, localities 179 (immature, identification doubtful), 

182, 183 (immature, identification doubtful). Baitoa
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 411

formation (early Miocene), Dominican Republic 

(USGS 8668). Deposits of early Miocene age, Simi 

area, Colombia (USGS 11594, 11612). Cercado formation (middle Miocene), Dominican Republic. Middle 

Miocene deposits, northeastern and southwestern Colombia, Darien, Panama (USGS 8429, 8430). Middle (?) Miocene deposits, Martinique. Angostura formation (middle Miocene), Ecuador. Punta Gavilan 

formation (late Miocene), Venezuela. Melajo clay 

member of Springvale formation (late Miocene), 

Trinidad.

Strioterebrum wolfgangi (Toula)

Plate 62, figures 29, 30; plate 66, figure 1

Terebra (Acus) bipartita Sowerby, variety bipartita s.s. (part), 

Dall, U.S. Natl. Mus. Proc., v. 18, p. 38, 1895 (1896) 

(Miocene, Dominican Republic [Canal Zone]).

Terebra wolfgangi Toula, K. k. Geol. Reichsanstalt Jahrb., 

v. 58, p. 705, pi. 28, fig. 7a, 7b, 1909 (Miocene, Canal 

Zone). Olsson, Bull. Am. Paleontology, v. 9, no. 39, p. 37, 

pi. 1, figs. 11, 12, 1922 (Miocene, Canal Zone).

Not Terebra wolfgangi Toula, Brown and Pilsbry, Acad. Nat. 

Sci. Phila. Proc., v. 63, p. 340, pi. 22, figs. 1, 3-6,1911 [Miocene, Canal Zone; = Strioterebrum spiriferum (Dall)]. 

Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 33, pi. 4, 

fig. 6, 1917 [Miocene, Dominican Republic; = Strioterebrum spiriferum (Dall)].

Terebra gausapata Brown and Pilsbry, Olsson, Idem, v. 9, no. 

39, p. 38, pi. 1, figs. 8, 10, 1922 (Miocene, Canal Zone, 

Panama).

Of medium size, moderately slender to slender, late 

whorls slightly bulging. Protoconch acute, 3- to 3i/£>- 

whorled. Sutural band appearing on first or second 

post-protoconch whorl. Sutural band narrow, set off 

by relatively shallow groove, slightly modifying whorl 

profile. Axial riblets narrow, relatively widely spaced, 

slightly retractive on sutural band of late whorls, 

slightly arcuate on remainder of whorl. Spiral bands 

below sutural band narrow, separated by shallow or 

moderately shallow depressions; slightly swollen on 

axial riblets. Similar bands absent on sutural band, or 

present in subdued form. Apertural features like those 

of Strioterebrum spiriferum.

Height (almost complete) 38.8 mm, diameter 9.5 mm 

(moderately slender figured specimen). Height (incomplete) 37 mm (estimated restored height 42 mm), 

diameter 8.2 mm (slender figured specimen). Height 

(incomplete) 42.7 mm (estimated restored height 55 

mm), diameter 10 mm (largest specimen).

Type: Technische Hochschule, Vienna.

Type locality: Presumably Gatun Locks excavation, 

Canal Zone, middle part of Gatun formation.

Four immature specimens from the lower part of 

the Gatun formation, two immature from the middle 

part, some 110 from four localities in the upper part in 

the Mount Hope area, and 12 additional specimens

from the same area, alleged to be from Ponton in the 

Dominican Republic (Dall's Terebra bipartita) , are 

identified as Strioterebrum wolfgangi. Unfortunately 

no mature specimens from the middle part other than 

the type are available, and the protoconch of the type 

is not preserved. As shown by Toula's illustration, the 

type is a slender form. Though some of the upper 

Gatun fossils, like that shown on plate 62, figure 29, 

are as slender as the type, the prevailing form (pi. 62, 

fig. 30), and the only form in the small lower Gatun 

sample, is less slender. The axial riblets of the upper 

Gatun fossils are not as closely spaced as those of the 

type.

In general features S. wolfgangi is similar to 8. spiriferum, with which it is associated at localities 175, 

I77b, and the locality alleged to be in the Dominican 

Republic. S. wolfgangi, however, is smaller and its 

whorls are not so flat-sided. The protoconch of the 

upper Gatun fossils and of an immature middle Gatun 

specimen is acute and polygyrate, whereas that of 8. 

spiriferum is blunt and paucigyrate, and the axial riblets of mature shells (but not those of the type) are 

more widely spaced.

S. waltonense Gardner (1926-47, p. 282, pi. 38, figs. 

8-10, 1938; Shoal River formation, Fla.) also has an 

acute, polygyrate protoconch, but its whorls are flatsided and its sutural band is wider than that of 8. wolfgangi. The middle Miocene Tehuantepec species identified as S. waltonense (Perrilliat Montoya, 1963, p. 31, 

pi. 5, figs. 9, 10) is similar to the upper Gatun 8. wolfgangi, except for its blunt, paucigyrate protoconch.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, locality 136. Middle part, locality 155, Gatun Locks area 

(Toula's record). Upper part, eastern area, localities 

173a, 174,175,177b, Dall's collection alleged to be from 

Ponton, Dominican Republic. Limon formation (late 

Miocene), Bocas del Toro area, Panama.

Strioterebrum gausapatum (Brown and Pilsbry) 

Plate 62, figures 23, 24

Terebra gausapata Brown and Pilsbry, Acad. Nat. Sci. Phila. 

Proc., v. 63, p. 340, pi. 22, figs. 8, 9, 1911 (Miocene, Canal 

Zone). Barrios, Colombia Servicio Geol. Nac., Bol. Geol., 

v. 6, nos. 1-3 (Informe 1082), p. 297, 1960 (Miocene, 

Colombia).

Not Terebra gausapata Brown and Pilsbry, Olsson, Bull. Am. 

Paleontology, v. 9, no. 39, p. 38, pi. 1, figs. 8, 10, 1922 

[Miocene, Canal Zone, Panama; = Strioterebrum wolfgangi (Toula)].

Not Terebra (Strioterebrum) gausapata Brown and Pilsbry, 

Gardner, U.S. Geol. Survey Prof. Paper 142, p. 281, pi. 38, 

fig. 4, 1937 (1938) (Miocene, Fla.).

Terebra acuaria Toula, K. k. Geol. Reichsanstalt, Jahrb., v. 61, 

p. 505, pi. 31, fig. 19, 1911 (Miocene, Canal Zone).
412 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Terebra (MyurelJa) acunrla Toula, Cossmann, Jour. Conchyliologie, v. 61, p. 12, pi. 1, figs. 19-24, 1913 (Miocene, 

Canal Zone, Martinique).

Terebra gausapata herviderana Spieker, Johns Hopkins Univ. 

Studies in Geol., no. 3, p. 35, pi. 1, fig. 1, 1922 (Miocene, 

Peru).

fTerebra (Strioterebrum) herviderana Spieker, Olsson, Bull. 

Am. Paleontology, v. 19, no. 68, p. 148, pi. 15, figs. 3, 7, 

1932 (Mio.cene, Peru).

Terebra (Strioterebrum) sp. b, Woodring, Carnegie Inst. Washington Pub. 385, p. 140, pi. 3, fig. 15, 1928 (Miocene, 

Jamaica).

Terebra (Strioterebrum) pavonia Olsson, Bull. Am. Paleontology, v. 19, no. 68, p. 146, pi. 15, figs. 8, 9, 1932 (Miocene, 

Peru).

Terebra (Strioterebrum) sp. A, Jung, Idem, v. 49, no. 223, p. 

590, pi. 79, figs. 7, 8, 1965 (Miocene, Venezuela).

Of medium size, slender, late whorls slightly bulging. Protoconch acute, 314=- to 4-whorled. Sutural band 

appearing on first post-protoconch whorl, strong and 

strongly noded on succeeding early whorls. Sutural 

band narrow, set off by deep groove of variable width, 

modifying whorl profile. Axial riblets narrow, relatively widely spaced, slightly retractive on sutural 

band of late whorls, slightly arcuate on remainder of 

whorl. Spiral bands below sutural band few and widely 

spaced, or more numerous and crowded; if numerous, 

that adjoining sutural band wider than others; slightly 

swollen on crest of riblets, or absent on crest. Similar 

bands generally absent on sutural band, exceptionally 

present, as on type. Columella weakly uniplicate; 

behind the fold a wide, low swelling.

Height (incomplete) 33.6 mm (estimated restored 

height 38 mm), diameter 7.5 mm (larger figured specimen). Height (almost complete) 27.3 mm, diameter 

6.7 mm (smaller figured specimen).

Type: Acad. Nat. Sci. Phila. 1678.

Type locality: Gatun Locks excavation, Canal Zone, 

middle part of Gatun formation.

Unlike the preceding two species, /Strioterebrum 

gausapatum is relatively rare. Though it is found in 

the lower and middle parts of the Gatun formation, 

only 30 specimens are assigned to it, and half of them, 

all from the lower part, are immature shells consisting 

of the protoconch and a few post-protoconch whorls. 

The larger figured specimen (pi. 62, fig. 24) is worn.

In protoconch and apertural features S. gausapatum 

is indistinguishable from the slender form of S. wolfgangi. S. gausapatum, however, is consistently slender; 

the sutural band more strongly modifies whorl profile; 

and on early whorls the sutural band is stronger and 

more strongly noded by the axial riblets. Moreover, 

3. gausapatum is the only Gatun species of the genus 

including specimens that have distinct, though weak, 

spiral sculpture on the sutural band. That, however, is

an exceptional feature, shown by only two available 

specimens: the type a small, incomplete shell, illustrated by Brown and Pilsbry in two views of different 

magnification and a somewhat larger topotype (locality 159d).

The type of Terebra acua.ria, like the type of S. 

gamapatum, is small (height 7.6 mm).

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 137, 

138, 138a. Middle part, eastern area, localities 146, 151, 

155b, 155c, 159d. Bowden formation (middle Miocene), 

Jamaica. Middle Miocene deposits, Colombia. Cantaure 

formation (middle Miocene), Venezuela. Zorritos formation (middle Miocene), Peru.

Strioterebrum indoeayapum Olsson 

Plate 62, figure 28; plate 66, figure 10

Strioterebrum indoeayapum Olsson, Neogene mollusks from 

northwestern Ecuador, Paleontological Research Inst., 

p. 79, pi. 11, figs. 2, 2a, 1964 (Miocene, Ecuador).

Of medium size, slender, whorl profile slightly modified by sutural band. Protoconch acute, 3i/£>- to 4i/£>- 

whorled. Sutural band appearing on first or second 

post-protoconch whorl. Sutural band moderately wide, 

slightly bulging, set off by deep, narrow groove. Axial 

riblets narrow, slightly retractive on sutural band of 

late whorls, straight or slightly arcuate, on remainder 

of whorl. Spiral bands below sutural band of variable 

width and spacing, that adjoining sutural band generally wider than others; slightly swollen on crest of 

riblets or absent on riblets, generally absent on sutural 

band. Similar bands exceptionally present in subdued 

form on sutural band of early whorls. Columella 

strongly biplicate, upper fold alined with thread limiting siphonal f asciole.

Height (almost complete) 36.8 mm, diameter 8.7 mm 

(figured specimen). Height (incomplete) 39.2 mm 

(estimated restored height 45 mm), diameter 9 mm 

(largest specimen).

Type: USNM 644128.

Type locality: Telembi, Eio Cayapas, Ecuador, Angostura formation.

Though the Gatun specimens reach a size twice as 

large as those in the type lot, they are identified as 

Strioterebrum indoeayapum. The groove setting off the 

sutural band of the type is wider and shallower than 

on Gatun shells. That feature is variable on other specimens in the type lot, but not on Gatun specimens.

Among the Gatun species of the genus that have an 

essentially similar sculptural pattern, 8. indoeayapum 

is readily distinguished by its acute, polygyrate protoconch, and strongly biplicate columella, strongly 

biplicate even on immature shells of a few post-proto-
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 413

conch whorls. It occurs in the three parts of the Gatun 

formation, a total of some 340 specimens. All of the 

125 specimens from the middle part are small, having 

a restored estimated height of 30 mm or less. In that 

respect they are similar to the shells from the type 

locality. Seventy-four of these small shells, 44 of which 

have preserved protoconchs, were collected at locality 

147b.

Though the sculpture and columella of Olsson's 

Terebra spirifera (Olsson, 1922, p. 38, pi. 1, fig. 13; 

Miocene, Costa Rica, Bocas del Toro area, Panama) 

suggest 8. indocayapum, its spire is long, acute, and 

evenly tapering. Terebra spinfera midiensis (Olsson, 

1922, p. 39, pi. 1, fig. 7; Miocene, Costa Rica) has only 

two spiral bands below the sutural band and strong 

axial threads between them.

At first glance S. indocayapum might be identified 

as S. loBvifasciolum (Maury) (1917, p. 27, pi. 3, fig. 19; 

Cercado formation, Dominican Republic), which 

reaches a size as large as the lower Gatun shells. Its 

protoconch, however, is blunt and paucigyrate, and its 

sutural band is slightly narrower than that of the 

Gatun species. S. Icevifasciolum may be a small subspecies of the common western Atlantic S. dislocatum 

(Say), which has an age range of late Miocene to the 

present time in southeastern United States. In the 

Caribbean region 8. dislocatum is unknown before 

early Pliocene time. The early Pliocene Venezuelan 

species described as 8. gatunense kugleri (Weisbord, 

1962, p. 428, pi. 40, figs. 12, 13, pi. 45, figs. 24, 25) is 

identified as 8. dislocatum.

8. coleri (Engerrand and Urbina) (1910, p. 120, pi. 

59, figs. 35, 36), based on small, incomplete shells from 

middle Miocene deposits near Palenque, Chiapas, may 

be related to 8. dislocatum, as Engerrand and Urbina 

thought, or may be related to 8. indocayapum. Its protoconch is unknown. The same reservation is applicable 

to the small Chipola species 8. pupiforme (Gardner, 

1926^7, p. 280, pi. 38, fig. 3, 1938).

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 137, 138, 138a, 138b, 138c, 138d, 138e, 138f, 138g. 

Middle part, eastern area, localities 139b, 139c, 146, 

147b, 147f, 147g, 147h, 159d; western area, locality 

161c. Upper part, eastern area, locality 178. Middle 

Miocene deposits, Darien, Panama (USGS 8434). Angostura formation (middle Miocene), Ecuador.

Strioterebrum oresignum oresignum Olsson 

Plate 62, figure 37

Strioterebrum oresignum Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst, p. 78, 

pi. 10, figs. 6, 6a, 6b, 1964 (Miocene, Ecuador).

Of medium size, slender, sutural band modifying 

whorl profile. Protoconch not preserved. Sutural band 

relatively wide, set off by narrow, shallow groove. Axial 

riblets narrow, closely spaced, slightly retractive on 

sutural band of late whorls, slightly arcuate on remainder of whorl. Spiral bands below sutural band 

weak to moderately strong, separated by shallow depressions. Similar bands absent on sutural band. Columella smooth, or indistinctly uniplicate and bearing an 

indistinct, wide swelling behind the fold.

Height (incomplete) 43.5 mm (estimated restored 

height 48 mm), diameter 10.2 mm (figured specimen).

Type: USNM 643923.

Type locality: Cueva de Angostura, Rio Santiago, 

Ecuador, Angostura formation.

The somewhat worn figured specimen, the most 

nearly complete shell among 21 found in the middle 

part of the Gatun formation, closely resembles the type 

of Stiioterebrum oresignum, except that the sutural 

band of the Gatun shell is slightly less swollen. Despite 

the wide sutural band of S. oresignum oresignum, well 

shown on mature shells, some immature specimens assigned to that form may be misidentified S. gausapatum.

Occurrence: Middle part of Gatun formation (middle Miocene), eastern area, localities 139c, 139e, 139f, 

155. Angostura formation (middle Miocene), Ecuador.

Strioterebrum oresignum hadrum Woodring, n. subsp. 

Plate 62, figures 22, 35

Sutural band more swollen than that of Strioterebrum oresignum oresignum, producing a turreted 

whorl profile. Axial riblets wider and more widely 

spaced.

Height (incomplete) 46.4 mm (estimated restored 

height 55 mm), diameter 12.8 mm (type). Height (incomplete) 24.2 mm (estimated restored height 35 mm), 

diameter 8.2 mm (figured paratype).

Type: USNM 646045; paratype, USNM 646046.

Type locality: 136 (USGS 16912, north side of 

Transisthmiaii Highway, knoll about 30 meters north 

of highway, 1.2 kilometers northwest of Sabanita, 

Panama), lower part of Gatun formation.

As compared with other Gatun forms, the sutural 

band of this lower Gatun /Strioterebrum is exceptionally swollen and the axial sculpture is coarse. The type, 

like all the other 38 specimens, except the paratype, 

is worn, although the type is one of the few Gatun 

specimens of the genus that has an almost complete 

aperture. Therefore the spiral sculpture below the 

sutural band appears to be weak. A better idea of that 

sculpture is afforded by the immature, incomplete paratype (pi. 62, fig. 22), which is only slightly worn.
414 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

The swollen sutural band suggests alliance to /S. 

nelsoni (Hanna and Israelsky), but the apical angle of 

that species is wider and its axial ribs are distinctly 

arcuate. S. nelsoni occurs in the late Miocene Tumbes 

formation of Peru and in deposits of Pliocene age in 

Ecuador. The type has been illustrated by Pilsbry and 

Olsson (1941, p. 14, pi. 1, fig. 3). S. armillatum 

(Hinds) (in Sowerby, 1845, p. 173, pi. 43, fig. 49) is a 

close living ally of S. nelsoni.

Occurrence: Lower part of Gatun formation (middle 

Miocene), localities 136, 136a, 137a.

Strioterebrum monidum (Woodring) 

Plate 63, figures 3, 4

Terebra (Strioterebrum) monida Woodring, Carnegie Inst. 

Washington Pub. 385, p. 141, pi. 3, fig. 17, 1928 (Miocene, 

Jamaica).

Small, moderately slender, whorls flat. Protoconch 

not preserved. Sutural band exceptionally wide, making up a little less to a little more than half of whorl 

height, set off by deep, narrow groove. Axial ribs wide, 

slightly protractive, separated by narrow V-shaped 

depression, gradually attenuated downward and 

slightly sigmoidal on body whorl. Closely spaced, narrow spiral bands of variable strength on flanks of 

riblets below sutural band, stronger on last few whorls 

than elsewhere. Similar bands present or absent on 

sutural band. Columella uniplicate.

Height (almost complete) 23.5 mm, diameter 5.8 

mm (figured specimen).

Type: USNM 369347.

Type locality: Bowden, Jamaica, Bowden formation.

The exceptionally wide sutural band and closely 

spaced axial ribs of this species are unmistakable. It 

is represented by three specimens from the lower part 

of the Gatun formation and one from the middle part 

in the western area. The sutural band of the lower 

Gatun shells makes up half or a little more than half 

of whorl height, whereas that of the middle Gatun 

shell makes up a little less than half.

The extraordinary number of bore holes, made by 

an unknown predator, in some shells of this species 

from Jamaica and Panama, 1,000 kilometers apart, is 

noteworthy. All of some 20 Jamaican shells have more 

than one hole and one has 13. The illustrated lower 

Gatun shell has 14 holes, only a few of which penetrate 

the entire shell wall. Five and two holes, respectively, 

are shown by the other two lower Gatun shells, but 

the middle Gatun specimen, the smallest of all, has 

none.

Two specimens from the Cercado formation of the 

Dominican Republic represent a slender form of /S. 

monidum, possibly a subspecies. Middle Miocene deposits in southeastern Costa Rica yielded one specimen that has narrower and more numerous axial riblets than those so far mentioned. It presumably is to 

be assigned to a distinct species. The treatment of the 

Cercado and Costa Rican forms is slightly different 

from that adopted in 1928.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138c, 

138d. Middle part, western area, locality 161c. Bowden 

formation (middle Miocene), Jamaica. Cercado formation (middle Miocene), Dominican Republic, slender form (USGS8534).

Strioterebrum aff. S. raptum (Gardner) 

Plate 61, figure 6

Small, slender, late whorls slightly bulging. Protoconch unknown. Sutural band set off only by pits between axial riblets. Axial riblets narrow, closely spaced, 

extending uninterruptedly from suture to suture, protractive, slightly arcuate on late whorls. Spiral bands 

below sutural band weak on last few whorls, indistinct 

or absent on earlier whorls. Similar bands absent on 

sutural band. Columella weakly uniplicate.

Height (almost complete) 10 mm, diameter 3.5 mm 

(figured specimen).

A small, presumably immature, lower Gafrun shell is 

characterized by the weakly set off sutural band set 

off only by pits between the axial riblets. In that feature it is similar to Strioterebrum raptum (Gardner, 

1926-47, p. 285, pi. 38, figs. 18, 19, 1938), a species of 

the Shoal River formation of Florida. The whorl 

height of the Gatun fossil increases at a less rapid rate 

and its spiral sculpture below the sutural band is more 

distinct and coarser. The spiral sculpture is discernible 

on only a few well preserved Florida shells, and even 

on them it is very weak and fine.

Occurrence: Lower part of Gatun formation (middle Miocene), locality 138c.

Family PYRAMIDELLIDAE

The pyramidellids are omitted. With the exception 

of Miocene species from the Dominican Republic, the 

fossil species of that family in the Caribbean region, 

and many of those still living there, are undescribed 

or inadequately covered.

Pyramidellids occur in the marine member of the 

Bohio (?) formation, the Bohio formation, the moderately deep-water facies of the Caimito formation on 

Barro Colorado Island, and the La Boca and Gatun 

formations a total of about 17 species, 12 of which 

are found in the Gatun. Brown and Pilsbry (1913, p. 

509-510, figs. 4a-4c) proposed two names for Gatun 

representatives of Tuvbonilla: T. bartschiana, and T.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 415

gatunensis. They probably are forms of one species. 

Like many other small species, pyramellids are most 

abundant at locality 147b.

Family ACTEONIDAE 

Genus Acteon Montfort

Montfort, Conchyliologie systematique, v. 2, p. 315, 1810. 

Type (orthotype): Voluta tornatilis Gnielin (Voluta tornatilis 

Linne), living, eastern Atlantic Ocean.

Subgenus Acteon s.s. 

Acteon (Acteon) aff. A. tampae Ball

The upper part of the Bohio formation on Barro 

Colorado Island yielded a small, inflated, incomplete 

Acteon. The sculpture, as usual, consists of narrow 

spiral grooves, which are obscurely pitted on the calcite-replaced shell. The inflated outline and small size 

suggest alliance to A. tampce (Dall, 1915, p. 32, pi. 4, 

fig. 10), of the early Miocene Tampa limestone of 

Florida. The body whorl of the type of that species 

the only specimen has two relatively wide spiral 

grooves near the suture.

The estimated restored height of the Bohio fossil is 

7 mm, the estimated restored diameter 4 mm.

Occurrence: Bohio formation (late Oligocene), 

locality 42.

Acteon (Acteon) punctostriatus (C. B. Adams) 

Plate 62, figure 11

Tornatella puncto-striata C. B. Adams, Boston Jour. Nat. History, v. 3, p. 323, pi. 3, fig. 9, 1840 (living, Mass.). Clench 

and Turner, Harvard Univ. Mus. Cornp. Zoology, Occasional Papers on Mollusks, v. 1, p. 333, pi. 40, figs. 5, 6, 

1950 (living, Massachusetts).

Actceon punctostriatus (C. B. Adams), Dall, Wagner Free Inst. 

Sci. Trans., vol. 3, pt. 1, p. 14, 1890 (Miocene to present 

time). Pilsbry, Manual Conchology, v. 15, p. 157, pi. 19, 

figs. 22, 23, 1893 (1894) (living, Massachusetts to West 

Indies).

Acteon punctostriatus (C. B. Adams), Pilsbry, Acad. Nat. Sci. 

Phila. Proc., v. 73, p. 310, 1922 (Miocene, Dominican 

Republic). Olsson and Harbison, Acad. Nat. Sci. Philadelphia Mon. 8, p. 157, 1953 (Pliocene, Florida).

Tornatella, punctata d'Orbigny [Acteon'], in de la Sagra, Ramon, 

Histoire physique, politique et naturelle de 1'ile de Cuba, 

Mollusques, v. 1, p. 230, pi. 17, figs. 10-12, 1842 (living, 

Cuba). Not Acteon punctatus Lea, 1833.

Action cubcnsis Gabb, Am. Philos. Soc. Trans., n. ser., v. 15, 

p. 245, 1873 (Miocene, Dominican Republic).

Tornatella (Actceon) cubemis (Gabb), Guppy, Geol. Soc. London Quart. Jour., v. 32, p. 518, 1876 (Miocene, Dominican 

Republic).

Acteon riomcensis Maury, Bull. Am. Paleontology, v. 5, no. 29, 

p. 11, pi. 3, fig. 1, 1917 (Miocene, Dominican Republic). 

Woodring, Carnegie Inst. Washington Pub. 385, p. 118, 

pi. 2, fig. 3, 1928 (Miocene, Jamaica).

Very small, moderately slender, suture slightly channeled. Early protoconch whorls submerged, about one 

whorl visible, merging into first post-protoconch whorl. 

Sculpture consisting of finely engraved, microscopically pitted, spiral strise, limited to about lower twothirds of body whorl. Columellar fold strong for size 

of shell.

Height 3.7 mm, diameter 2 mm (figured specimen).

Type: Mus. Comp. Zool. 155925.

Type locality: Living, New Bedford, Mass.

Twenty-two specimens of this small species were 

found in the lower and middle parts of the Gatun formation, but half of them are minute shells consisting 

of the protoconch and one or two post-protoconch 

whorls. The largest shell (height almost 7 mm) was 

badly damaged during handling.

If Acteon, punctostriatus now ranges from Massachusetts to the Gulf of Mexico and the West Indies, 

A. riomcensis cannot be distinguished satisfactorily 

from it, contrary to the view that was adopted in 1928. 

The typical form, at the north end of the range, is 

thin shelled and has a weak columellar fold. Lots collected farther south include shells that are thicker and 

have a correspondingly stronger fold. A. riomcensis 

appears to be indistinguishable from such shells. The 

Gatun fossils are more slender than topotypes of A. 

riomcensis and the usual run of dredged shells. Unlike 

some topotypes of A. riomcensis, and also some Jamaican fossils and living shells, none of the few Gatun 

fossils has spiral sculpture extending over the entire 

body whorl.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138a, 

138c. Middle part, eastern area, localities 139b, 139c, 

147b. Cercado formation (middle Miocene), Dominican Eepublic. Bowden formation (middle Miocene), 

Jamaica. Caloosahatchee marl (Pliocene), Florida. 

Living, Massachusetts to Gulf of Mexico and West 

Indies.

Subgenus Lissacteon Monterosato

Monterosato, Naturalista Siciliano, v. 9, p. 188, 1890. 

Type (monotype): Action exilis Jeffreys, living, eastern and 

western North Atlantic Ocean.

Acteon (Lissacteon?) species

Mature size and shape unknown. Exposed part of 

protoconch (one whorl) exceptionally large, almost 

flat topped. End marked by abrupt appearance of 

sculpture. First post-protoconch whorl inflated. Sculpture covering entire whorl, consisting of narrow, 

closely spaced, spiral grooves, pitted by fine axial 

threads. Columella iionplicate.

Height 2 mm, diameter 1.3 mm.
416 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

The minute shells from the middle part of the Gatun 

formation include five specimens, all consisting only of 

the protoconch and the first post-protoconch whorl, of 

a noteworthy Acteon noteworthy for its exceptionally large protoconch and nonplicate columella. The 

large protoconch and strong, abruptly appearing sculpture indicate that it is not closely allied to A. exilis, 

the type of Lissacteon. In fact, it has no known close 

allies.

The validity of Lissacteon as a subgenus is questionable. The type species has a suggestion of a weak columellar fold, but there is no discontinuity in the 

strength of the fold shown by species of the genus.

Occurrence: Middle part of Gatun formation (middle Miocene), eastern area, locality 14/Tb; western area, 

localities 161c, I70a.

Genus Rictaxis Ball

Dall, Am. Jour. Conchology, v. 7, p. 136, 1871.

Type (orthotype): Rictaxis punctoccelata (Carpenter), (Tornatella punctoccelata Carpenter), living, eastern Pacific

Ocean.

Rictaxis oryza (Gabb)

Actceonidea oryza Gabb, Acad. Nat. Sci. Phila. Proc., v. 24, 

(1872), p. 273, pi. 11, figs. 8, 8a, 1873 (Miocene, Dominican Republic). Gabb, Am. Philos. Soc. Trans., v. 15, p. 

245, 1873 (Miocene, Dominican Republic).

Rictaxis oryza (Gabb), Pilsbry, Acad. Nat. Sei. Phila. Proc., 

v. 73, p. 310, pi. 23, fig. 12, 1922 (Miocene, Dominican 

Republic).

Small, slender. Visible part of protoconch consisting 

of one whorl. End marked by appearance of sculpture. 

Sculpture, of narrow, closely spaced, spiral grooves, 

pitted by fine axial threads, covering entire whorl. 

Aperture long, narrow. Columella bearing well up a 

narrow fold, merging downward into long, narrow 

columellar lip.

Estimated restored height 6 mm, diameter 2.7 mm.

Type: Acad. Nat. Sci. Phila. 3181.

Type locality: Dominican Republic, Miocene.

Though this rare species, heretofore represented only 

by the type, is unequivocally identified, the seven specimens all from the lower and middle parts of the 

Gatun formation are unsatisfactory for illustrating. 

Four minute shells show the protoconch and first or 

first and second, post-protoconch whorl. A mature body 

whorl (locality 139c), which has more closely spaced 

spiral grooves than the type, shows the apertural features.

Rictaxis oryza is the monotype of Actceonidea Gabb, 

proposed when the species was described. It is sole 

named Caribbean species of a genus that survived until 

Pliocene time in Florida, where it is represented by 

R. myakkanus (Dall) (Olsson and Harbison, 1953, p.

158, pi. 25, figs. 5, 5a), and is now living in the eastern 

Pacific Ocean. An immature shell from the late Miocene Melajo clay member of the Springvale formation 

of Trinidad (height 3.3 mm), which has a large protoconch, may represent a second Caribbean species.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138c, 

138e. Middle part, eastern area, localities 139b, 139c, 

139g. Miocene (presumably Cercado or Gurabo formation), Dominican Republic.

Family RINGICTTLIDAE 

Genus Ringicula Deshayes

Deshayes in Lamarck, Histoire naturelle des animaux sans

vertebres, 2d eel., vol. 8, p. 342, 1838. 

Type (logotype, Gray, Zool. Soc. London Proc., p. 140, 1847):

Auricula ringens Lamarck, Eocene, western Europe.

Subgenus Ringiculella Sacco

Sacco, I Molluschi del terreni terziarii del Piemonte e della

Liguria, pt. 12, p. 16, 1892.

Type (logotype, Cossmann, Essais de paleoconchologie comparee, pt. 1, p. 115, 1895): Marginella auriculata Menard,

Miocene to present time, western Europe.

Ringicula (Ringiculella?) species

A crushed inflated Ringicula was found in the Caimito formation on Pato Horqueto Island. The outer 

lip is not preserved. Both columellar folds are narrow. 

The parietal callus is thick, but the inner part of the 

upper parietal wall, where the parietal fold of Ringiculella is seated, cannot be exposed without risk of 

damaging the shell. All except the uppermost part of 

the body whorl is sculptured with finely engraved, 

spiral strise. The height is 3.3 mm, and the diameter, 

minus the outer lip, but somewhat increased by crushing, 2.5 mm.

Occurrence: Caimito formation, Gatun Lake area, 

locality 55a.

Ringicula (Ringiculella) semistriata d'Orbigny 

Plate 62, figure 12

Ringicula semi-striata d'Orbigny, in de la Sagra, Histoire physique, politique, et naturelle de Tile de Cuba, Mollusques, 

v. 2, p. 103, pi. 21, figs. 17-19, 1841 (living, Jamaica).

Ringicula semistriata d'Orbigny, Gabb, Acad. Nat. ScL Phila. 

Jour., 2d ser., v. 8, p. 358, 1881 (Pliocene, Costa Rica). 

Pilsbry, Manual Conchology, v. 15, p. 399, pi. 46, figs. 43, 

44, 1893 (1894) (living, Jamaica).

Ringicula tridentata Guppy, Sci. Assoc. Trinidad Proc., vol. 2, 

no. 2, p. 76, 1873 (Miocene, Jamaica) ; reprint, Bull. Am. 

Paleontology, v. 8, no. 35, p. 60, 1921.

Ringicula (Ringiculella) tridentata Guppy, Woodring, Carnegie 

Inst. Washington Pub. 385, p. 132, pi. 2, fig. 22, 1928 

(Miocene, Jamaica).
GASTROPODS: EULIMIDAE, MARGESTELLIDAE TO HELMINTHOGLYPTIDAE 417

Ringicula hypograpta Brown and Pilsbry, Acad. Nat. Sci. Phila. 

Proc., v. 64, p. 505, fig. 2, 1913 (Miocene, Canal Zone).

Small, inflated. Protoconch not distinguishable from 

first post-protoconch whorl. Lower half to two-thirds 

of whorl, exceptionally entire early whorl, sculptured 

with faint, finely engraved, spiral striae. Outer lip 

strongly thickened, inner edge extending toward aperture below middle; or moderately thickened and inner 

edge not extending toward aperture. Parietal callus 

relatively thin. Basal columellar fold wide or narrow. 

Upper columellar and parietal folds narrow.

Height 3 mm, diameter 2 mm (figured specimen).

Type: British Museum (Natural History).

Type locality: Living, Jamaica.

The three parts of the Gatun formation yielded 46 

specimens of Ringicula. All except three are minute or 

otherwise immature. Two of the relatively large shells, 

including that illustrated, have a thick outer lip. The 

other (locality 138c) has a thinner lip and narrower 

basal columellar fold. It is improbable, however, that 

it represents a different species. The entire second or 

third whorl of some minute well-preserved shells is 

faintly sculptured.

No topotypes of R. semistri-ata, are available. The illustrated specimen closely resembles shells from southern Florida that doubtless are to be referred to that 

species, despite the exaggerated apertural features 

shown in d'Orbigny's illustration. It also resembles 

topotypes of R. tridentata, although they have more 

uniformly heavy apertural features. R. tridentata was 

formerly accepted as a valid species.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 138a, 138c, 138d, 138e, 138g. Middle part, eastern 

area, localities 139b, 139c, 139g, 14Yb, 14Yi, 153a, 159d; 

western area, locality I70a. Upper part, eastern area, 

locality 1YY. Gurabo formation (middle Miocene), 

Dominican Republic (USNM 113Y50, USGS 8702). 

Bowden formation (middle Miocene), Jamaica. Moin 

formation (Pliocene), Costa Rica. Living, North Carolina to Gulf of Mexico and West Indies.

Family SCAPHANDRIDAE 

Genus Acteocina Gray

Gray, Zool. Soc. London Proc., p. 160, 1847. 

Type (orthotype): Acteon wctherellii [wetherilli] Lea, Miocene, 

New Jersey.

An incomplete shell and several minute ones from 

the La Boca formation are listed as Acteocina sp.

The type of Acteon ivetherilli apparently the only 

specimen was illustrated by Lea (1833, p. 213, pi. 6, 

fig. 224) and by Richards and Harbison (1942, p. 221, 

pi. 21, fig. 21). Despite the features shown in Lea's

drawing, the shell is now in poor condition: corroded 

and the protoconch is missing. Olsson and Harbison 

(1953, p. 159) thought it is Acteocina canaliculata 

(Say). That species is the type of Utriculastra, proposed as a subgenus of Retusa (Thiele, 1925, p. 269).

The northern A. canaliculata and the southern A. 

candei, which overlap in North Carolina, have frequently been confused. Wells and Wells (1962) found, 

however, that they do not overlap ecologically and 

that they are distinguishable on shell and radular features. Though the type of Retusa (cited on p. 423 of 

present report) differs in shell features and is not 

known to have a radula, they assigned A. canaliculata 

to that genus, as has been done by others.

Acteocina cf. A. bullata (Kiener) 

Plate 48, figures 4, 6

Relatively large, slender, subcylindrical. Spire moderately high. Protoconch small, missing on most specimens. Fold at base of columella of moderate width.

Height 5.3 mm, diameter 2.4 mm (larger figured 

specimen).

The protoconch is preserved on only three of the 25 

specimens of this Acteocina, which occurs in the upper 

part of the Bohio formation of Barro Colorado Island. 

The shell material is replaced by calcite. It probably is 

an early form of A. bullata, the next species described. 

It is unlikely that the calcite would reproduce the faint 

spiral sculpture of that species.

Occurrence: Bohio formation (late Oligocene), locality 42d.

Acteocina bullata (Kiener), small form 

Plate 62, figures 7, 9

Tornatclla bullata Kiener, Species general et iconographie des 

coquilles vivantes, Genre Tornatelle, p. 5, pi. 1, fig. 4, 

1834 (living, West Indies).

Tornatina bullata (Kiener), Pilsbry, Manual Conchology, v. 15, 

p. 183, pi. 22, figs. 17-19, pi. 50, fig. 30, 1893 (1894) (living, Florida Keys to West Indies). Guppy and Dall, U.S. 

Natl. Mus. Proc., v. 19, p. 305, 1896 (Oligocene [Miocene], Jamaica).

Acteocina stibbullata Pilsbry and Johnson, Acad. Nat Sci. Phila. 

Proc., v. 69, p. 150, 1917 (Miocene, Dominican Republic 

[Jamaica]). Pilsbry, Idem, vol. 73, p. 310, fig. 5, 1922 

(Miocene, Dominican Republic [Jamaica]). Woodring, 

Carnegie Inst. Washington Pub. 385, p. 120, pi. 2, fig. 4, 

1928 (Miocene, Jamaica).

Acteocina, cf. subbullata Pilsbry and Johnson, Olsson, Neogene 

mollusks from northwestern Ecuador, Paleontological 

Research Inst., p. 74, pi. 22, fig. 8, 1964 (Miocene, Ecuador).

Acteocina subbullata costaricana Olsson, Bull. Am. Paleontology, v. 9, no. 39, p. 34a, pi. 15, figs. 20-21, 1922 (Miocene, 

Costa Rica).
418 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Kelatively large, slender, subcylindrical. Spire low, 

even so low that almost nothing except protoconch 

shows above body whorl in apertural view. Protoconch 

small. Faint microscopic spiral lineation visible on 

body whorl. Fold at base of columella narrow or moderately narrow.

Height 4.9 mm, diameter 2.2 mm (figured specimen). Height 6.4 mm, diameter 2.7 mm (largest 

specimen).

Type: Presumably at Museum of Natural History, 

Geneva.

Type locality: Living, West Indies.

Eight specimens, collected at localities in the middle 

and upper parts of the Gatun formation, are identified 

as Acteocina buttata. Though the largest are not much 

more than half as large as large living shells, they 

show no basic features to distinguish them from small 

living shells, and some lots of living shells consist of 

specimens not much larger than the Gatun fossils. The 

fossils are similar also to small specimens of the Miocene Jamaican A. subbullata, which reaches a height 

of 10 mm. A. subbullata formerly was accepted as a 

valid species, but that view is now rejected.

Three of the Gatun fossils, including that illustrated, have a low spire, but the spire of the others is 

higher.

Occurrence: Middle, and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

localities 139b, 147b, 147e, 147g, 159d. Upper part, 

eastern area, locality I77b. Thomonde formation (early 

Miocene), Haiti (USGS 9907). Bowden formation 

(middle Miocene), Jamaica. Middle Miocene deposits, 

Costa Rica. Angostura formation (middle Miocene), 

Ecuador. Limon formation (late Miocene), Costa Rica 

USGS 18693). Living, Florida Keys to West Indies.

Acteocina elachista Woodring, n. sp. 

Plate 62, figures 1, 2

Bullina (Tornatina) aff. candei (d'Orbigny), Toula, K. k. Geol. 

Reichsanstalt Jahrb., v. 61, p. 510, pi. 31, fig. 24, 1911 

(Miocene, Canal Zone).

Very small, slender, rounded subcylindrical, spire 

moderately high. Last half of body whorl more or less 

loosely coiled. Protoconch large. Narrow band adjoining suture depressed, its outer edge sharply defined. 

Middle part of outer lip slightly constricted. Fold at 

base of columella narrow.

Height 2.3 mm, diameter 1.2 mm (type).

Type: USNM 646052.

Type locality: 147b (USGS 6033c, Panama Railroad, about 3,500 feet (1,065 meters) southeast of 

Gatun railroad station, Canal Zone), middle part of 

Gatun formation.

If this minute Acteocina were represented by a few 

specimens, it might be appropriate to consider them an 

aberrant small race of the living Caribbean species 

A. candei (d'Orbigny) (Pilsbry, 1893-1894, p. 185, pi. 

22, figs. 21, 22, pi. 50, figs. 27-29, 1894). It is represented, however, by 49 specimens collected at eight 

localities in the lower and middle parts of the formation, as many as 22 at one locality (138a). Moreover, 

the loose coiling of the body whorl of the largest indicates that they are mature. The small size, slender outline, and exceptionally large protoconch are uniform 

features that distinguish these fossils from A. candei. 

The same features distinguish them from the Miocene 

Jamaican fossils described as A. anetaspira (Woodring, 1928, p. 121, pi. 2, fig. 6). It is suggested that 

A. anetaspira be suppressed as a synonym of A. candei.

Toula's specimen is missing in his collection.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 138, 

138a, 138e, 138f. Middle part, eastern area, localities 

147b, 159d; western area, localities 161c, I70a.

Acteocina rusa Gardner 

Plate 62, figures 3, 4

Acteocina rusa Gardner, U.S. Geol. Survey Prof. Paper 142, 

p. 264, pi. 37, fig. 17, 1937 (1938) (Miocene, Florida).

Very small, slender, subcylindrical. Spire moderately 

high, turreted. Last half of body whorl more closely 

coiled than first half. Protoconch large. Post-protoconch spire whorls and uppermost part of body whorl 

sculptured with narrow, low axial threads. Fold at base 

of columella narrow.

Height 2.8 mm, diameter 1.3 mm (figured specimen).

Type: USNM 351030.

Type locality: USGS 3742, Shell Bluff, Shoal River, 

5 or 6 miles west of Mossyhead, Walton County, 

Florida, Shoal River formation.

Two specimens of this axially sculptured Acteocina 

were found in the Gatun formation: a minute shell, 

consisting of the protoconch and first post-protoconch 

whorl (the apertural face of which is missing), in the 

lower part of the Gatun formation and the illustrated 

specimen in the middle part.

The illustrated specimen is a little smaller and more 

slender than the type of A. rusa, and has a higher 

spire. Almost 100 Florida fossils, however, show that 

the height of the spire is variable and that the Gatun 

specimen closely resembles some Florida shells that are 

a little smaller than the type.

A. harpa (Dall) (1871, p. 136, pi. 15, fig. 1, an inadequate representation), living in the eastern Pacific 

Ocean from British Columbia to San Diego, is the only 

species with which A. rusa needs to be compared. A.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 419

harpa is larger; mature, but not immature, shells are 

less cylindrical; and the axial threads are more closely 

spaced and extend down to the middle of the body 

whorl. Moreover, the slender fold at the base of the 

columella disappears 011 mature shells. It seems strange 

that 110 comparable form has turned up in the eastern 

Pacific Panamic province.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, locality 138c. 

Middle part, eastern area, locality 139b. Shoal River 

formation (middle Miocene), Florida.

Genus Cylichnella Gabb

Gabb, Acad. Nat. Sci. Philadelphia Proc., v. 24 (1872), p. 273,

1873. 

Type (monotype): Cylichnella bidentata (d'Orbigny) (Bulla

Mdentata d'Orbigny), living, West Indies.

The radula and anatomy of Cylichnella ~bidentata, 

based on Brazilian specimens that are larger than those 

from the West Indies, were described by Marcus (1958, 

p. 1-9).

Cylichnella atacata stibara Woodring, n. subsp. 

Plate 62, figure 13

Cylichna? aff. Cylichnella Mdendata d'Orbigny (error for Mdentata), Toula, K. k. Geol. Reichsanstalt Jahrb., v. 61, 

p. 510, pi. 31, fig. 25, 1911 (Miocene, Canal Zone).

Moderately large, moderately inflated, rounded subcylindrical. Apex slightly depressed, covered with thin 

coat of callus. About lower third of body whorl sculptured with finely engraved, spiral striae. Upper 

columellar fold strong, basal fold moderately strong.

Height 3.2 mm, diameter 1.8 mm (type).

Type: USNM 646054.

Cylichnella atacata stibara is the most abundant and 

most widely distributed cephalaspid in the Gatun formation. Some 130 of the 365 specimens were collected 

at the type locality and 80 at locality 138c.

It is larger than the nominate subspecies of Cylichnella atacata (Woodring, 1928, p. 124, pi. 2, fig. 9; 

Bowden formation, Jamaica), and the columellar folds 

are stronger, even at the size of the nominate subspecies. On mature shells of C. triticumtritonis 

(Maury) (1917, p. 14, pi. 3, fig. 4; Cercado formation, 

Dominican Republic) the basal columellar fold disappears, or almost disappears. The species now living 

in the Carribbean region, C. bidentata (d'Orbigny) 

(Warmke and Abbott, 1961, p. 144, pi. 27, fig. c), is 

more slender and its basal columellar fold is weak or 

absent. The identification of the living species is longstanding, despite d'Orbigny's illustrations (1841- 

47(?), p. 125, pi. 4, figs. 13-16, 1841), which show

spiral striae over the entire body whorl and a basal 

columellar fold stronger than the upper fold.

Two slender shells from the Gurabo formation of 

the Dominican Republic are identified as C. atacata 

stibara, on the basis of the columellar folds (TJSGS 

8702).

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, localities 137, 138, 138a, 138c, 138d, 138e. Middle part, 

eastern area, localities 139b, 139c, 139f, 139g, 146, 

147b, 147f, 147g, 147h, 147i (identification doubtful), 

155, 155c; western area, localities 161c, 162. Upper 

part, eastern area, localities 163, I73a; western area, 

locality 185. Gurabo formation (middle Miocene), 

Dominican Republic.

Genus Hoxania Leach

Leach, Annals and Mag. Nat. History, v. 20, p. 268, October,

1847. 

Type (monotype) : Bulla cranchii Leach ( Bulla utriculus

Brocchi), Miocene to Pliocene, western Europe, living,

eastern Atlantic Ocean.

Roxania chipolana (Ball) 

Plate 62, figure 14

Bullina (Altderospira) chipolana Dall, U.S. Natl. Mus. Proc.,

v. 18, p. 32, 1895 (1896) (Miocene, Florida, Canal Zone).

Brown and Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 63,

p. 339, 1911 (Miocene, Canal Zone, Ball's record). 

Micromelo (Altderospira) chipolana (Dall), Dall, Wagner Free

Inst. Sci. Trans., v. 3, pt. 6, p. 1,632, pi. 59, fig. 23, 1903

(Oligocene [Miocene], Florida) 

Abderospira, chipolana (Dall), Gardner, U.S. Geol. Survey Prof.

Paper 142, p. 274, pi. 37, fig. 37 (Dall's illustration), 1937

(1938) (Miocene, Florida).

Small, inflated, ovate, tapering toward apical end. 

Apex narrowly perforate. Entire body whorl sculptured with finely engraved, microscopically pitted, 

spiral striae. Sculpture strongest at both ends of whorl, 

faint on middle part of some mature shells. Columellar 

lip detached, forming an umbilical depression.

Height 3.5 mm, diameter 2.2 mm (figured specimen).

Type (lectotype, herewith designated) : USNM 

113894.

Type locality: USGS 2213, 1 mile below Baileys 

Ferry, Chipola River, Calhoun County, Florida, Chipola formation.

Though Roxania chipolana was collected at eight 

localities in the three parts of the Gatun formation, it 

is abundant only at locality 147b (26 specimens). Dall 

identified the specimen from locality 177 as his species. 

It is rare in Florida. In the USNM collections it is 

represented only by the type lot of five specimens. The 

lectotype is the largest in the type lot and was illustrated by Dall. None of the Gatun shells is as large as
420 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

the lectotype (height 4.6 mm). That illustrated is more 

ovate than Florida shells. Others, however, closely 

resemble a small Florida shell.

R. chipolana is the type of Abderospira, proposed by 

Dall when the species was described. The type of 

Roxania is much larger than this fossil species.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, locality 138c. Middle part, eastern area, localities 139c, 146, 

147b, 147g. Upper part, eastern area, localities I73a, 

177, I77c. Chipola formation (early Miocene), Florida.

Genus Scaphander Montfort

Montfort, Conchyliologie systematique, v. 2, p. 335, 1810. 

Type (orthotype): Scaphander lignarius (Linne) (Bulla lignaria Linne), living, eastern Atlantic Ocean.

Submenus Scaphander s.s.

Scaphander (Scaphander) cf. S. jacksonensis Palmer 

Plate 48, figures 1, 2, 12

Of medium size, narrowly ovate, apex depressed. 

Immature shells of moderate size, and presumably mature shells, sculptured with finely engraved, widely 

spaced, spiral striae. On smaller immature shells striae 

of varying spacing limited to both ends of shell, intervening part showing closely spaced, microscopic spiral 

lineation.

Height (incomplete) 15 mm (estimated restored 

height 20 mm), diameter 10.5 mm (largest specimen). 

Height (incomplete) 11 mm (estimated restored 

height 14 mm), diameter 7.5 mm (larger figured specimen).

Eighteen specimens of this Scaphander were found 

in the marine member of the Bohio (?) formation. 

Eleven, however, including the largest, lack, or almost 

completely lack, shell material. The specimens that 

retain shell material range in height from 4 to an estimated 14 mm. One of them, a slender immature specimen, is questionably identified.

Though it evidently is a new species, it would be inappropriate to designate an immature specimen as the 

type. It is more narrowly ovate that is, more subcylindrical than the late Eocene S. jacksonensis 

Palmer (in Harris and Palmer, 1946-47, p. 449, pi. 

64, figs. 6, 7, 1947), but is sculptured like that species. 

It is even more narrowly ovate than the early Oligocene /S. primus Aldrich (1885, p. 148, pi. 2, figs. 7a, 7b), 

and has more closely spaced spiral striae.

The subgeneric name Mirascapha has been proposed 

for a narrowly ovate Eocene species (Stewart, 1927, p. 

438; type (orthotype) : Cylichna costata Gabb, Eocene, 

Calif.). Fossil and living species of Scaphander, however, show a great range in diversity of outline.

Occurrence: Marine member of Bohio (?) formation 

(late Eocene or early Oligocene), localities 40a, 40d, 

41, 41b.

Scaphander (Scaphander) cryus Woodringf, n. sp. 

Plate 48, figures 5, 10, 11

Of medium size, narrowly ovate, apex depressed. 

Sculpture consisting of closely spaced, shallowly engraved, spiral striae and closely spaced, microscopic, 

axial threads, weakly overriding threads between 

striae.

Height 13.5 mm, diameter 7.8 mm (type).

Type: USNM 646093.

Type locality 42d (USGS 18837, Barro Colorado 

Island, northern part of island, stream heading west 

of Miller Trail near Miller 17, about 100 meters above 

mouth, Canal Zone), Bohio formation.

The upper part of the Bohio formation at the type 

locality yielded 25 specimens of Scaphander cryus, all 

replaced by calcite. They range in altitude from 2.5 

to 13.5 mm. Ten additional specimens, which have 

practically no shell material, were collected at locality 

42g.

The axial sculpture, which shows to better advantage on the paratype than on the type, is exceptionally pronounced. It presumably was stronger on the 

original aragonite than on the replacing calcite.

Five poorly preserved fossils from the La Boca 

formation are doubtfully referred to this species.

Occurrence: Upper part of Bohio formation (late 

Oligocene), localities 42d, 42g. La Boca formation 

(early Miocene; identification doubtful), localities 99b, 

115a, 116a.

Family PHILINIDAE

Genus Floribella Woodring, n. gen.

Type. DolaMla aldrichi Dall, early Miocene, 

Florida, Canal Zone, Cuba.

Exceptionally large, reaching height of 80 mm, 

circular ovate. Earliest part (i/2 to 1 whorl) tightly 

coiled, remainder (i/2 to % whorl) separated from 

tightly coiled part by gradually widening gap. Early 

part thick, remainder progressively thinner. Earliest 

parf more or less covered by callus, crudely sculptured 

spirally, remainder bearing low, narrow, closely spaced 

spiral undulations, generally replaced by more or less 

exaggerated growth lines on about last half whorl. 

Callus extending along outer margin of thin part as 

wide, thin band. Aperture greatly expanded. Excessively thin outer lip broken and nicked, growth lines 

showing short projection at upper margin. Inner 

(columellar) margin bordered by swollen ridge on 

thick part of shell, disappearing on thin part.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 421

At first, doubtless under the influence of Dall's assignment, Floribella was thought to be an aplysid related to Dolabella. On further consideration, prompted 

by A. A. Olsson, the conclusion was reached, chiefly 

on the basis of shell outline, that it is a remarkable 

philinid. The shell outline and sculpture of spiral undulations indicate alliance to a much smaller late 

Eocene species from the Paris basin: Philine corrugata Cossmann (1889, p. 315, pi. 11, fig. 16; Cossmann 

and Pissarro, 1910-13, pi. 54, fig. 240-5). The enormous 

size of Floribella and its wide band of callus extending 

along the outer margin of the shell are unique.

Floribella aldrichi (Ball) 

Plate 50, figures 1, 2, 5-8

Dolabella, aldrichi Dall, Wagner Free Inst. Sci. Trans., v. 3, 

p. 18, pi. 10, fig. 7a, 1890 (Miocene, Florida). Gardner, 

U.S. Geol. Survey Prof. Paper 142, p. 276, pi. 37, fig. 45 

(Dall's illustration), 1938 (Miocene, Florida).

Shell from La Boca formation small, only early part 

preserved, thick (7.5 mm, not including inner marginal 

swollen ridge). Swollen ridge strong, 5 mm wide. 

Callus moderately thick. Low, narrow, closely, spaced 

undulations on part not covered with callus.

Height (incomplete) 33.5 mm, diameter (incomplete) 31 mm (figured La Boca specimen). Height (almost complete) 80 mm, diameter (not quite complete) 

72 mm (largest Florida specimen).

Type (lectotype, herewith designated) : USNM 

328453.

Type locality: USGS 2564, Chipola Eiver on McClelland farm, 1 mile below bridge at old Bailey's 

Ferry, Chipola River, Calhoun County, Florida, Chipola formation. Through an error Dall mentioned only 

Alum Bluff (USGS 2211), on Apalachicola Eiver, 

Liberty County, Fla.

Dall had five small, incomplete specimens of his 

Dolabella aldrichi. The specimen used for the drawing 

is designated the lectotype. An additional somewhat 

larger, incomplete shell (USGS 7183, Alum Bluff) 

was available to Gardner, who used Dall's illustration. 

Another small one is in the Aldrich collection, 

formerly at John Hopkins University, but now in the 

U.S. National Museum. In 1951 C. K. Locklin collected 

eight fine shells, five of them large, at the type locality. 

Two of the large shells were nested so neatly that they 

looked like an abnormal double shell. Two of his shells 

are shown on plate 50, figures 5-8. The fragile outer 

lip is missing on all. How far the band of callus extended along the outer margin is indeterminable, as it 

ends in a callus break. Where the inner edge of the 

callus is intact on several shells, the band has a width

of 11.5 to 16.5 mm. An intact inner edge is shown on 

plate 50, figure 6.

The single La Boca shell (pi. 50, figs. 1, 2), replaced 

by calcite, is thick, but the thickness (7.5 mm) falls 

within the range of Florida specimens at the same 

growth stage (3.5 to 8mm). A large mold from the 

Giiines limestone of Cuba (incomplete height 65 mm), 

recognized by Druid Wilson, is identified as Floribella 

aldriclii, although its undulations are coarser than 

those of mainland specimens. Wherever the species 

has been found its age is early Miocene.

Occurrence: La Boca formation (early Miocene), 

locality 115a. Giiines limestone (early Miocene), Cuba 

(USGS 4290, Vento, Habana Province). Chipola formation (early Miocene), Florida.

Family BULLIDAE 

Genus Bulla Linne

Linne, Systema naturae, 10th ed., p. 725, 1758. 

Type (Opinion 196, Internat. Comm. Zool. Nomenclature, 1954): 

Bulla ampulla Linne, living, western Pacific Ocean.

A minute shell from the moderately deep-water 

facies of the Caimito formation on Barro Colorado 

Island is listed as Bulla? sp.

Bulla umbilicata Roding, small form 

Plate 62, figure 18

Bulla paupercula Sowerby, Gabb, Am. Philos. Soc. Trans., n. 

ser., v. 15, p. 246, 1873 (Miocene, Dominican Republic).

Bullaria paupercula (Sowerby), Maury, Bull. Am. Paleontology, 

v. 5, no. 29, p. 18, pi. 3, fig. 8, 1917 (Miocene, Dominican 

Republic). Maury, New York Acad. Sci., Scientific Survey 

of Porto Rico and Virgin Islands, v. 3, pt. 1, p. 75, 1920 

(Miocene, Puerto Rico). Hubbard, Idem, pt. 2, p. 163, 

1920 (Miocene, Puerto Rico). Pilsbry, Acad. Nat. Sci. 

Phila. Proc., v. 73, p. 313, 1922 (Miocene, Dominican 

Republic). Olsson, Bull. Am. Paleontology, v. 9, no. 39, 

p. 34c, pi. 4, fig. 14, 1922 (Miocene, Costa Rica).

Small, almost symmetrically elliptical. Apex narrowly perforate, border of perforation rounded. Wall 

of perforation sculptured with spiral striae. Other 

parts of shell too chalky to show whether they are 

sculptured.

Height 8.5 mm, diameter 5.2 mm (figured specimen).

The widespread small Miocene form of Bulla umbilicata is represented by four chalky and somewhat 

corroded Gatun shells that are even smaller than those 

from other localities, and presumably are immature. 

Better preservation at other localities shows that the 

base of the shell is weakly striate. Many of several 

hundred specimens from the Cercado formation of the 

Dominican Republic retain traces of mottled coloration. They have a maximum height of 16 mm, but 

Maury (1917, p. 236) cited 21 mm for her illustrated
422 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

specimen. Some Costa Rican shells also show traces of 

mottled coloration.

This small form of B. unibilicata has been identified 

as the unillustrated and inadequately described B. 

paupercula Sowerby (1850, p. 52). Sowerby's species, 

however, is large and rounded (height 38 mm, diameter 25.8 mm). It evidently is B. solida Gmelin, which, 

as noted by Pilsbry (1922, p. 313), was named B. 

saraTiberlinerce by Maury (1917, p. 19, pi. 3, fig. 9).

Should it be considered desirable to have a name for 

the small form of B. umMlicata, perhaps B. wnbilicata 

waltonensis Gardner (1926-47, p. 272, pi. 37, figs. 34, 

35, 1938; Shoal River formation, Florida), described 

as a subspecies of B. striata Bruguiere, is available, 

although Florida shells have a narrower apical perforation than those from the Caribbean region.

B. umbilicata (Roding, 1798, p. 15) is traditionally 

known as B. amygdala (Dillwyn, 1817, p. 480). As 

pointed out by Render (1962, p. 585), Roding's name 

has many years precedence.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, locality 136. 

Middle part, eastern area, locality 159d; western area, 

localities 161, 161c. Thomonde formation (early Miocene), Haiti (USGS 9907). Quebradillas (now Aymamon) limestone and Ponce limestone (early Miocene), 

Puerto Rico. Cercado and Gurabo formations (middle 

Miocene), Dominican Republic. Middle Miocene 

deposits, Costa Rica.

Family ATYIDAE 

Genus Atys Montfort

Montfort, Conchyliologie systematique, v. 2, p. 343, 1810. 

Type (orthotype): Atys cymbulus Montfort (= Bulla nancum 

Linne) living, western Pacific Ocean.

Submenus Roxaniella Monterosato

Monterosato, Nornenclatura generica e specifica di alcune conchiglie inediterranee, p. 145, 1884. 

Type (monotype) : Roxaniella jeffreysi (Weinkauff) (OylicJina

jeffreysi Weinkauff), living, Mediterranean Sea.

It is by no means certain that Roxaniella is properly 

allocated as a subgenus of Atys.

Atys (Roxaniella) rhadina Woodring, n. sp. 

Plate 48, figure 17

Of medium size, exceptionally slender, subcylindrical. Apical depression moderately wide. Faint traces 

of spiral striae at both ends of shell. Aperture exceptionally narrow. Outer lip rising well above apex. 

Columellar lip molded against shell wall.

Height 6.8 mm, diameter 2.6 mm (type).

Type: USNM 646095.

Type locality: 42d (USGS 18837, Barro Colorado 

Island, northern part of island, stream heading west of 

Miller Trail near Miller 17, about 100 meters above 

mouth, Canal Zone), Bohio formation.

The type, replaced by calcite, is the only specimen 

of this slender, Oylichna-like species, found in the 

upper part of the Bohio formation of Barro Colorado 

Island. It occurs also in the Mint Spring marl member of the Marianna limestone of Mississippi, although 

the Bohio specimen is slightly more slender than those 

from the Mint Spring. Atys gracUis Dall (Gardner, 

1926-47, p. 270, pi. 37, fig. 30, 1938), of the Miocene 

Alum Bluff group of Florida, is considerably smaller 

(height 5 mm).

The outer lip of both species rises to a greater height 

above the apex than that of the type species of Roxaniella.

Occurrence: Bohio formation (late Oligocene), 

locality 42d. Mint Spring marl member of Marianna 

limestone (middle Oligocene), Mississippi.

Subgenus Aliculastrum Pilsbry

Pilsbry, Manual of Conchology, v. 16, p. 237, 1896. Substitute 

name for Alicula Ehrenberg, 1831, not Alicula Eichwald, 

1830.

Type (monotype of Alicula Ehrenberg): Bulla cylindrica Helbling, living, western Pacific Ocean.

The Caimito formation yielded a small, crushed 

shell listed as Atys sp., and the La Boca formation a 

poorly preserved shell listed as Atys? sp.

Atys (Aliculastrum) eurys Woodring, n. sp. 

Plate 62, figure 10

Of medium size, almost symmetrically elliptical. 

Apical depression moderately wide. Both ends of shell 

sculptured with finely engraved, spiral striae. Outer 

lip rising high above apex. Basal lip slightly drawn 

out along emergence of narrow, slight swelling resembing prosobranch siphonal fascicle. Columellar lip 

detached, enclosing relatively wide umbilicus.

Height 7.3 mm, diameter 3.6 mm (type).

Type:USNM 646057.

Type locality: 159d (USGS 24173, Gatun [presumably Gatun Locks excavation], Canal Zone), middle 

part of the Gatun formation.

Four specimens of Atys eurys were collected in the 

middle part of the Gatun formation, but only the type 

is mature. It is a distinctive species, characterized by 

its high outer lip, siphonal fasciole-like swelling, and 

wide umbilicus. These features, as well as the larger 

size distinguish it from A. dalli (Woodring, 1928, p.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 423

128, pi. 2, figs. 17, 18; Bowden formation, Jamaica). 

A. caribwa (d'Orbigny) (Pilsbry, 1893-94, p. 274, pi. 

48, fig. 12a, 1894), now living in the Caribbean region, 

is similar to A. eurys in outline, but has no siphonal 

fasciole-like swelling and is not umbilicate. In fact, no 

closely allied species is recognized.

Occurrence: Middle part of Gatun formation 

(middle Miocene), eastern area, locality 159d; western 

area, localities 161c, 170a.

Subg-enus. Weinkauffia Monterosato

Monterosoto, Nomenclatura generica e specifica di alcune conchiglie mediterranee, p. 145, 1884. 

Type (monotype): Weinkauffia diaphana (Aradas) (Bulla diaphana Aradas and Maggiore, 1839, not Bulla diaphana

Montagu, 1803, = Bulla turgidula Forbes, 1844), living,

Mediterranean Sea.

Atys (Weinkauffia) cadus Woodring, n. sp. 

Plate 62, figures 8, 16

Very small, inflated, middle part bulging. Apical 

depression wide. All except middle third of shell 

sculptured with spiral striae, finely engraved toward 

middle of shell, wider at extreme ends, especially 

apical end. Outer lip rising high above apex. Apertural 

features like those of Atys eurys, but umbilicus narrower.

Height 2.1 mm, diameter 1.4 mm (type). Height 3.6 

mm, diameter 1.9 mm (paratype).

Type: USNM 646058; paratype, USNM 646059.

Type locality: 139b (USGS 22017, East side of road 

leading from Transisthmian Highway to refinery site 

on Payardi Island, Panama, about 100 meters southwest of refinery gate), middle part of Gatun formation.

Atys cadus is represented by three specimens collected at the type locality. The outer lip of the type is 

broken back. The apertural face of the paratype the 

largest specimen is damaged.

This species is distinguished from A. oedemata Ball 

(Gardner, 1926-47, p. 269, pi. 37, fig. 28, 1938; Chipola 

formation, Florida) by its high outer lip, distinct 

siphonal fasciole-like swelling, and wide umbilicus. A. 

doliolum Maury (1917, p. 18, pi. 3, fig. 7, an inadequate 

illustration; Cercado formation, Dominican Eepublic) 

has no suggestion of a siphonal fasciole-like swelling, 

but has a small, slender fold at the base of the columella. A. sulculorum Pilsbry and Johnson (Pilsbry, 

1922, p. 313, fig. 12) is considered to be a synonym of 

A. doliolum.

Occurrence: Middle part of Gatun formation 

(middle Miocene), eastern area, locality 139b.

Family RETUSIDAE 

Genus Retusa Brown

Brown, Illustrations of the conchology of Great Britain and

Ireland, pi. 38, figs. 1-6, 1827. 

Type (logotype, Gray, Zool. Soc. London Proc., p. 161, 1847):

Bulla obtusa Montagu, living, eastern Atlantic Ocean.

Subgenus Cyiichnina Monterosato

Monterosato, Nomenclatura generica e specifica di alcune conchiglie mediterranee, p. 143, 1884.

Type (logotype, Bucquoy, Dautzenberg, and Dollfus, Les mollusques marins du Rousillon, v. 1, p. 524, 1886): Cylichna 

umMlicata (Montagu) (Bulla umbilicata Montagu, 1803, 

not Bulla umbilicata Rb'ding, 1797,=Volvaria sulcylindrica Brown, 1827), living, eastern Atlantic Ocean.

Poorly preserved shells from the La Boca formation 

are listed as Retusa? sp.

Retusa (Cyiichnina) aff. R. adamsi (Palmer)

Very small, inflated, subcylindrical. Apical depression funnel-shaped. Both ends of shell faintly striate, 

remaining part showing faint, microscopic, spiral 

lineation. Top of outer lip almost flush with apex. 

Basal lip rounded. Columellar lip not exposed.

Height 3.3 mm, diameter 2 mm.

A minute, barrel-shaped shell, replaced by calcite, 

from the marine member of the Bohio(?) formation 

closely resembles Retusa adamsi (Palmer, 1937, p. 482, 

pi. 75, figs. 22, 23) in size and outline. Whether the 

resemblance is as close in features of the columellar 

lip is indeterminable. R. adamsi is a middle Eocene 

species. No comparable late Eocene or Oligocene 

species is known in southeastern United States.

Occurrence: Marine member of Bohio (?) formation 

(late Eocene or early Oligocene), locality 41b.

Retusa (Cyiichnina) quercinensis biforis Pilsbry and Johnson 

Plate 62, figure 6

Retusa biforis Pilsbry and Johnson, Acad. Nat. Sci. Phila. Proc., 

v. 69, p. 151, 1917 (Miocene, Dominican Republic). Pilsbry, Idem, v. 73, p. 311, fig. 8, 1922 (Miocene, Dominican 

Republic).

Very small, moderately slender, subcylindrical. 

Apical depression funnel-shaped. Base faintly striate 

or apparently not sculptured. Outer lip rising slightly 

above apex. Basil lip rounded. Columellar lip detached, 

narrow or wide.

Height 2.8 mm, diameter 1.5 mm (figured specimen).

Type: Acad. Nat. Sci. Phila. 3192.

Type locality: Dominican Kepublic, Miocene, presumably Cercado or Gurabo formation.

Thirty-nine minute shells, collected at localities in 

the middle and upper parts of the Gatun formation 

29 at locality 147b are referred to Retusa biforis.

368-278 O - 70 - 9
424 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

They are regarded as representing a small Caribbean 

race of R. quercinensis Dall (Gardner 1926-47, p. 271, 

pi. 37, fig. 32, 1938). The type of the nominate subspecies is from the Oak Grove sand member of the 

Shoal Kiver formation of Florida. As noted by Gardner, it is an immature shell. Nevertheless her only 

illustration is a reproduction of Ball's drawing. The 

maximum height of mature shells among hundreds of 

topotypes is five millimeters, whereas the type is only 

half as large.

Though the Caribbean subspecies is distinguished 

only by its small size, the Gatun material and the type 

seem to represent a fair sample. The variation in the 

width of the columellar lip is duplicated by Florida 

shells.

A similar species is living in the Caribbean region, 

but is represented in the USNM collections by only one 

shell, dredged off Barbados at a depth of 90 to 100 

fathoms (USNM 502054, height 2.8 mm). It is thinshelled, slender, and slightly more cylindrical than the 

fossils. R. verrillii (Dall) (1889, p. 54; 1902, p. 502, 

pi. 29, fig. 1), from depths of 50 to 124 fathoms off 

North Carolina, is larger than any of the fossils (height 

7.2 mm) and has a very narrow columellar lip.

R. decapitata Dall (Gardner, 1926-47, p. 271, pi. 37, 

fig. 31, 1938; Chipola formation) and R. anthera 

(Gardner) (Idem, p. 271, pi. 37, fig. 33; Shoal Kiver 

formation) are considered to by synonyms of R. 

quercinensis. Despite page precedence, R. quercinensis 

is given precedence over R. decapitata, as it is represented by far more specimens.

Occurrence: Middle and upper parts of Gatun formation (middle Miocene). Middle part, eastern area, 

localities 139b, 147b, 147g, 147h; western area, locality 

I70a. Upper part, eastern area, locality 175. Miocene, 

Dominican Eepublic.

Genus Sulcoretusa Burch

Burch, Conchological Club Southern Calif. Minutes, no. 47, p.

16, 1945. Substitute name for Sulcularia Dall, 1921, not

Sulcularia Rafinesque, 1831. 

Type (orthotype of Sulcularia Dall): Retusa sulcata (d'Orbigny) (Eulla sulcata d'Orbigny), living, North Carolina to

West Indies.

/Sulcoretusa, unlike the other cephalaspid genera 

under consideration, has a limited geographic and age 

range. It is an endemic American genus, now living in 

western Atlantic and eastern Pacific waters, and is uiikiiowii earlier than Miocene.

Sulcoretusa sulcata lipara (Woodring) 

Plate 62, figure 5

Sulcularia lipara Woodring, Carnegie Inst. Washington Pub. 

385, p. 123, pi. 2, fig. 8, 1928 (Miocene, Jamaica).

Moderately small, moderately slender, subcylindrical, 

lower half swollen. Apical perforation funnel-shaped. 

Axial threads narrow, closely spaced. Outer lip rising 

slightly above apex. Columellar lip narrow.

Height 2.7 mm, apical diameter 0.9 mm, basal diameter 1.3 mm (figured specimen).

Type: USNM 369323.

Type locality: Bowden, Jamaica, Bowden formation.

Four names have been proposed for Miocene forms 

of Sulcoretusa: Retusa sulcata fossilis Pilsbry (1922, 

p. 311; Cercado formation, Dominican Eepublic), 

Sulcularia lipara, Sulcularia prosulcata Gardner 

(1926-47, p. 265, pi. 37, fig. 19, 1938; Chipola and 

Shoal Eiver formations, Florida), and Sulcularia sulcata harveyensis Mansfield (1930), p. 27, pi. 1, figs. 6, 7; 

Mansfield's late Miocene Cancellaria zone, Florida). 

The differences between them, as well as those 

distinguishing them from the western Atlantic Sulcoretusa sulcata (d'Orbigny) (Pilsbry, 1893-94, p. 

221, pi. 23, figs. 73, 74, 1894) and the eastern Pacific 

Sulcoretusa paziana (Dall) (1919a, p. 297) are subtle, 

perhaps too subtle. In any event the Miocene forms are 

treated as subspecies of Sulcoretusa sulcata. S. sulcata 

lipara is the most distinctly swollen form.

Nineteen specimens are from the middle part of the 

Gatun formation and one from the lower part. Though 

they are not as uniformly swollen as Jamaican shells, 

they are identified as S. sulcata lipara.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, locality 138a. 

Middle part, eastern area localities 139b, 139c, 139g, 

146, 147b. Bowden formation (middle Miocene), 

Jamaica.

Genus Volvulella Newton

Newton, Systematic list of the Frederick E. Edwards collection 

of British Oligocene and Eocene Mollusca, British Mus. 

(Nat. History), p. 268, 1891. Substitute name for Volvula 

A. Adams, 1850, not Volvula Gistl, 1848. (Newton thought 

Volvula, A. Adams is a homonym of Volvulus Oken, 

1815.)

Type (logotype of Volvula A. Adams, A. Adams, Annals and 

Mag. Nat. History, 3d ser., v. 9, p. 154, 1862): Volvula 

acuminata (Bruguiere) (Bulla acuminata Bruguiere), 

living, eastern Atlantic Ocean. The type designation was 

first recorded by Harry (1967, p. 133).

During recent years RJiizorus Montfort (1810, p. 

338; type (orthotype) : Rliizorus adelaidis Montfort, 

living, Mediterranean Sea) has come into usage for 

this genus, presumably following Iredale's lead (1936, 

p. 332). As implied many years ago by Bucquoy, Dautzenberg, and Dollfus (1882-86, p. 533, 1886), and Dall 

(1889, p. 51), and recently by Harry (1967, p. 133), 

RJiizorus is to be treated as a nomen dubium. In any 

event, Moiitfort's description and illustration of the
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 425

type species is not convincing as a species of Volvulella.

Submenus Volvulella s.s.

Volvulella (Volvulella) aff. V. conradiana (Gabb)

Very small, slender. Apical spine short. Basal third 

of shell sculptured with faint spiral strise.

Height 2.7 mm, diameter 1 mm.

A minute Volvulella, collected at locality 40d in the 

marine member of the Bohio (?) is similar to the 

middle Eocene V. conradiana (Palmer, 1937, p. 494, 

pi. 74, figs. 15-18) in size and outline. It is a little 

more slender than an unnamed Oligocene species in 

tEe Mint Spring marl member of the Marianna limestone and the Byram formation. No comparable species 

is so far recorded from late Eocene deposits in southeastern United States. A more adequate sample is 

needed for identification of the Bohio (?) species.

A more inflated, imperfect specimen (height 2.8 

mm, diameter 1.4 mm) from locality 42 is identified 

as Volvulella f sp.

Occurrence: Marine member of Bohio (?) formation, 

locality 40d.

Volvulella (Volvulella) oxytata (Bush) 

Plate 62, figures 19, 20

fVolvula persimilis Morch, Malak, Blatter, v. 22, p. 179, 1875 

(living, West Indies).

Volvula cylindrica Gabb, Am. Philos. Soc. Trans., n. ser., v. 15, 

p. 246, 1873 (Miocene, Dominican Republic; not Volvula 

cylindrica Carpenter, 1865). Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 16, pi. 3, fig. 5, 1917 (Miocene, Dominican Republic). Olsson, Idem, v. 9, no. 39, p. 34c, 1922 

(Miocene, Costa Rica).

Volvula oxytata Bush, Connecticut Acad. Arts and Sci. Trans., 

v. 6, p. 468, pi. 45, fig. 12, 1885 (living, N. Car.). Dall, 

Harvard Univ. Mus. Comp. Zoology Bull., v. 18, p. 50, 

1889 (living, North Carolina, West Indies). Pilsbry, 

Manual of Conchology, v. 15, p. 235, pi. 26, fig. 63, 1894 

(living, North Carolina to West Indies). Pilsbry, Acad. 

Nat. Sci. Phila. Proc., v. 73, p. 312, 1922 (Miocene, Dominican Republic). Woodring, Carnegie Inst. Washington 

Pub. 385, p. 125, pi. 2, fig. 10, 1928 (Miocene, Jamaica). 

Gardner, U.S. Geol. Survey Prof. Paper 142, p. 267, pi. 

37, figs. 23, 24, 1937 (1938) (Miocene, Florida).

Bulla (Volvula) cf. oxytata Bush, Toula, K. k. Geol.. Reichsanstalt Jahrb., v. 58, p. 709, pi. 28, fig. 4, 1909 (Miocene, 

Canal Zone).

Volvula cercadensis Van Winkle, Bull. Am. Paleontology, v. 8, 

no. 36, p. 6, 1921; new name for Volvula cylindrica Gabb, 

1873, not Volvula cylindrica Carpenter, 1865.

Volvulella persimilis (Morch), Harry, Veliger, v. 10, p. 136, 

fig. 4, 1967 (living, North Carolina to West Indies).

Of medium size, subcylindrical, slender or moderately slender. Apical spine moderately long. Both ends 

of shell sculptured with faint spiral strise, generally

more distinct on basal part than on apical part, or even 

absent on apical part.

Height 3.4 mm, diameter 1.2 mm (figured slender 

specimen). Height 3.6 mm, diameter 1.4 mm (figured 

moderately slender specimen).

Type: Peabody Museum, Yale University.

Type locality: Off Cape Hatteras, N. Car., depth 

7-17 fathoms.

In his 1889 publication Dall expressed the opinion 

that it is highly probable that Volvula oxytata is V. 

persimilis, and Harry recently suppressed V. oxytata 

in favor of V. persimilis. His identification of Morcli's 

species, however, was based chiefly on a process of 

elimination. Until Morch's specimen is found in Copenhagen or London and illustrated, Miss Bush's unequivocal name is preferred.

Volvulella oxytata is a long ranging and widely 

distributed species. It is represented in the Gatun formation by 70 specimens. As for many other small 

species, it is abundant at locality 147b (27 specimens), 

but none from that locality is quite as large as the two 

illustrated. The moderately slender illustrated shell 

has an imperfect outer lip that was damaged and repaired during life. The apical part therefore is abnormal.

Occurrence: Lower and middle parts of Gatun formation (middle Miocene). Lower part, localities 137, 

138, 138a, 138c, 138e, 138g. Middle part, eastern area, 

localities 139b, 139c, 147b, 147g, 147i, 155c; western 

area, localities 161, 161c (identification doubtful), 162, 

170a. Early Miocene part of Uscari shale, Costa Eica. 

Thomonde formation (early Miocene), Haiti (USGS 

9945, 9946). Chipola formation (early Miocene), 

Florida. Middle Miocene deposits, Costa Eica. Cercado 

and Gurabo formations (middle Miocene), Dominican 

Eepublic. Bowden formation (middle Miocene), 

Jamaica. Limon formation (late Miocene), Bocas del 

Toro area, Panama. Duplin formation (late Miocene), 

North Carolina. Caloosahatchee formation (Pliocene), 

Florida. Living, North Carolina to West Indies.

Volvulella (Volvulella) micratracta Brown and Pilsbry

Volvulella micratracta Brown and Pilsbry, Acad. Nat. Sci. 

Phila. Proc., v. 64, p. 504, fig. 1, 1912 (1913) (Miocene, 

Canal Zone).

Very small, inflated. Apical spine moderately long. 

Both ends of shell sculptured with strong, widely 

spaced, spiral strise: three on basal part, two on apical 

part of type.

Height 1.5 mm, diameter 0.7 mm (type).

Type: Acad. Nat. Sci. Phila. 3842.

Type locality: Gatun Locks excavation, Canal Zone, 

middle part of Gatun formation.
426 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

The type is the only Gatun specimen of this inflated, 

distinctive species: distinctive on account of its strong, 

widely spaced, spiral strise. The "appearance of very 

shallow longitudinal plication" on the swollen part of 

the shell, recorded by Brown and Pilsbry, presumably 

is the result of slight difference in shell composition. 

If the type is mature, it is a minute species.

A minute translucent shell, from the Limon formation in Puerto Limon, Costa Rica (USGS 21036), 

is identified as Volvulella micratracta. It is a little 

more slender than the type, and has five basal spiral 

strise and six apical and a more acute apical spine. The 

main part of the shell bears microscopic spiral lineation.

Brown and Pilsbry compared their species with V. 

minuta Bush (1885, p. 469, pi. 45, fig. 11), a similar 

small, inflated western Atlantic species that lacks 

strong, widely spaced, spiral strise, I am unable to 

confirm the apical, axial sculpture shown in Harry's 

(1967, p. 138, fig. 11) drawing of a specimen of V. 

minuta identified by Bush (USNM 44773), and such 

sculpture is not apparent 011 other specimens of her 

species. Therefore Harry's synonymizing of V. minuta 

with the larger, apically sculptured western Atlantic 

species (Bulla acuta d'Orbigny, a junior homonym), 

for which he adopted the name V. recta (Morch), is 

rejected. V. recta, however, is a name of debatable 

availability, as it is based on a lapsus calami.

Occurrence: Middle part of Gatun formation 

(middle Miocene), Gatun Locks excavation (Brown 

and Pilsbry's record). Limon formation (late Miocene), Costa Rica.

Volvulella (Volvulella) cylindrica parallela 

(Pilsbry and Johnson)

Plate 62, figure 17

Volvula parallela Pilsbry and Johnson, Acad. Nat. Sci. Phila. 

Proc., v. 69, p. 151, 1917 (Miocene, Dominican Republic). 

Pilsbry, Idem, v. 73, p. 313, fig. 11, 1922 (Miocene, Dominican Republic).

Moderately large, cylindrical, very slender, height 

three times diameter. Apical spine short. Shell bearing 

microscopic spiral lineation, and at both ends faint 

spiral strise, those on apical part more distinct and 

more widely spaced than those on basal part.

Height 4.7 mm, diameter 1.6 mm (figured specimen).

Type: Acad. Nat. Sci. Phila. 3188.

Type locality: Dominican Republic, presumably 

Cercado or Gurbao formation.

A polished, cylindrical shell from the middle part 

of the Gatun formation (presumably from the Gatun 

Locks excavation) is referred to Volvulella cylindrica 

parallela, which was based on three specimens in 

Gabb's Dominican Republic collection. It is doubtful

whether these Miocene fossils can be distinguished 

from the nominate subspecies of Carpenter's eastern 

Pacific species (Harry, 1967, p. 141, figs. 6-9). They 

closely resemble shells from the Panamic part of the 

range of the nominate subspecies. No similar species 

is living in the Caribbean region.

V. cylindrica parallela occurs also in strata of 

middle Miocene age in southeastern Costa Rica 

(USGS 5882g and other collections from Rio Banana).

Occurrence: Middle part of Gatun formation 

(middle Miocene), eastern area, locality 159d. Miocene, 

presumably Cercado or Gurabo formation, Dominican 

Republic. Middle Miocene deposits, Costa Rica.

Volvulella (Volvulella) phoinicoides (Gardner) 

Plate 62, figure 15

Volvula phoinicoides Gardner, U.S. Geol. Survey Prof. Paper 

142, p. 268, pi. 37, fig. 26, 1937 (1938) (Miocene, Florida).

Of medium size, cylindrical, very slender, height 

three times diameter. Apical spine very short or absent. 

Entire shell bearing microscopic spiral lineation, generally somewhat coarser on basal part. Apical end of 

aperture wide, outer lip extending to apex with gentle 

slope.

Height 3.3 mm, diameter 1.1 mm (figured specimen).

Type: USNM 351037.

Type locality: USGS 3742, Shell Bluff, Shoal River, 

Walton County, Florida, Shoal River formation.

Volvulella phoinicoides has the outline of V. cylindrica parallela^ but differs in apical features and sculpture. Immature shells and mature shells that are 

broken back show that a short spine is present at an 

early stage regardless of its presence or absence at 

maturity. The apical features are more or less intermediate between those of Volvulella s.s. and those of 

the subgenus Paravolvulella (Harry, 1967, p. 141; type 

(orthotype) : Volvulella (Paravolvulella) texasiana 

Harry, living, western Gulf of Mexico). The apical end 

of the outer lip of Paravolvulella extends horizontally 

to the apex of the shell and is slightly concave.

None of the 21 shells from the Gatun formation is as 

large as the largest Florida shells, which reach a height 

of 4.5 mm. This species evidently left no descendents.

Occurrence: Lower, middle, and upper parts of 

Gatun formation (middle Miocene). Lower part, locality 138c. Middle part, eastern area, localities 139b, 

139c, 147b, 147g. Upper part, eastern area, locality 

I77b. Shoal River formation (middle Miocene), 

Florida.

Family SPIRATELLIDAE

Genus Spiratella Blainville

Blainville, Dictionnaire des sciences naturelles, v. 9, p. 407, 

1817.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 427

Type (monotype): Clio helicina Phipps, living, Atlantic and 

Pacific Oceans.

Spiratella inflata elevata (Collins) 

Plate 66, figures 5, 7, 9

Limacina elevata Collins, Johns Hopkins Univ., Studies in 

Geology, no. 11, p. 181, pi. 7, figs. 9-11, 1934 (Miocene, 

Mexico).

Very small, asymmetrically planispiral, body whorl 

moderately inflated. Spire slightly depressed below 

level of body whorl, first whorl slightly projecting. 

Umbilicus very deep, narrow. Apertural part of body 

whorl broken back.

Height 0.5 mm, diameter 0.8 mm (figured specimen).

Type: USNM 645189.

Type locality: USGS 23737, Head of small stream 

flowing into Arroyo Tomas Martinez, about 3 kilometers northeast of Santa Rosa and about 28 kilometers northwest of Santa Lucrecia, Vera Cruz, 

Mexico, deposits of middle Miocene age.

The lower part of the Gatun formation at locality 

136a yielded three minute specimens of Spiratella inflata elevata. Whether this is a valid subspecies is 

questionable. Though the body whorl is less inflated 

than that of the prevailing form of the nominate subspecies, S. inflata elevata occurs with the nominate 

subspecies at the locality near Santa Rosa, in the 

Isthmus of Tehuantepec (Collins, 1934, p. 179, pi. 7, 

figs. 3-8). The nominate subspecies has been found also 

in Miocene deposits in the Dominican Republic (Pilsbry, 1922, p. 308, fig. 1). The inflation of the body 

whorl of the specimen from the Cercado formation 

of the Dominican Republic (USNM 483145), mentioned by Collins (1934, p. 180), is more or less intermediate. Whether intermediates are represented among 

the 40 specimens of S. inflata elevata from the type 

locality is indeterminable at the present time, as the 

collection is temporarily not in Washington.

Occurrence: Lower part of Gatun formation, locality 

136a. Middle Miocene deposits, Mexico.

Family CAVOLINTDAE 

Genus Vaginella Daudin

Daudin, Bull. Sciences, Soc. Philomatique Paris, no. 43, p. 145, 

1800.

Type (monotype): Vaginella depressa Daudin, Miocene, western Europe.

Vaginella lophota Woodring, n. sp. 

Plate 49, figures 10, 11

Of medium size, slender. Posterior part moderately, 

or exceptionally more strongly, inflated, anterior half 

progressively more strongly compressed toward aperture. Sides strongly keeled throughout their length. 

Apertural lips broken.

Height (not quite complete) 5.1 mm, width 2.3 mm, 

diameter 1.4 mm (type).

Type: USNM 646096.

Type locality: 54j (USGS 18833, Barro Colorado 

Island, southwestern part of island, stream crossing 

Conrad Trail at Conrad 2, about 365 meters above 

mouth, Canal Zone), Caimito formation.

This is the first Oligocene American species of 

Vaghietfa, which has an age range of Late Cretaceous 

to Miocene. It is characterized by the strong keel extending along the length of the shell and by the 

marked compression of the anterior half. Eight specimens were found in the moderately deep-water facies 

of the Caimito formation on Barro Coloado Island. 

Little shell material remains, except on the type and 

on a posterior fragment (locality 541) that is more 

strongly inflated than the others.

V. chipolana Dall (Gardner, 1926-47, p. 257, pi. 37, 

figs. 1, 2, 1938; Collins, 1934, p. 214, pi. 13, figs. 16-19; 

Chipola formation, Florida) is larger, wider, more 

inflated, and the keel extends only part way up the 

shell. It is perhaps the same species as V. depressa, the 

type of the genus.

Occurrence: Caimito formation (late Oligocene), 

Barro Colorado Island, localities 54j, 541.

Vaginella undulata (Gabb)

Balantium undulatum Gabb, Am. Philos. Soc. Trans., n. ser., 

v. 15, p. 200, 1873 (Miocene, Dominican Republic).

Vaganella undulata (Gabb) (error for Vaginella), Pilsbry, 

Acad. Nat. Sci. Phila. Proc., v. 73, p. 309, fig. 2, 1922 

(Miocene, Dominican Republic).

Vaginella undulata (Gabb), Collins, Johns Hopkins Univ. Studies in Geology, no. 11, p. 219, pi. 14, figs. 14, 15, 1934 

(Miocene, Dominican Republic).

Vaffinella carib~beana Collins, Idem, p. 220, pi. 14, figs. 16, 17, 

1934 (Miocene, Canal Zone).

Of medium size, very slender, slightly concave, 

clorso-ventrally compressed. Ventral surface bearing 

low undulations, three on type of Vaginella undulata 

and at least five on type of V. caribbeana.

Height 6.5 mm, diameter 1.8 mm (type of V. undulata) .

Type: Acad. Nat. Sci. Phila. 2892.

Type locality: Dominican Republic, presumably 

Cercado or Gurabo formation, Miocene.

Nothing can be added to Collins' description and 

illustration of the fragment that served as the type of 

V. caribbeana (USNM 645196), which is the only 

specimen from the Gatun formation. It has more undulations and somewhat stronger undulations than the 

type (the only specimen) of V. widulata. Those dif-
428 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

ferences surely are not of taxonomic significance. 

Collins suspected he was proposing an unnecessary 

name.

Occurrence: Middle part of Gatun formation 

(middle Miocene), eastern area, locality 159d. Presumably Cercado or Guabo formation (middle Miocene), Dominican Republic.

Genus Cavolina Abilgaard

Abilgaard, Skrivter af Naturhistorie-Selskabet, v. 1, p. 174, 175,

1791.

Type (monotype): Cavolina natans Abilgaard (= Monoculus 

telemus Linne), living, Atlantic and Pacific Oceans.

Subgenus Paleocavolina Woodring, n. subgen.

Type: Cavolina (Paleocavolina) xenica Wooclring, n. sp. Caimito formation, Oligocene, Canal Zone.

Relatively large. Ventral side moderately inflated, 

smooth, little less than apertural half strongly bent. 

At least lateral margins rimmed. Apical and apertural 

margins defective. Dorsal side slightly inflated, end of 

apex missing. Remnants of marginal rim remaining. 

Narrow swollen band adjoining groove setting off 

marginal rim. Apical part bearing low, narrow ridges. 

Traces of similar sculpture 011 preserved patch of shell 

near swollen band close to apical margin.

The combination of rimmed margin, swollen band 

on the dorsal side near the margin, low, narrow ridges 

on the dorsal side, and long, strongly bent ventral side 

distinguishes Paleocavolina from the subgenera Cavolina s.s. and Gamopleura Bellardi (1873 (1872), p. 58; 

type (monotype) : Gamopleura taurinensis (Sismonda) 

(HyalcBa taurinensis Sismonda), Miocene, Italy). The 

length of the apical spine is unknown. No trace of an 

interlocking projection and notch is apparent, but the 

lateral margins are not completely preserved.

Cavolina (Paleocavolina) xenica Woodring, n. sp. 

Plate 49, figures 3, 4, 6, 9

Features of the monotypic subgenus.

Ventral side: height 5.5 mm, width 6.3 mm, diameter 

3.5 mm (type). Dorsal side: height 6.4 mm, estimated 

restored width 6.5 mm, diameter 1.8 mm (paratype).

Type: USNM 646097; paratype, USNM 646098.

Type locality: 54j (USGS 18833, Barro Colorado 

Island, southwestern part of island, stream crossing 

Conrad Trail at Conrad 2, about 365 meters upstream 

from mouth, Canal Zone), Caimito formation.

Locality of paratype: 54h (USGS 18841, Barro 

Colorado Island, western part of island, mouth of first 

stream north of Zetek House, Canal Zone), Caimito 

formation.

Cavolina xenica is represented by a ventral side and 

a dorsal side collected in the moderately deep-water 

facies of the Caimito formation on Barro Colorado 

Island at localities 1.6 kilometers apart (54j and 54h 

respectively; Woodring, 1958, pi. 1). Patches of shell 

material are missing on both sides. It may seem to be 

rash to base a species and subgenus on defective material, but it is a distinctive species.

If Cavolina cookei Simonelli (1895, p. 19, figs, a-c; 

Malta) is of Aquitanian age, C. xenica is the oldest 

known Cavolina. The alleged late Oligocene species 

from Buton (Beets, 1942, p. 305-306, pi. 29, figs. 117- 

123; 1953, p. 251-252, pi. 1, figs. 5-7) are similar to 

Miocene and living species. Their age now is thought 

to be younger than Oligocene (Beets, 1953, p. 239).

Occurrence: Caimito formation (late Oligocene), 

Barro Colorado Island, localities 54h, 54j.

Subgenus Cavolina s.s.

Cavolina (Cavolina) triaspis Woodring, n. sp. 

Plate 63, figures 16, 17

Small. Wide central part of ventral side strongly 

inflated and arched, flanked on each side by a low 

ridge shaped like an inverted V. Apertural margin 

defective. Dorsal side practically flat. Central part 

barely raised, flanked on each side by a low swelling. 

Apertural extremity, which presumably overhung ventral side, missing. Apical spine broken, its base narrow.

Height (not quite complete) 4.2 mm, width 4.5 mm, 

diameter 2.4 mm (type).

Type: USNM 646067.

Type locality: I77d (USGS 6036, Mount Hope, 

about 1/6 mile (270 meters) south of railroad station, 

Canal Zone), upper part of Gatun formation.

The type, from the upper part of the Gatun formation in the Mount Hope area, is the only specimen of 

Cavolina triaspis. It is the second American representative of a group of Miocene western European species 

typified by O. lisulcata (Kittl) (1886, p. 65, pi. 2, figs. 

29-32; Polish Austria) and the very similar G. audeninoi Vinassi de Regny (Sacco, 1904, p. 13, pi. 4, figs. 

2a-2c; Italy). C. audeninoi trinitatis Rutsch (1934, p. 

312, pi. 8, figs. 6-8), from the middle Miocene part of 

the Brasso formation of Trinidad, was the first American form. The ventral side of the European species, 

as well as the Trinidad form, is almost evenly tipartate; that is, the swollen, arch-like central part is 

flanked on each side by a lower, swollen arch. Their 

dorsal side is more strongly sculptured than that of 

C. triaspis.

Occurrence: Upper part of Gatun formation (middle 

Miocene), eastern area, locality I77d.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 429

Cavolina (Cavolina) cf. C. ventricosa (Guppy) 

Plate 63, figures 11, 12

Small, both sides strongly inflated, dorsal side more 

strongly than ventral. Little shell material remaining 

on ventral side. Dorsal side bearing a low, central 

swelling of moderate width, set off by wide, shallow 

depression. Apertural part far overhanging ventral 

side. Posterior lateral margins upturned. Apical spine 

wide-based, bent toward dorsal side, tip missing.

Height 3.9 mm, width 4.3 mm, diameter 3.3 mm 

(figured specimen).

An early collection from the Mount Hope area contains another specimen of Cavolina s.s., evidently representing a new species. It is in poor condition. Though 

much of the dorsal side is preserved, it is badly cracked 

and bits of shell are missing.

The exceptionally strong inflation of the dorsal side 

is the most noteworthy feature of this species. In general features it resembles O. ventricosa (Woodring, 

1928, p. 114, pi. 1, figs. 8, 9; Collins, 1934, p. 184, pi. 7, 

figs. 16-18; Bowden formation, Jamaica), but the dorsal inflation is greater; the central swelling is wider; 

and a pair of minor lateral swellings is absent.

Occurrence: Upper part of Gatun formation (middle 

Miocene), eastern area, locality 177.

Family ELLOBIIDAE 

Subfamily EIXOBIINAE 

Genus Ellobium Roding

Roding, Museum Boltenianum, p. 105,1798.

Type (logotype, Winckworth, Malacolog.. Soc. London, v. 26, 

p. 139, 1945) : Ellobium midce Roding (=Voluta aurlsmidce Gmelin = Bulla aurismidw Linne), living, western 

Pacific Ocean.

Winckworth attributed the type designation to Gray 

(1847, p. 179), but that designation was not in suitable 

form for Ellobium. The type species is gigantic compared with the American species.

Ellobium aff. E. pellucens (Menke) 

Plate 48, figure 24

Of medium size, moderately slender. No trace of 

sculpture apparent. Columella bearing two folds: a 

basal fold and one some distance above it.

Height 17.5 mm, diameter 8 mm (figured specimen).

The La Boca formation yielded the first American 

fossil representative of the salt-marsh pulmonate genus 

Ellobium. It is in poor condition: partly crushed and 

some shell is missing. In outline and lack of sculpture 

it closely resembles immature specimens of the living 

Caribbean species E. pellucens Menke (1828, p. 78). If 

it were known to be mature, it would be appropriate

to consider it to be a small predecessor of the nominate 

form. The original aragonite is replaced by calcite, 

but if this fossil were related to the eastern Pacific 

E. stagnalis (d'Orbigny) (1835, p. 23), af least a trace 

of the fine, crimped sculpture of that sprcies should be 

apparent.

Occurrence: La Boca formation (early Miocene), 

locality 116a.

Family HELMINTHOGLYPTIDAE

Subfamily XANTHONYCINAE

Genus Averellia Ancy

Ancy, [Letter to editor], Conchologists' Exchange, v. 1, p. 54, 

1887. Substitute name for Coelospira Ancy, 1886, not 

Ccelospira Hall, 1863.

Type (orthotype of Ccelospim Ancy, 1886) : Helix macneili 

Crosse, living, Nicaragua and Costa Rica.

Submenus Lecallia Woodring, n. sub gen.

Type: Averellia (Lecallia) stewarti Woodring, n. sp., marine 

member of Bohio(?) formation, late Eocene or early 

Oligocene, Canal Zone.

Of medium size, flattened discoidal, periphery asymmetrically rounded. Umbilicus about a third of shell 

diameter. Whorls 4*4, slowly enlarging in diameter, 

upper surface slightly inflated. Growth lines exaggerated at intervals. Peristome missing.

Lecallia is inferred to be related to Trichodiscina von 

Martens (1890-1901, p. 133, 1892, substitute name for 

Trichodiscus Strebel and Pfeffer, 1880, not Trichodiscus Ehrenberg, 1830; type (logotype of Trichodiscus 

Strebel and Pfeffer, 1880) : Helix coactiliatus Ferrusac, living, northern Mexico to Trinidad). Trichodiscina is currently assigned subgeneric rank under Averellia. The anatomy of the type species of Averellia is 

unknown. The upper surface of the shell is concave 

and the body whorl near the aperture is creased by two 

strong internal lamellae. The jaw and radula of the 

type species of Trichodiscina have been described 

(Baker, 1922, p. 57, pi. 17, figs. 7, 9) and the genitalia 

of a closely related species, A. cordovana (Pfeiffer) 

(Baker, 1927, p. 242, pi. 20, figs. 53-57). Lecallia is 

larger than the species of Trichodiscina and the upper 

surface of its whorls is flatter.

Averellia (Lecallia) stewarti Woodring, n. sp. 

Plate 48, figures 9, 14, 18

Features of the monotypic subgenus.

Height 6 mm, diameter 15.5 mm (type).

Type: USNM 646088.

Type locality: 41b (USGS 18839, East side of Palenquilla Point, head of cove north of triangulation station and southwest of Corozo Island, Canal Zone), 

marine member of Bohio (?) formation.
430 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

The collection from locality 41b includes the type of 

this land snail, associated with 40 marine species. 

Though parts of the shell, which is replaced by calcite, 

and the peristome are missing, it is in reasonably good 

condition and evidently was buried close to the mouth 

of the stream that carried it into the sea. The upper 

edge of the body whorl descends where it is broken off, 

indicating that not much more than the peristome is 

missing on that part of the whorl.

If the affinities of Averellia stewarti have been interpreted properly, it is the first fossil helminthoglyptid from the Caribbean region. In any event it is the 

oldest Central American land snail so far found. The 

species is named for R. H. Stewart, geologist of the 

Panama Canal Company.

Occurrence: Marine member of Bohio (?) formation 

(late Eocene or early Oligocene), locality 41b.

ADDITIONS AND CORRECTIONS TO FAMILIES COVERED IN 

PRECEDING CHAPTERS

Some 20 additional species of gastropods are now 

available, as a result of large collections from the 

Gatuncillo, La Boca, and Gatun formations, received 

after chapter B was prepared. A few of these species 

of special interest, better specimens of some already 

described, and some corrections for the preceding chapters are covered in the following pages. The pages 

cited in headings and citations refer to earlier chapters.

Family TURRITELLIDAE

Genus Turritella Lamarck

Turritella abrupta Spieker (p. 106)

Plate 63, figures 22, 24

Collections from the lower and middle parts of the 

Gatun formation contain larger and better specimens 

of Turritella dbrupta than that shown on plate 23, 

figure 16. All, however, consist of only a few whorls. 

The largest (2y2 whorls, height 56 mm) is estimated to 

have had a height of about 150 mm, not as large as the 

largest from Colombia and Venezuela.

This species is distributed along the south side of the 

Caribbean Sea, thence northward to the Tehuantepec 

area,, and along the Pacific coast from Chiriqui to 

Peri. The specimen from the Tehuantepec area, mentioned on page 107, has been found in the Johns Hopkins University collection, now in the U.S. National 

Museum.

Additional occurrence: Lower and middle parts of 

Gatun formation (middle Miocene). Lower part, localities 138c, 138d, 138e. Middle part, eastern area, localities 139e, 139g. Agueguexquite formation (middle Miocene), Tehuantepec area, Mexico.

Family VERMETIDAE

Genus Pataloconchus H. C. Lea (p. 161)

Petaloconchus sculpturatus H. C. Lea

Plate 63, figure 23

Petaloconchus sculpturatus H. C. Lea, Am. Philos. Soc. Trans., 

n. s., v. 9, p. 233, pi. 34, fig. 3, 1846 (Miocene, Virginia).

Petaloconchus domingensis Sowerby, Geol. Soc. London Quart. 

Jour., v. 6, p. 51, pi. 10, fig. 9, 1850 (Miocene, Dominican 

Republic).

Pctaloconchus aff. P. floridanus Olsson and Harbison, Woodring, present report, p. 161, pi. 29, fig. 9, 1959.

Further sampling of the lower and middle parts of 

the Gatun formation show that the provisionally identified species of Petaloconchus is P. sculpturatus. Specimens that have the normal growth habit of P. sculpturatus, in the form of an open tapering cylinder, and 

at some localities specimens of irregular growth habit, 

were collected at localities 138c, 138e, and 139c. One of 

them is illustrated.

Two specimens from the original lot of P. doming ens is, labelled by Sowerby and formerly in the Calvert collection, are now in the U.S. National Museum 

(560276).

Occurrence: Lower, middle, and possibly upper part 

of Gatun formation (middle Miocene). Lower part, 

localities 138c, 138e. Middle part, eastern area, locality 

139c. For other Gatun localities see p. 162. Widely distributed in deposits of middle and late Miocene age 

from Virginia to Trinidad.

Family THIARIDAE

Subfamily THIARINAE

Genus Hemisinus Swainson (p. 157)

Subgenus?

Hemisinus amaras Woodring, n. sp. 

Plate 63, figures 8, 13

Large, moderately inflated. Shoulder sharply carinate, sutural channel wide, shaped like a shallow, asymmetrical gutter. Early whorls missing. Lower half of 

body whorl sculptured with narrow, flat bands, becoming indistinct above level of aperture. Outer lip broken 

back, slightly sinuate at sutural channel, according to 

growth lines. Basal lip bearing a narrow, shall notch 

adjoining columellar lip.

Height (incomplete) 20.5 mm (estimated restored 

height 27 mm), diameter 12.5 mm (type).

Type: USNM 646071.

Type locality: 161d (USGS 8366, cuts west of Gatun 

Dam, station 3a, Canal Zone), middle part of Gatun 

formation.
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 431

The type the only specimen of this unique species 

of Hemisinus was overlooked in the preparation of 

chapter B. The combination of sharp carina, wide, gutter-shaped sutural channel, and absence of axial sculpture is unique. The basal notch is not as wide and deep 

as it appears to be, as the basal lip is slightly broken 

back. Though truncation of the spire is common in 

thiarids, the spire of this fossil was broken back as 

far as it now is after burial, possibly during collecting. 

Rock matrix is exposed at the broken spire.

H. tnmcatus (Gabb) (Pilsbry, 1922, p. 379, pi. 34, 

figs. 14 16) is the only Caribbean fossil or living species of comparable size and general outline. That species, however, is not carinate or is slightly carinate. It 

has a steeply sloping shelf instead of a gutter-shaped 

sutural channel, and almost, or entirely, microscopic 

spiral sculpture. It has been collected in the Dominican Republic only by Gabb and is the monotype of his 

Ectraclieliza (Gabb, 1872 (1873), p. 271, misprinted 

971) a name of doubtful utility, as indicated by Pilsbry. H. steerei Conrad (1874, p. 32, pi. 1, fig. 14), a 

species in the remarkable fauna of the Pebas formation of late Miocene (?) age on the upper Amazon, has 

a narrower sutural channel and stronger spiral sculpture, according to Conrad's illustration. His type is 

missing.

Occurrence: Middle part of the Gatun formation 

(middle Miocene), western area, locality 161d.

Family CERITHIIDAE (p. 170) 

Subfamily CAMPANILINAE

Genus Campanile Bayle

Bayle, in Fischer, Manuel de conchyliologie et de paleontologie 

conchyliologique, p. 680, 1884.

Type (logotype, Cossmann, Soc. Royale Malacologique Belgique 

Annales, v. 24, p. 25, 1889): GeritMum giganteum Lamarck, Eocene, western Europe.

Campanile is an Italian masculine noun. The generic 

name therefore takes that gender.

The tough matrix encasing the fossil from the La 

Boca formation, mentioned on page 170 as a large cerithid, has yielded to treatment not available when chapter B was prepared. It is not a cerithid, but.is a large 

specimen (4 whorls, height 54 mm, diameter 29 mm) 

of Turritetta cf. T. collazica (p. 98).

Recently acquired collections contain five specimens 

of Campanile from the Emperador limestone member 

of the La Boca formation and a mold of iy2 large 

whorls (height 52 mm, diameter 47 mm) from the La 

Boca proper. Though the mold presumably represents 

Campanile, it is listed as Campanile? sp., as its sculpture is unknown.

Campanile cf. C. herculeanus (Cooke) 

Plate 51, figure 15

Very large for a mid-Tertiary cerithid, slender. 

Early whorls missing. Remaining whorls covered with 

calcareous crust or part of whorl consisting only of 

mold. Molds showing a sutural band and a band near 

lower edge of whorl. Both bands on encrusted whorls 

bearing tubercles, enlarged and distorted by crust. 

Tubercles on sutural band larger than those on lower 

band. Aperture missing, growth lines obscured by crust.

Height (incomplete, 5 whorls) 87.5 mm (estimated 

restored height 140 mm), diameter 54.5 mm (figured 

specimen). Height (incomplete 6 whorls) 104 mm 

(estimated restored height 150 mm), diameter 52 mm 

(largest specimen).

The affinities of this species are indeterminable on 

account of the uneven development of the calcareous 

crust enlarging and distorting the tubercles, and con 

cealing any intervening minor spiral sculpture that 

may have been present. Nevertheless it is reasonably 

certain that the tubercles, especially those on the lower 

band, originally were stronger than those of Campanile herculeanus (Cooke) (1919, p. 116, pi. 1, fig. 7), 

which occurs in the Anguilla formation of the island 

of Anguilla, considered to be of the same age as the 

La Boca formation and its Emperador limestone member. The sculpture of C. herculeanus duplicates that 

of the smaller and more slender Oligocene French species C. charpentieri (Grateloup) (Boussac, 1912, pi. 1, 

figs. 3, 3a, 11). The other mid-Tertiary American species C. halensis (Dall), C. vaughani (Dall) (1916, p. 

512, pi. 87, figs. 9, 10; p. 513, pi. 87, fig. 8, respectively; 

late Oligocene, Georgia), O. hernandcensis (Mansfield), 

G. hernandoensis blackwwterensis (Mansfield) (1937, p. 

150, pi. 6, fig. 3, pi. 7, fig. 8; p. 151, pi. 6, figs. 1, 2, 4; 

late Oligocene, Florida), and G. collazm (Hubbard) 

(1920, p. 143, pi. 23, figs. 1, 2; late Oligocene, Puerto 

Rico) have more subdued sculpture than C. herculeanus, and the Puerto Rican species differs also in 

whorl profile. Perhaps the Puerto Rican species is that 

for which Maury (1920, p. 55, pi. 8, figs. 1, 2) proposed 

the name Campanile (Portoricia) lamcum, the monotype of Portoricia, based on molds. Until the molds 

can be correlated with specimens that show the sculpture, her name is a nomen dubium.

C. herculeanus and the Emperador species, whatever 

it may be, are relics of Tethyan origin, the last American species, younger than any in Europe.

Occurrence: Emperador limestone member of La 

Boca formation (early Miocene), locality Il7c.
432 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Family CALYPTRAEIDAE

Genus Cmcibulum Schumacher

Submenus Dispotaea Say (p. 83)

Say, Acad. Nat. Sci. Phila. Jour., 1st ser., v. 4, p. 131, 1824.

Type (logotype, Harris, Bull. Am. Paleontology, v. 1, p. 64 

[explanation of pi. 7 of reprint of Say's account], 1896): 

Calyptrcea grandis Say, Miocene, Maryland [Virginia].

I am indebted to my colleague Druid Wilson for 

pointing out this type designation in an unlikely part 

of Harris' reprint. It puts Dispotcea on a better footing 

than the later designation cited on page 83 of the 

present account.

Family STROMBIDAE

Genus Orthaulax Gabb (p. 190)

Orthaulax gabbi Ball

Plate 49, figures 7, 12

Orthaulax gabbi Ball, Wagner Free Inst. Sci. Trans., v. 3, p. 

170, pi. 12, figs. 5, 5a, 5b, 1890 (Miocene, Florida). Cooke, 

U.S. Geol. Survey Prof. Paper 129-B, p. 29, pi. 3, figs. 5, 

6a, 6b (Ball's illustrations), 7, pi. 4, fig. 1, 1921 (Miocene, 

Florida; Canal Zone records doubtful). Gardner, U.S. 

Geol. Survey Prof. Paper 142, p. 560, pi. 55, figs. 1-3 

(Ball's illustrations), 1947 (Miocene, Florida). H. E. and 

E. H. Yokes, Tulane Studies Geology, v. 6, no. 2, p. 74, 

pis. 1-3, 1968 (Miocene, Florida).

Of medium size, strongly shouldered, shoulder 

ridged. Spire moderately high. Horizontal section at 

shoulder triangular.

Height (incomplete) 69 mm (estimated restored 

height 80 mm), maximum shoulder diameter 51 mm, 

minimum shoulder diameter 39.5 mm (figured specimen).

Type: Lectotype, herewith designated, USNM 

112218.

Type locality: USGS 2212, 1 mile west of Baileys 

Ferry, Chipola River, Calhoun County, Florida, Chipola formation.

The collection from the La Boca formation at locality lOlh, on the west side of Las Cascades Reach, includes a specimen identified as Orthaulax gabbi. It is 

almost entirely a natural cast, consisting of calcareous 

sandstone and irregularly disposed dense calcite, 

flecked with black mineral grains, like those in the 

calcareous sandstone. An axial section shows ghosts of 

the lower columellar part of a few whorls, but no other 

shell or callus outline.

A large sample of 0. gabbi, recently described by H. 

E. and E. H. Vokes, was found to show a considerable 

range of variation in shell outline. Nevertheless fully 

mature shells and many of intermediate size, including 

the lectotype, have a ridged shoulder and a triangular 

horizontal section at the shoulder. Those features are

well shown by a series of large specimens from Alum 

Bluff (USGS 2211), ranging in height, or estimated 

restored height, from 100 to 120 mm. The ridged shoulder is the most diagnostic feature of this species.

According to the catalog record, five specimens were 

in the type lot. The larger specimen illustrated by 

Dall is designated the lectotype. The smaller illustrated one is missing.

The fossils recorded as 0. cf. 0. pugncwc and 0. cf. 

0. aquadillensis on page 191 should have been recorded 

as Orthaulax sp., as their affinities are uncertain. They 

are so recorded in lists in the present chapter.

Occurrence: La Boca formation (early Miocene), 

locality lOlh. Chipola formation (early Miocene), 

Florida.

Family COLUBRARIIDAE 

Genus Colubraria Schumacher

Schumacher, Essai d'un nouveau systeme des habitations des

vers testaces, p. 251, 1817. 

Type (monotype): Colubraria granulata Schumacher (= Murex

maculosus Gmelin r= Buccinum muricatum Lightfoot),

living, western Pacific Ocean.

Colubraria obscura (Reeve) 

Plate 63, figures 20, 21

Triton obscurus Reeve, Conchologia iconica, vol. 2, Triton, species 63, pi. 16, fig. 63 and text, 1844 (living, East Indies).

Colubraria obscura (Reeve), Warmke and Abbott, Caribbean 

seashells, p. 116, pi. 21, fig. q, Livingston Publishing Co., 

1961 (living, West Indies, western Pacific Ocean).

Epidromws testaceus Morch, Catalogus conchyliorum * * * 

Comes de Yoldi, p. 107, Copenhagen, 1852 (living, West 

Indies).

Triton testaceus (Mb'rch), Morch, Malakozooligsche Blatter, v. 

24, p. 25, 1877 (assigned to subgenus Colubraria; living, 

West Indies).

Colubraria lucasensis Strong and Hertlein, Calif. Acad. Sci. 

Proc., 4th ser., v. 22, no. 6, p. 173, pi. 35, fig. 17, 1937 

(living, Baja Calif.). Emerson, Veliger, v. 8, p. 176, pi. 26, 

figs. 1-4, 1966 (living, Baja Calif, to Panama).

Colubraria soverbii (Reeve), Campbell, Nautilus, v. 74, p. 140, 

pi. 10, figs. 5, 6, text fig. 4, 1961 (living, Gulf of California 

to Guaymas, Mexico).

Large, inflated, whorls bulging. One varix on earliest 

preserved whorl, two irregularly arranged varices on 

other spire whorls, only terminal varix on body whorl. 

Early sculpture finely and evenly reticulate, axial and 

spiral threads of about equal strength, small nodes at 

intersections. At later stage axial threads stronger than 

spiral. On last few whorls both sets and accompanying 

nodes subdued. Interior of outer lip lirate. Base of 

parietal shield obscurely ridged. Narrow ridge near 

upper end of parietal wall. Siphonal canal short, bent 

backward, deeply emarginate.

Height (almost complete) 40.5 mm, diameter 15.5 

mm (figured specimen).
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 433

Type: Presumably at British Museum (Natural History).

Type locality: Living, East Indies.

One specimen identified as Colubraria obscura was 

collected from sandstone of the Gatun formation, about 

half a meter below the base of corallif erous conglomerate, 011 the south side of Rio Chargres, 175 meters below 

the spillway of Gatun Dam. The body whorl and part 

of the penult whorl are somewhat worn.

This is the first Tertiary record of a large species of 

Colubraria from the Caribbean region or southeastern 

United States, aside from the early Miocene Brazilian 

0. parcensis Maury (1925a, p. 127, pi. 6, fig. 1), which, 

according to Maury's illustration, has strongly bulging 

whorls and strong spiral sculpture.

0. obscura is recognized as living in the Caribbean 

region and in the western Pacific Ocean at least shells 

that are indistinguishable are found in both areas. On 

the same basis it also survived in the eastern Pacific 

Ocean, where it is represented by C. lucasensis. Though 

the type of G. lucasensis is small (height 27 mm), the 

shell from the Pacific coast of the Canal Zone illustrated by Emerson (figure 4) closely agrees with the 

Gatun fossil in size, outline, and sculpture.

Occurrence: Middle part of Gatun formation (middle Miocene), locality 160c. Living, Florida, Bermuda, 

Bahamas to Barbados; Gulf of California to Panama; 

western Pacific Ocean.

Family MTTRICIDAE (p. 214)

Subfamily MTTBICI1TAE 

Genus Fterynotus Swainson

Swainson, Zoological illustrations, 2d ser., v. 3, p. 100, 1833,

subgenus without species. 

Type (monotype, Swainson, Idem, p. 122, pi. 122, 1833): Murex

(Pteronotus) pinnatus Swainson (=Purpura alata

Rb'ding), living, western Pacific Ocean.

Subgenus Subpterynotus Olsson and Harbison

Olsson and Harbison, Acad. Nat. Sci. Phila. Mon. 8, p. 246, 

1953.

Type (orthotype): Murex textilis Gabb [Murex (Pteronotus) 

textilis], Miocene, Dominican Republic, Venezuela, Panama, Florida; Pliocene, Florida.

Subpterynotus is distinguished from Pterynotus s.s. 

by its long, almost straight siphonal canal and its correspondingly almost straight, short end of the siphonal canal preceding the terminal canal. The cloth-like 

sculpture is stronger than that of the type species of 

Pterynotus, but is similar to the sculpture of other 

species, such as the western Pacific P. pellucidus 

(Keeve) and P. tripterus (Born) (Keeve, 1845, pi. 14, 

figs. 54, 55, respectively), and the eastern Pacific Murex 

rhyssus Dall, a junior homonym renamed Pteropurpura vokesce Emerson (1964a, p. 5; illustrated as 

Pteropurpura rhyssa, Emerson, 1964, p. 153, pi. 20, fig. 

1). The proper subfamily and generic assignment for 

Pteropurpura vokesce will not be known until the 

radula is known. Its operculum has an apical nucleus 

like that of most muricine species, but its siphonal 

canal is closed, like that of the type species of Pteropurpura, an ocenebrine genus, although not all species 

of that genus have a closed canal.

Pterynotus (Subpterynotus) textilis (Gabb) 

Plate 63, figures 18, 19

Murex (Pteronotus) textilis Gabb, Am. Philos. Soc. Trans., n.

ser., v. 15, p. 202, 1873 (Miocene, Dominican Republic).

Dall, Wagner Free Inst. Sci. Trans., v. 3, p. 142, pi. 9,

fig. 4, 1890 (Pliocene, Florida). 

Murex textilis Gabb, Guppy, Geol. Soc. London Quart. Jour.

v. 32, p. 522, pi. 29, fig. 1 (an inadequate illustration),

1876 (Miocene, Dominican Republic). 

Murex (Pteropurpura) textilis Gabb, Dall, Wagner Free Inst.

Sci. Trans., v. 3, p. 243 (in discussion), 1892. Pilsbry,

Acad. Nat. Sci. Phila. Proc., v. 73, p. 353, pi. 28, fig. 4,

1922 (Miocene, Dominican Republic). 

Murex (Subpterynotus) textilis Gabb, Olsson and Harbison,

Acad. Nat. Sci. Phila. Mon. 8, p. 247, pi. 36, figs. 7, 7a,

1953 (Pliocene, Florida). Jung, Bull. Am. Paleontology,

v. 49, no. 223, p. 521, pi. 69, figs. 8, 10, 1965 (Miocene,

Venezuela). 

Murexiella (Subpterynotus) textilis (Gabb), B. H. Vokes, Tulane Studies Geology, v. 6, no. 3, p. 120, pi. 8, figs. 1-3,

1968 (Miocene, Dominican Republic, Florida; Pliocene,

Florida).

Of medium size, trivaricate, a low, wide swelling 

midway between varices. Whorls strongly bulging, 

concave near suture. Varices thin, alined on last three 

whorls, edge bent backward, short projections on edge 

alined with primary spiral threads, still shorter projections alined with some secondary threads. Primary 

spiral threads high-standing, secondary, or secondary 

and tertiary, threads lower. Fine axial threads producing an effect of loosely woven cloth. On apertural 

face of varices axial threads looped upward (away 

from aperture) or horizontal in narrow channels corresponding to spiral threads, and looped downward 

(toward aperture) in channels or on flat surfaces corresponding to spaces between spiral threads, upward 

looping more conspicuous than downward. On back 

face of varices and between varices axial threads 

crimped, on some whorls subdued between varices. 

Lower part of body whorl, including siphonal canal, 

missing.

Height (incomplete) 24.5 mm (estimated restored 

height 40 mm), diameter 19.5 mm (figured specimen).

Type: Acad. Nat. Sci. Phila. 3257.

Type locality: Dominican Kepublic, Miocene.
434 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Though the siphonal canal of the only specimen, 

found in the lower part of the Gatun formation, is 

missing, the sculpture is unmistakable. The occurrence 

of this species in the Chipola formation, recorded by 

Vokes, extends its time range; that in Panama expands the middle Miocene distribution; and late Miocene specimens from Florida, also recorded by Vokes, 

fill in a time gap. Eight shells of that age are in Druid 

Wilson's Acline collection of that age. On the Florida 

late Miocene and Pilocene shells the bands of downward loops on the apertural face of the varices are 

wider and therefore more conspicuous than the bands 

of upward loops. The sutural whorl-concavity is deeper 

on Pliocene specimens than on those of middle and 

late Miocene age. A progressive increase in size is 

shown by the available material: early Miocene, 15 

mm; middle Miocene, 33 to an estimated 40 mm; late 

Miocene, estimated 48 mm; Pliocene, 56 to 67 mm.

This remarkable species left no descendants. It may 

be related to the Miocene Italian Pterynotus latilabris 

(Bellardi and Michelotti) (Bellardi, 1873, p. 88, pi. 4,

%. 11).

Occurrence: Lower part of Gatun formation (middle 

Miocene), locality 138c. Chipola formation (early Miocene) , Florida. Cantaure formation (middle Miocene), 

Venezuela. Miocene (presumably Cercado or Gurabo 

formation), Dominican Republic. Deposits of late Miocene age, Florida. Caloosahatch.ee formation (Pliocene) , Florida.

Subfamily TYPHINAE

Genus Typhis Montfort

Subgenus Talityphis Jousseaume (p. 221)

Typhis (Talityphis) eucteanus Woodring, n. sp.

Plate 63, figures 9, 10

Small, slender, spire high. Early whorls worn. Remaining whorls bearing four varices, ending on shoulder in blunt, hollow spines. Terminal varix lamellar, 

others fairly wide. Tubes closer to preceding varix than 

to succeeding varix, generally broken almost to base. 

Spiral sculpture faint or absent on spire whorls. Body 

whorl bearing six weak, widely spaced spiral threads, 

most conspicuous on flanks of terminal varix. Terminal 

varix expanded, its edge frilled by the spiral threads. 

Siphonal canal short, tip missing.

Height (practically complete) 19 mm, diameter 9.7 

mm (type). Height (practically complete) 23.3 mm, 

diameter (incomplete) 11 mm (largest specimen).

TyperUSNM 646083.

Type locality: 138f (USGS 23663, south side of 

Transmisthmian Highway, hillside excavation at Colchoneria Yero, about 450 meters southwest of Cativa, 

Panama, lower part of Gatun formation.

This small, slender species was found in the lower 

part of the Gatun formation. Three were collected at 

the type locality and three others, available through 

the kindness of H. E. and E. H. Vokes, are in the 

Tulane collection from a nearby locality.

Typhis eucteanus is smaller and more slender than 

T. alatus Sowerby (1850, p. 48, pi. 10, fig. 4; height 

31.8 mm, diameter 9 mm), collected only by Heneken 

in the Dominican Republic, and the varices, other than 

the terminal varix, of the Gatun species, are less lamellar. It is of the same size as T. pterinus Gardner (1926- 

47, p. 528, pi. 53, fig. 14, 1947; Shoal River formation, 

Fla.), but is more slender and the varices of that species are like those of T. alatus. On the contrary the 

varices of T. puertoricensis Warmke (1964), a small 

species living in Puerto Rican waters, are high and 

wide.

Occurrence: Lower part of Gatun formation (middle 

Miocene), locality 1381

Family NASSARIIDAE 

Genus Psilarius Woodring

Woodring, Nautilus, v. 77, no. 4, p. 143, Apr. 14, 1964. Substitute 

name for Leptarius Woodring, p. 272 of present account, 

Feb. 27, 1964, not Leptarius Gill, 1863.

Attention is drawn to the substitute name for the 

junior homonym Leptarius.

Family OLIVIDAE 

Subfamily OLIVIFAE 

Genus Oliva Bruguiere 

Subgenus Strephonella Ball (p. 278) 

Oliva (Strephonella) colpotus Woodring, n. name 

Oliva plicata Guppy, in Guppy and Dall, U.S. Natl. Mus. Proc., 

v. 19, p. 308, pi. 30, fig. 12, 1896 (Miocene, Jamaica). Not 

Oliva plicata Fischer von Waldheim, Museum-Demidoff, 

v. 3, p. 161, Moscow, 1807.

Oliva dimidiata Pilsbry and Johnson, Acad. Nat. Sci. Phila. 

Proc., v. 69, p. 165, 1917 (Miocene, Dominican Republic). 

Not Oliva dimidiata Sowerby, in Darwin, The geology of 

the voyage of the Beagle, pt. 3, Geological observations 

on South America, p. 263, London, 1846.

Both Oliva plicata Guppy, the name used on page 

278 of the present account, and its subjective synonym 

Oliva dimidiata Pilsbry and Johnson are junior homonyms.

Family?

Genus Glyptostyla Dall

Glyptostyla panamensis Dall (p. 289)

Plate 48, figure 23

After publication of chapter C, A. A. Olsson exposed by careful work the columella of two specimens 

of Glyptostyla panamensis. They show that the folds
GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 435

are not like volutid folds, they are seated on a pad. The 

wide basal fold is followed upward by a gap and then 

by four or five closely spaced narrower, low folds. The 

effect of a wide basal fold and a wider upper fold on 

the type, as shown on plate 39, figure 21, is due to 

damaging of the pad when the specimen was prepared 

for the drawing published by Dall. The result is that 

Olsson's Peruficus, instead of being a synonym of 

Glyptostyla, as concluded on page 289, is a valid genus 

and doubtless is to be allocated to a different family.

Olsson, who had examined and photographed the 

lectotype of Strepsidura striata Newton (1922, p. 33, 

pi. 3, figs, 24, 25 (S. multistriata in explanation of 

plate); Eames, 1957, p. 48, pi. 7, fig. 3), an Eocene species from Nigeria, pointed out that it is very closely 

related to 'Glyptostyla panamensis. The remarkable 

columellar features are practically identifical, except 

that on Strepsidura spirata the gap above the basal 

fold is partly filled by a low fold. The Nigerian species 

has a conspicuous siphonal fascicle limited by a thread 

and its axial sculpture is weak. The end of the siphonal 

canal is broken on all the specimens of Glyptostyla 

panamensis. Much of the columellar side remains on 

the type, but the siphonal fascicle is worn. The fascicle 

is shown to best advantage on an immature specimen 

(locality 40d, height 24.6 mm). The limiting thread is 

weaker than that of the Nigerian species. It should be 

pointed out that the outer lip of both species is flaring. 

Strepsidura spirata is the type of Strepsiduropsis 

(Eames, 1957, p. 48), proposed as a subgenus of Strepsidura. The Nigerian fossil is considered to be a species 

of Glyptostyla. Whether the differences between it and 

G. panamensis are of specific or subgeneric rank is a 

matter of judgment. The family status of the genus is 

uncertain.

Occurrence: Marine member of Bohio (?) formation 

(late Eocene or early Oligocene), localities 40, 40a, 40c, 

40d.

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GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE 439

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368-278 O - 70 - 10
440 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

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INDEX

[Italic page numbers indicate both major references and descriptions]

Abderospira... ............................... 420

chipolana. ................................ 419

abrupta, Turritdla.... .......... 318, 323, 430, pi. 63

abundans, Pleurotoma. ....................... 378

acalypta, Cancdlaria... ............... 318, 841, pi. 53

acanthodes,MitreUa. .......................... 306

acaria ectypha, Agladrillia (Agladrillia) ........ 320,

324, 387, pis. 59, 64 

acicularis,Eulima. ......................... 326,327

acidna, Agladrillia (Eumetadrillia) . . . . 320, 388, pi. 60

adinica, Cymatosyrinx . ....................... 374

Cymatosyrinxlunata.. .................... 374

Acmaeal sp.. ................................. 314

acoZws, Conus. ....................... 318, 34#, pi. 55

Cymatophos"! . ............................ 325

ocra, Pleurofusia. ........... 318, 367-368, pis. 57, 65

/Icropora __--.._._....__.__ ____. .------.. 312

Actaecm cubensis... ........................... 415

punctostriatus . ........................... 415

Actaeonidea. .................................. 416

oryza.. ................................... 416

anetaspira ................................ 418

buttata. ............................ 306,324,417

bullata, small form ......... 320, 417-418, pi. 62

cf. A. &M«ata .............. 305, 306, 417, pi. 48

canaliculata ... ............................ 417

candei... ............................... 417,418

elachista..... ..................... 320, 4/8, pl.62

harpa.. ................................ 418,419

rosa. ..................... 320, 324, 4/S-4/9, pl.62

subbuUata. ............................. 417,418

cf. subbuUata..... ... ..................... 417

subbuttata costaricana. .................... 417

sp. ..................................... 316,417

. _ ............. _ .-.-...-...-.-. 415

exilis. .................................... 416

punctatus. ............................... 415

riomaensis... ............................. 415

tampae. ................................ 306,415

wetheritti.. ............................... 417

(Acteori) punctostriatus..... 320, 324, 4/5, pi. 62

afl. A. tampae.................. 305,306,4/5

(Lissacteon?) sp. ................... 320, 415-416

Acteonidae... ........ _ ..................... 415

acuaria, Terebra. ........................... 411,412

acuta, BuUa. ................................. 426

Eulima.. .................. 318,323, 327, pi. 51

Leiostraca ..... ............................ 327

Pleurotoma.. ................. ... ......... 364

Strombiformis. ............................ 327

acutirostra, Pleurotoma........................ 360

adamsf,l?eiMsa..--.. __ ..... _ ........... 305,423

( Cylichnina) afl . 1? ............. 304, 305, 4#3

Additions and corrections to families covered 

in preceding chapters.... ................... 430

Additions to annotated bibliography ......... SOI

Additions to localities at which fossils were 

collected.. ............................... 301-302

adela, Turritella.. ............................. 305

addaidis, Rhizorus. ........................... 424

adematum, Lioglyphostoma .................... 3 99

Adrana newcombi. ............................ 326

perprotracta . ............................. 326

aegis, Crassispira. .............. _ ----------- 377

aemulator, Conus. ......... ... ................ 351

Conus proteus .. ... ....................... 351

aemulator Continued Page 

aemulator, Conus... 318, 324, 351-352, pis. 55, 56 

manzanittaensis, Conus................... 351

Aeguipecten.................................. 312

Aforia........................................ 361

Agaronia testacea mancinetta................... 323

sp 314

agassizimultiliratus, Conus.................... 356

Agassizia clevei................................ 312

Agladrillia.................................... 385

collothyra................................. 385

estrdlana................................. 386

Idandi.................................... 384

mimys. ------------------.-.-.--.--------- 385

(Agladrillia)..... ......................... 322

acaria ectypha........ 320, 324, 3S7, pis. 59, 64

characta......... 320, 385, 386, 387, pis. 59, 64

enneacyma enneacyma................. 320,

385-386, pis. 59, 64

subsp-. ..- . 320,3S6 

phengoides.......... 320, 386-387, pis. 59, 64

(Eumetadrillia) acidna.......... 320, 388, pi. 60

isthmica.............. 320,387-388, pis. 60, 64

agriaagria, Cancellaria........................ 342

uaquala, Cancellaria...................... 342

aguadillana, Mangilia......................... 397

aguadittensis, Orthaulax ct. 0................. 432

Alabina asperaides asperoides.................. 323

canaliculata........................... 323

alaquaensis, Conus............................ 358

oZaiws, TypMs................................ 434

o&esws, TtypMs ( Talityphis) ............... 323

albida, Pleuroliria... .......................... 365

Pleurotoma^........................... 363,364

teKea, Pleurotoma......................... 364

Tunis.................................. 363,364

sanctidavidis, Tunis...................... 369

aZMdoides, Pleuroliria.-....................... 363

alcestis, Clathrodrillia......................... 375

aldricM,Dolabella._......................... 420,421

Floribella..................... 313,316,4s/,pi. 50

Semicassis................................ 311

Semicassisf (Tylocassis?) cf. S........... 310,311

alesidota, Crassi-pira........................ 379,380

macilenta, Pkurotoma sp. aff. P......... 378,379

magne, Pkurotoma (Drillia).-............. 379

alfaroi, Valuta................................ 325

eurytera, Voluta......................... 324,325

Alhajuela formation. ------------------------- 317

Aliculastrum................................. 4##

allodapum, Glyphostoma (Rhiglyphostoma)....- 320,

400, pis. 60,66 

attilira, Turritella ct. T....................... 306

altilira, Turritella (Bactrospira)..... 317,323,325

(Torcula)......................... 317

praecellens, Turritella (Bactrospira)....... 323

subsp., Turritella (Bactrospira)........... 306,

307,313,314,316 

(Torcula)....................... 307,316

altiuscula, Solariella.......... ............... 306

alveata,Architectonica......................... 305

(Stellaxis) aff. A...................... 305

amaras, Hemisinus............... 318,430-4S/,pi- 63

Turritella (Bactrospira?).... 310,311,313,314,316

(Torcula?)...................... 310,311,316

Amarophos................................... 325

ftottrws----------------------------------- 325

Amarophos Continued Page 

dentalis................................... 325

ame, Dirocerithium............................ 303

arnica, Oemmula............................ 307,360

Gemmulact. O...................... 306,307,360

amicta rintriada, Glyphostoma................. 395

Nannodiella.............................. 395

amphitrites, Bursa (Colubrellina) caelata-...-.. 323 

Ampullinopsis................................ 307

spenceri.................................. 307

amydra, Persicula (Rabicea) venezuelana....... 310,

311,314,316,333, pi. 48 

amygdala, Bulla.............................. 422

Marginella................................ 331

Anachis (Costoanachis) mirafugax............. 323

mira.............................. 323

anapetes, Latirus (Polygona).................. 325

Ancilla (Eburna) pinguis..................... 323

Ancistrosyrinx................................ S71

cedonullireevi............................. 371

datti...................................... 371

elegans vax................................ 371

miranda.................................. 371

andersoni, Bathygalea......................... 325

Pleuroliria................................ 365

anefrupt'ra, Acteocina.......................... 418

angulifera, Littorina........................... 311

Littorina afl. i....................... 310,311

annella, Crassispira.- ------------------------- 381

anomoia, Cancellaria (Cancellaria)............. 318,

334-335, pi. 52 

anorhepes, Dolostoma..... 320,324,397,398,pis. 60,66

antealesidota, Crassispira...................... 379

anthera, Retusa............................... 424

antittea, Heterostegina......................... 309

Miogypsina............................... 309

Antillophos candei..... ------------------------ 356

(AnftZZopftos) candei gatunensis.. 311,316,323,336 

Antillophos"! (Antillophosl) cf. .4. caredej

gatunensis.............. 310,311,314,316

Antillophos (Antillophos) mexicanus........... 323

Antillophos^ (Antillophosl) sp. (small) -------- 314

antiquum, Morum............................. 305

(Cancellomorum) ct.M................ 305

("Oniscidia") cf. M_.----------------- 305

apalachee, Prunum....----------------------- 316

(Microspira) aff. P........ 314,316,330,pi. 48

apenes, Semicassis (Echinophoria)............. 307

aperta, Calyptraea............................ 303

Calyptraea cf. C...---------------------- 303

Aphera................................. 322,334,344

islacolonis.................... 318,323,344, pi. 56

..................-- . 344

.................. ...---- 344

apimela, Cancellaria (Cancellaria)-ZIS,SS7-SS8, pi. 52 

Apiocypraea. _---.---.-- .------------------- 322

apium, Conus. --------- ...----------------- 347

Architectonica.. ------------------------------- 306

aZceafa.----------------------------------- 305

fungia.........-.---- . 305

cf. /I./Magia...--------------------------- 305

(Architectonica) nobilis.............. 307,313,322

ct. A.nobilis..- ---~ 316 

mobito fcarsiewi..---------------------- 325

nobilts.. -------------------------- 323

subsp........-.---------------- 313,314

rhicna................................ 307

(Pseudotoriniat) sp...--.----------------- 314

441
442 INDEX

Arckitectonica Continued Page 

(Stellaxis) afif. A. alveata.. ................ 305

arcuata, Cruziturricula.. ...................... 369

Pleurofusia (Cruziturricula).............-- 368

Pleurotoma ............................... 369

"Tunicula".. ............................ 369

arcuatus, Conus...... ................... 346,355,356

armillatum , Strioterebrum ..................... 414

arteaga, Mangilia. ............................ 393

Asperdaphne versivestita ....................... 403

asperoides asperoides, Alabina ................. 323

canaliculata, Alabina.. .................... 323

asperum, Strioterebrum........................ 409

asterodisca, Lepidocyclina.. .................... 308

astuta, Melanella. ............................. 328

atacata, Cylichnella ........................... 419

stibara, Cylichnella........... 320, 324, 419, pi. 62

Atlanta (Atlantidea) lissa. .................... 323

Atlantic and Pacific muck.. . ..._..._.._.__. 325-386

atlanticus, Conus spurius... ................... 348

atraktoides, Cancellaria. ...................... 342

atriformis, Hemisinus. ........................ 306

Atyidae. --......_.----._.......---_...-___-.- 422

4tys-.----. ----------------------------------- 422

caribaea. ................................. 423

daZZi - _... _- 422 

doliolum.. ................................ 423

gracilis .... . ............................... 422

oedemata ... ............................... 423

sulculorum. ... _- .. _-_____- -_.- 423

sp. ----.---- ---- -- 422

.4^*7 sp-. . .- __ -. 316,422 

Atys (Aliculastrum) eurys..... 320, 488-488, pi. 62

sp.......--..-.......... ...--- 306

(Roxaniella) rhadina. ...... 305, 306, 422, pi. 48

(Weinkauffia) cadus. ............ 320, 483, pi. 62

audeninoi, Cavolina. .......................... 428

trinitatis, Cavolina........................ 428

aulaca, Balds (Balds) ................ 318,328, pi. 51

aulakoessa, Clathrodrillia.. .................... 383

auricincta, Eulima. ........................... 327

auriculaptera, Cancellaria.. ........ ........... 338

aurora, Calliostoma.-.. ....................... 325

austini, Conus................................ 346

<zfc-e«a , Marginella ................ ... .......... 330

Volvarina. . ............................... 331

Averellia. ..................................... 429

coactiliata.. ............................... 305

cordovana ................................. 429

(Lecallia) stewarti..... 304, 305, 429-430, pi. 48

B 

Balantium undulatum. ....................... 427

balboae, Cancellaria ... ........................ 340

bartschi . 

calkinsi.

egregia ... .................................

jamaicensis ...............................

makista...... .............................

maoica.. ..................................

(Balds) aulaca................... 318,328,

ceHa ... _ 318,328-329, 

jacululum.. .......... 318,323,327-328,

lipara..... ................... 318,328,

barbadoides, "Cythara".. ......................

barretti, Pleuroliria. ..........................

bartschi, Balds ...... ..........................

Daphnella. .............. ... ..............

bartschiana, TurboniVa. .... ...................

barystoma, Cancellaria (Narona) ...... 318,34?,

basilissa, "Cythara"...........................

Bathygalea andersoni.. ........................

datti... ...................................

(Miogalea) hadra.. ......... ... ............

Bathytoma..... ... ......................... ...

belemnitum, Terebellum. ......................

belemnttumf, Terebellum (Seraphs) ............

328

328

329

329

328

329

328

pi. 51

pi. 49

pi. 51

pi. 51

393

365

328

402

414

pi. 53

391

325

325

325

363

303

303

375

Page 

berjadinensis cocoditana, Turritella............. 313

TurritellacL T..................... 313,314

bermudensis, Conus........................... 349

bidentata, Cylichna? afif. Cylichnella....-.---.-. 419

Cylichnella................................ 419

biforis, Retusa................................. 423

( Cylichnina) querdnensis............. 320,

324,4^3-424, pi. 62 

bipartita. Terebra............................. 404

bipartita, Terebra (Acus)................. 411

spirifera, Terebra (Acus)................ 409,410

bisulcata, Cavolina............................ 428

bitropis, Hemipleurotcma...................... 362

Bittium permutabile........................... 313

scotti....... ___...... 313,314

BivetieJla............................... 337,340.341

Bivetopsia.................................. 340, 341

blackwaterensis, Campanile hernandoensis..... 431

blountensis, Crassispira....................... 379

blountiana, Cancellaria......-....-....---.-.-. 342

bocasensis, Agladrillia......................... 388

Bohio formation... . 305-306

bolivarensis bolivarensis, Neverita............... 305

tapina, Neverita (Glossaulax).............. 305

fioZMs, AWca (Natica1?)........................ 323

Borsonia (Paraborsonia) centaurana........... 374

Borsoniinae.-.-.-. ............. .. . 360,372

bothrus, Amarophos........................... 325

boussaci, Conus marginatus.................... 358

Brachycythara galae........................... 393

turrita.................................... 391

brassoensis, Paraborsonia.................... 307,374

Paraborsonia afif. P............. 306,307, 378-874

Tunis.................................... 362

firazw, Conus............. 318,324,348-349,350, pi. 56

Conus molis.............................. 348

brunneus subclausus, Polinices................ 323

Buccinum taurinum........................... 406

Buchema..................................... 375

-BwZZa......................................... 42;

aCMta .. ........... 426

amygdala................................. 422

paupercula............................. 421,422

sarahberlinerae............................ 422

soZMa..................................... 422

striata.................................... 422

umbilicata, small form.___ _ . 320,

324,42^-422, pi. 62

waltonensis........................... 422

-BztZZa?sp. ._ _ _. ._ 306,421

-BwZZa (VoZTOZa) cf. uxytata..................... 425

Bullaria paupercula........................... 421

bullata, Acteodna....................... 306,324,417

small form, 4cteocma.--..... 320,4/7-4/8, pi. 62

4cteociraa cf. 4............... 305,306,4/7, pi. 48

Cancellaria (Trigonostoma) afif. Cancellaria. 345 

ToraafeZZa................................ 417

Tornatina................................ 417

bullbrooki, Cancellaria......................... 342

Crassispira consors........................ 379

Drillia consors............................ 378

Bullidae . _ _ . _ 421 

Bullina (Abderospira) chipolana............... 419

(Tornatina) afif. candei.................... 418

burckhardti, Conus............................ 357

burckhardti?, Conus........................... 357

burckhardti burckhardti, Conus................. 318,

324,357-358, pi. 57 

harrisi, Conus............ 318,324,358-359, pi. 57

Buridrillia.................................... 377

Buridrillia?................................... 377

Bursa (Colubrellina) caelata amphitrites__-_. 323

C 

cacuminatus, Conus........................... 351

cadus, Atys (Weinkauffia)............. 320,423, pi. 62

caelata amphit rites, Bursa (Colubrellina)....... 323

Caimito formation......................... 306-307

Page 

caleta, Turritella.............................. 305

Turritella o,i. T........................... 305

calkinsi, Balds................................ 328

calligonoides, Crassispira...................... 379

Calliostoma aurora............................ 325

(Calliostoma) metalium.................... 325

callothyra, Agladrillia......................... 385

Calophos...................................... 322

calypta, Tricolia.............................. 305

Calyptraea aperta............................. 303

cf. C. aperta.............................. 303

centralis............................ 311,313,323

cf. C. centralis.................. 310,311,313,314

Caly ptraeidae...._______ ____- _____ _. 432

camaronensis, Lepicythara..................... 391

Campanile.................................... 431

charpentieri............................... 431

collazus................................... 431

halensis................................... 431

herculeanus............................ .316,431

cf. C. herceuleanus............ 312,316,4#/,pi. 51

hernandoensis............................. 431

blackwaterensis........................ 431

vaughani.................................. 431

Campanile? SV-.--------.................... 314,431

Campanile (Portoricia) laricum................ 431

Campanilinae................................ 431

camura, Malea................................ 323

canaliculata, Acteodna........................ 417

Alabina asperoides........................ 323

Cancellaria................................... 884

acalypta.......................... 318,S4/,pl. 53

affn'a affn'a................................ 342

uaguala............................... 342

atraktoides................................ 342

auriculaperta............................. 338

balboae.................................... 340

blountiana................................ 342

bullbrooki................................. 342

cassidiformis.............................. 340

chrysostoma............................... 340

coerwis ......... ..................... 342

corrugata................................. 343

Cancellaria? cf. C.dariena..................... 336

Cancellaria dariena trachyostraca............... 336

var....................... . ._ 336

darienensis................................ 336

defuniak.................................. 335

 . . - 344 

omifera............................. 336,337

epistomifera......................... 336,337

harpiformis............................... 340

hettneri................................. 339,340

karsteni................................. 339,340

laevescens................................. 335

lagua..................................... 341

lavelana................................... 341

macneili................................... 335

montserratensis....... .... .. ..... 340

nancellaria................ 318, 841-848, pi. 53

perdidana................................ 343

propevenusta.............................. 343

pj/cto... ...... .. ... 338

reticulata.................................. 335

r<we«i . .. ......... ... 335,338

runchaena............................... 343

scheibei................................... 339

similis.................................. 337,340

soZzda..-.........-......-.-------------.- 338

tessellata.................................. 344

venezuelana............................... 340

pewwsia... _ _-.- --- - --- 343 

werenfelsi----.-........................... 340

Cancellaria? sp__. ------ 314

Cancellaria (Aphera) islacolonis............... 344

(Cancellaria) anomoia...... 318, 884-885, pi. 52

apimela................ 318, 387-388, pi. 52

dariena............................. 335,336
INDEX 443

Cancellaria (Cancellaria) Continued Page 

epistomifera dariena. .................. 318,

323,325, SSS-SS7, pi. 52

afl. C. epistomifera dariena- 322, 325, 336, 337 

epistomifera lipara. ...... 318, 323, SS7, pi. 52

afl. C.macneili............. 318, SS5, pi. 52

tapeina.................. 318, SS5, 339, pi. 51

sp.................................. 322, SSS

(Charcolleria) terryi.... 318, 323, 848-344, pi. 54

sp.-_.-___.__.-_----__-_-------------- 343

CEwcZia). _ _ ____-__.___.-___ 322

codazzii............. 318, 323, 889-340, pi. 54

dinota.. .................. 318,5^0, pis. 54, 56

maldonadoL........................ 339,340

(Narona)................................. 322

barystoma................... 318, S4#, pi. 53

decaptyx............................ 318, S4#

(Pyrudia)........................ ....... 322

dbarcoladbarcola..... 318,323, SSS, pis. 52,53

diadela... ........... 318, 325, 388-389, pi. 53

diadelaf............................. 322,325

(Trigonostoma) afl. Cancellariabullata..... 345

Cancellariidae- . ------- - 334

candei,Acteodna...... ...................... 417,418

Antillophos............................... 356

gatunensis,Antillophos (Antillophos)....... 311,

316,323,336 

Antillophos? (Antillophosf) cf. A------ 310.

311,314,31 fi 

Bullina (Tornatina) ati................... 418

caradida, Clavatula............................. 401

caparorais, Turritella........................ 311,313

Carabaformation..---...-.__-_----.------.. 307-308

carbasea, Ficus................................ 317

carbasea, Ficus...................... 317,323,325

micronematica, Ficus............ 310,311,313,314

Ficus cf. P............................ 316

caribaea, Atys................................. 423

caribbeana, Vaginella.......................... 427

carinata, Turritella............................ 303

Turritella cf. T........................... 303

CariJiodriZHa S75

eZocafa. _ -- .-- 376 

felis............................... -. 376

haliostrephis............................... 376

winchesterae............................... 376

(Buchema) tainoa......................... 375

(CarinodriJlia) cf. C. elocata...... 318, S76, pi. 58

zoofci................. 318 875-376, pis. 58, 65

sp. - 325, 376-377, pi. 63

cascadewsis, Goniopora___....-...--.....--- 308

Goniopora cf. GL.... . 308

cassidiformis, Cancellaria...................... 340

Cassigerinella................................. 303

Cassis sp -..--..-----.-.---------------------- 314

Ctenatus, Conus.............................. 352

Conus cf. C............................. 318, SS2

Carolina...................................... 4%8

audeninoi................................. 428

trinitatis.............................. 428

bisulcata.................................. 428

coofcei.-.......... -..-- .--.---- - 428

ventricosa................................. 429

(Cavolina) triaspis............... 320, 488, pi. 63

cf. C. ventricosa.............. 320, 429, pi. 63

(Paleocavolina) xenica....... 306, 307, 4#8, pi. 49

Cavolinldae--..-.--.-- .------------ 427

cedonutti, Cochlespira (Andstrosyrinr)......... 318,

322,324, 871-372, pi. 57 

cedonulli, Pleurotoma.......................... 371

cedonulli reevi, Ancistrosyrinx.................. 371

cembra, Terebra (Oreoterebra) subsuldfera...... 320,

324,405, pi. 61 

(Paraterebra)......................... 405

centaurana, Borsonia (Paraborsonia)........... 374

centralis, Calyptraea..................... 311,313,323

CalyptraeacL C................. 310,311,313,314

cercadensis, Conus............................ 349

Nassarius (Uzita)........................ 323

cercadensis Continued

Persicula................................. 333

Volvula................................... 425

cercadica, Cythara............................. 390

Cytharact................................ 390

Melanella (Eulima)....................... 326

Cerithiidae ------------------------------- 431

Cerithium (Thericium) mimeticum............. 306

sp..-------------------------------- 314

cetta, Balds (Balds)............ 318, 328-389, pi. 49

Chagres sandstone, including Toro limestone

member-.... 322,325

Changes in formation and age assignments... 300 

characta, Agladrillia (Agladrillia).............. 320,

S85, 386, 387, pis 59, 64 

Charcolleria................................. 322, 848

chariessa, "Cythara".......................... 393

charpentieri, Campanile....................... 431

childreni, Pleurotoma......................... 360

chilona, Cypraea........................ 307,311,313

CypraeacL C........... 307,310,311,313,314,316

chipolana, Abderospira........................ 419

Bullina (Abderospira)................-.... 419

Micromelo (Abderospira)................... 419

Ro-ania................ 320, 324, 419-420, pi. 62

Vaginella............................... 307,427

chipolanum, Crucibulum (Crucibulum)........ 323

Morawi-- -- - ---------------- 313

(Cancellomorum) cf. M__.......... 313,314

("Oniscidia")ct.M................... 316

chipolanus, Conus.......... 306,307,311,316,354,355

Corawsafl. C -. . -- - 305,

306,307,310,311,314,316,354, 355

chipolanus imitator lineage Conus.......... 355,356

chiraensis, Conus............................. 349

Oostrombus............................... 303

Oostrombus afl. 0......................... 303

eft iriquiensis, Strombina (Sincola)............. 323

chloris, Leucosyrinr........................... 370

chrysostoma, Cancellaria...................... 340

dbarcola dbarcola, Cancellaria (Pyruclia)...... 318,

323, SSS, pis. 52, 53 

Cirsotrema sp ------------------- 314

cZarfci, Conus................................. 357

Perplicaria............................... 334

Clathrodrillia........................... 374,SS/,396

alcestis................................... 375

aulakoessa................................ 383

empera................................... 383

gatunensis.......... 318, 324, 881-388, pis. 58, 65

gibbosa................................. 382,383

islalindae................................. 384

cf. C.islalindae.. ............... 320, SS4, pi. 58

afl. C.lelandi.................. 320, SS4, pi. 59

mareana.................................. 383

onjoia...----_.--.-------------------- 382

puertocolombiana....................... 382,383

»aa»edrai...------ 318, SSS, 384, pis. 59, 65

afl. C. saavedrai............. 320, 888-884, pi. 58

subvaricosa............................... 383

serawsta.. --------- -- ------------------ 383

sp... ._..__ . . - 316,381

Clathrodrillia? sp.............. 325, S84-5S5, pi. 63

Clathurella panamella......................... 385

vendryesiana.............................. 396

(Glyphostoma) dentifera................... 400

Clavatula Candida............................. 401

Clavinae..... . ..-.--- ------ 360,374

Clavine turrid... - .-...--. -------- 306

cZ^ftra, Strioterebrum....................... 316,409

Strioterebrum cf. S_----------- 316, 409, pi. 50

cZewei, ^Iffassisia _ _ . 312

Clypeaster concavus...............-..-----.-.- 308

coactiliata, Averellia. _ . 305 

coactiliatus, Helix............................. 429

cochlearis, Pleurotoma..........-...-.---.---.. 363

Cochlespira..---..-..-....--.-.. --..-- S70-S71

elegans................................... 371

engonata................................ 370,371

radiate.. .. ....... - 371,372

Page 

Cochlespira? sp............................. 306, 371

Cochlespira (Andstrosyrinx) cedonulli......... 318,

322,324,S7/-57#, pi. 57 

Cochlespiropsis............................... 371

cocoditana, Turritella berjadinensis............. 313

Turritella cf. T. berjadinensis........... 313,314

codazsu, Cancellaria (Euclia).. 318, 323,SS9-S40, pl.54 

coensis, Cancellaria........................... 423

coleri, Strioterebrum ........................... 413

coUaris, Pleurotoma ........................... 367

collazica, Turritella.... ....................... 313

Turritella cf. T ____ _ 313,314,316,431

collazus, Campanile........................... 431

collina, Volvarina............................. 331

colombianum, Strioterebrum ------------------- 410

colpotus, Oliva (Strephonella)............... 323,434

Colubraria.................................... 452

Colubraria lucasensis....................... 432,433

Colubraria obscura...... 318, 322, 323, 4S#, 4SS, pi. 63

Colubraria paraensis.......................... 43,3

Colubraria soverbii............................ 432

C olubrariidae ________ ___ . _ _ __ -_ 432

comatotropis, Microdrillia...................... 390

compsacosta, Cytharella? cf. C_- 320, S92-39S, pi. 60 

Ithycythara............................. 392,393

concavitectum, Conus......................... 350

concavus, Clypeaster.......................... 308

Conidae . - ___ _ 345

coniforme, Prunum........................... 332

coniformis, Marginella......................... 332

Prunum.................................. 332

conradiana, Glyphostoma...................... 399

Volvulella............................... 305,425

(Volvulella) afl. V. _____ _ 304,305,4#5

consentanea, Alangilia......................... 394

consobrinus, Conus...................... 346,352,359

consobrinusf, Conus............---......-.-... 352

consobrinus consobrinus, Conus................ 318,

324, S5&-55S, pi. 56 

ultimus, Conus......................... 352,353

consors, CrassispiraaQ........................ 378

Crassispira? cf. Crassispira?............... 378

bullbrooki, Crassispira..................... 379

consors, Crassispira (Hindsiclava)......... 318,

324,325,3TS-5S0,382, pis. 58,65 

hobsoni, Crassispira....................... 379

magna, Crassispira........................ 379

pennae, Crassispira....................... 379

trinitatensis, Crassispira................... 379

Drillia ................................. 378

bullbrooki, DriUia......................... 378

portoricoensis, Drillia...................... 378

trinitatensis, Drillia....................... 378

Melongena melongena..................... 323

Pleurotoma............................... 378

Twrris (CrassispzYa). .,.. ............. 378

-- - -. -- 306,345-546

acoZus__________.......... 318, 349, pi. 55

aemulator................................. 351

aemulator......... 318,324, S51-S52, pis. 55,56

manzanittaensis....... ................ 351

agassizi multiliratus. 356

aZagaerasi*_......._...............-..- 358

apium......... . .... ... .- -.. -- 347

arcuatus............................ 346,355,356

austini. ................................ 346

bermudcnsis............................... 349

bravoi................ 318,324, 848-349, 350, pi. 56

burckhardti................................ 357

burckhardtil............................... 357

burckhardti burckhardti.... 318,324,357-558, pi. 57

harrisi................ 318,324,358-559,pi. 57

cacuminatus.............................. 351

catenatus................................. 352

cf. C. catenatus......................... 318,552

cercadensis................................ 349

chipolanus.............. 306,307,311,316,354,355

afl. C. chipolanus......................... 305,

306,307,310,311,314,316,354,355
444 INDEX

Conus Continued Page 

chipolanus-imtiator lineage.............. 355,356

chi raensis. ..._.......................... 349

clarki..................................... 357

concavitectum............................. 350

consobrinus......................... 346,352,359

consobrinus?.............................. 352

consobrinus consobrinus.... 318,324, 368-363, pl.56

uttimut............................. 352,353

corrugatus................................ 358

Aalll.................................... 354,355

........................... 354

......................... 350,351

emersoni................................ 346,353

fergusoni............................... 346,351

floridanus................................ 356

costaricensis.......................... 356

frisbeyae.................................. 357

gabbi..................................... 359

gaza...._.__.............................. 356

gradlissimus.............................. 359

granazonatoides. _......................... 352

granozonutus.............................. 359

harrisi.................................... 358

harveyensis............................... 358

haytensis................................. 351

imitator............................. 325,346,354

imitatorl............................ 322,325,356

aff. C.imitator.......................... 354,355

imitator imitator ...... 318,324, 364-366, 358, pi. 55

lius................................. 354,355

cf. imitator lius........................... 354

imitator subsp.. - -- - - 355

jaspideus................................. 347

larvatus_______............................ 359

lavilki.................................. 352,353

leoninus.................................. 348

cf. C. lius......-..--.-.................... 355

mahogani................................. 354

manzanillaensis........................... 352

marginatus............................. 348,356

cf. C.marginatus......................... 348

marginatus boussad....................... 358

mazei..................................... 359

mcgintyi.................................. 359

molis......... 318,324,346,348, 360-361, 352, pi. 55

aff. C.molis.............................. 351

molis bravoi............................... 348

multiliratus gaza........................ 356,357

muttiliratus...... 318,324, 366-367, 358,pi. 57

spiekeri............................... 357

wa.Zi................................. 357

muttittriatut.............................. 356

musaensis............... 318, 324, 347-348, pi. 57

patricius................................ 346,347

peruvianus............................. 307,359

cf. C. peruvianus....................906,307,369

planiceps............................... 316,350

cf. C. planiceps..................... 314,316,350

planiliratus............................. 346,356

proteus................................... 348

aemulator............................. 351

pygmaeus................................. 347

pyriformis.............................. 346,347

recognitus............... 318, 324, 346-347, pi. 55

riosantiagensis.......................... 348,349

sauridens............................... 304,349

cf. C. sauridens..................... 303,304,349

scaliae.................................... 353

________________________ 346,347

-_--__-_-.....-_.________. 355

.......................... 355

spurius............. 318, 324, 346, 348, 350, pi. 55

group._..--_..--_._._____________.__ 347

atlanticus............................. 348

.......................... 359

.......................... 346

sulculus..--__--_..-.____________.__ 306,307,350

cf. C. sulculus.............. 305,306,307,2

Conus Continued Page 

symmetricus............ 318, 324, 363-364, pi. 57

domingensis......................... 353,354

semiobsoletus........................ 353,354

taphrus......................... 318, 354, pi- 57

tornatus.................................. 358

toroensis.................................. 353

tortuosopunctatus............... 346,357,358,359

tortuosostriatus.. 318, 322, 324, 325, 352, 359, pi. 57 

turbinopsis............................... 357

veatchi.................................... 351

wittiamgabbl.............................. 347

sp___ .....__.________________ 306,348,351,352

(Lithoconus) sp........................... 353

cookei, Cavolina............................... 428

cordovana, Averellia........................... 429

coroni, Ctenilyria.............................. 304

corrugata, Cancellaria......................... 343

Philine................................... 421

corrugatus, Conus............................. 358

cossmanni, Purpura.......................... 389

costaricana, Acteocina subbullata............... 417

costaricensis, Conus floridanus................. 356

Cythara terminula........................ 390

costata, Cylichna.............................. 420

couviana couviana, Persicula................... 334

stenygra, Persicula (Eabicea).. 318, SSS-SS4, pi. 51 

Crassispira............._..._..._............. 377

ae0.8__ _-____ _________.___.___-_______- 377

alesidota................................ 379,380

annella................................. 381

antealesidota.............................. 379

blountensis............................... 3 79

calliwnoides.--........................... 379

aff. censors..._-_-___________.------_.___- 378

Crassispira? cf. Crassispira? censors........... 378

pira censors bullbrooki.................. 379

hobsoni............................... 379

magna................................ 380

pennae................................ 379

trinitatensis........................... 379

---.........._..... ---- 377

hispaniolae................................ 381

inaequistriata ............................. 378

jamaicensis............................... 381

maciknta............................... 379,380

maonisriparum............................ 381

militaris................................ 379,380

perspirata................................. 379

praeconsors... _ - _---- ------..--- - 379

ritanida.................................. 381

(Crass.sp.ra) henekeni leptalea............ 318,

S77-S78, pis. 58, 65 

(Crassispirella) cymation.......... 318,S8., pi. 58

fyZoessa....---._-.--_-----318, S8/,pl. 58

sp.................................. 314,380

(Hindsiclava censors censors..------------. 318,

324, 325, 378-380, 382, pis. 58, 65 

pyrgomi..... ___.____..__-_.325, 380, pi. 62

Crassispirella................................. S80

cratera, Terebra............................... 408

Crepidula plana.............................. 323

sp........................................ 314

cristata, Pleurotoma......................... 370,371

Crucibulum................................... 432

sp. _-- ---___-_-_-_---.---- 310,317

(Crucibulum) chipolanum................. 323

(Dispotaea) springvaleense................. 323

cruziana, Cruziturricula....................... 369

Pleurofusia (Cruziturricula)...--.----.-. 368,370

Cruziturricula.............................. 368-369

arcuata................................... 369

cruziana.................................. 369

Cryotunis.................................. 360,394

Cryoturrisf hillsboroughensis.................. 394

cryptopleura, "Mangelia"..................... 393

cr_<Ms, Scaphander (Scaphander)--------------- 305,

306, 316, 420, pi. 48 

?, Scaphander (Scaphander).............. 316

Page 

Ctenilyria..................................... 304

coroni..................................... 304

ctenista.... .._..,_..._.................. 304

ctenista, Ctenilyria............................ 304

cttbensis, Actaeon............................. 415

Tornatetta (Actaeon)...................... 415

Cucaracha formation.... .... ....-.......- 311

cucurrupiensis, Terebra (Panaterebra)......... 320,

324,407-405, pi. 61 

(Parafere&ra) cf. T.................... 407

Culebra formation.......................... 309-311

cybele, Drillia................................. 375

Cyclophorid, genus?--.---.-----.----..---.-- 314

Cyclostremiscus (Ponocyclus) pentagonus...... 323

cydia, Drittia.................................. 375

Cylichna? aff. Cylichnella bidentata............ 419

Cylichna costata..... ............._.._..... 420

Cylichnella..... ............................... 419

atacata............_..................... 419

stibara................... 320,324,4/0, pi. 62

bidentata.................................. 419

triticumtritonis............................ 419

Cylichnina.................................... 4%3

cylindrica, Volvula............................ 425

parallela, VolvuleUa (Volvulella)........... 320,

322,324,4-tf, pi. 62 

cymatias, "Mangelia"................ ......... 394

cymation, Crassispira (Crassispirella). 318,381, pi. 58 

Cymatium nicobaricum........................ 307

(Septa) ogygium........................... 307

Cymatophos................................... 322

Cymatophos? acolus........................... 325

Cymatophos subsemicostatus............ ....... 323

veatchi.................................. 311,316

Cymatophos? cf. C. veatchi... ........ 310,311,314,316

Cymatophos veatchi veatchi..................... 323

Cymatosyrinz-..-........................... 374,388

ac.inica..-................................ 374

lunata.................................... 374

aclinica............................... 374

Cymatosyrinxf sp. ind------------------------ 375

cymella, Cythara.............................. 393

Cymia........................................ 399

Cypraea chilona......................... 307,311,313

cf. C. chilona............ 307,310,311,313,314,316

(Muracypraea) henekeni................... 324

Cypraedia........... ....___........_.... 304

subelegans................................ 303

aff. C. subelegans.------------.*--------.- 303

"Cythara" barbadoides......................... 393

basiHssa. _ _ _ __ ............................ 391

Cythara cercadica............................. 390

cf. cercadica.........--.-.--------.-------. 390

"Cythara" chariessa........................... 393

Cythara cymella............................... 393

heptagona................................. 390

"Cythara" isabellae............................ 393

Cythara, terminula costaricensis................. 390

sp._._.._.___.........-..-....---..-..-.-- 390

"Cythara" (Brachycythara?) cf. terminula...... 391

Cytharella ................................ 392,393

Cytharellaf................................... 392

cf. C.compsacosta........... 320 39S-39S, pi. 60

Cytharella limata.............................. 393

D

fita.Zi, _4.nci-.ros_/rin.r ...-...-..-. ...... 371

^.2/s.................._..-.... 422

-SaZcis.....-....._--.---.----------- 329

Bathygalea................................ 325

ConMS.............--------------- 354,355

Pleurotoma............................... 378

(Drilia)............................. 378,379

oJaWi, Solenosteira.........---------------- 323

medioamericana, Solenosteira ......... ... 323

subsp., Solenosteira.-.... . ... ....... 325

Daphnella.................................... 402

barttcM................................... 402

lymneiformis.-.................... ..... 402
INDEX 445

Daphnella Continued

pagera........................... 320,403, pi. 61

sp______.__-_---__----__-----___-----_ 320,403

" Daphnella" sp............................... 402

Dapknella?sv_......................... 320,402,403

Daphnellinae.-.---------------------------- 360,402

Darbya....................................... 377

Darbyaf...................................... 377

(Buridrillia?) sp_................ 325, 377, pi. 63

dariena, Cancellaria? cf. C.................... 336

Cancellaria (Cancellaria)................ 335,336

Cancellaria (Cancellaria) epistomifera...... 318,

323,325,335-337, pi. 52 

Cancellaria (Cancellaria) aff. C. epistomifera........................ 322,325,336,337

trachyostraca, Cancellaria.................. 336

var. Cancellaria........................... 336

darienensis, Cancellaria...................... 336

dariensis, Mitra (Concilia)..................... 323

decapitata, Retusa............................. 424

decaptyx, Cancellaria (Narona).............. 318,342

decussata gatunensis, Distorsio (Rhysema). 322,323,325 

defuniak, Cancellaria.......................... 335

Ithycythara.. 320, 324, 39J-392, 393, pis. 60, 64 

delecta, Xenophora............................ 323

dentalis, Amarophos........................... 325

dentifera, Clathurella (Glyphostoma)........... 400

dentiferum, Glyphostoma (Glyphostoma).......- 320,

324, 399-400, 401, pis. 60, 66 

dentilabris, Terebralia......................... 311

dentilabrisf, Terebralia...................... 310,311

depressa, Solariella............................ 306

Vaginella........ ....................... 307,427

diadela, Cancellaria (Pyruclia)................. 318,

325, 338-339, pi. 53 

diadelaf, Cancettarta (Pyruclia).............. 322,325

Diarecallus.................................... 322

dicheres, Terebra (Oreoterebra).... 313, 316, 404, pi. 50

dimidiata, Oliva............................... 434

Dimya........................................ 312

dinota, Cancellaria (Euclia)...... 318, 340, pis. 54,56

Dolostoma........................ 320, 898, pi. 61

Dirocerithium................................. 304

ame....................................... 303

whttfteldi.................................. 303

dislocata, Terebra.............................. 409

dislocatum, Strioterebrum.................... 410,413

Dispotaea..................................... 432

Distorsio (Rhysema) decussata gatunensis. 322,323,325 

DolabellaaldrichL........................... 420,421

doliolum, Atys................................ 423

Dolostoma................................... 322,397

anorhepes........... 320, 324, 397, 398, pis. 60, 66

dinota............................ 320, 398, pi. 61

domingensis, Conus........................... 354

domingensisf, Conus........................ 350,351

domingensis, Conussymmetricus............. 353,354

Petaloconchus............................. 430

dominicensis, Tunis (Bela).................... 394

Drmia...................................... 361,374

censors___._....-......-.. .-..-.-.. 378

bullbrooki............................. 378

portoricoensis.......-........---.--..- 378

Mnttatensts.............-. - -- 378

cybele..................................... 375

cydia...................................... 375

euphanes.................................. 375

eupora.............. ............ ... 379

"Drittia" eurysoma............................ 374

Drtttiafusinus................................ 369

gatunensis................................ 381

isthmica................................... 387

Ussotnpis...-....-..-..-..-.-.- --- 388,389

macilenta rectaxis........................ 378,379

puertocolombiana.......................... 381

riogurabonis............................... 375

aff. D. riogurabonis........................ 375

subperpolita............................... 375

gem machapoorensis.................. 362

sanctiandreae......................... 362

Drittia Continued

venusta................................... 381

zooki..................................... 375

"Drittia" sp._................................ 375

Drillia (Crassispira) zooki..................... 375

(Neodrillia) riogurabonis eurysoma......... 318,

324, 374-375, pis. 58, 64 

Drillia? (Neodrillia?) sp..................... 305,375

dumbauldi Terebra........................... 408

E 

eburneola Marginella......................... 330

Eburnospira.................................. 330

Echinolampas cf. E. lycopersicus.............. 312

semiorbis................................. 317

Echinophoria................................. 307

Ectinochilus.................................. 304

gaudichaudi............................... 303

ci. E. gaudichaudi......................... 303

Ectracheliza................................... 431

ectypha Agladrittia (Agladrillia) acaria........ 320,

324, 387, pis. 59, 64 

ecuadoriana, Pleuroliria (Polystria)............ 322,

325, 355, pi. 62 

Polystria oxytropis........................ 365

Scobinella. ...................... 325,373, pi. 62

egouen, Egouena.............................. 331

Egouena..................................... 331

egouen.................................... 331

egregia, Balds................................ 329

eileta, Hemipleurotoma........................ 361

elachista,Acteocina................... 320,4/8, pi. 62

elegans var., Ancistrosyrim................... 371

Cochlespira............................... 371

Limacina.............................. 427

Mangelia................................. 393

Spiratellainflata............ 320, 324, 407, pi. 66

Cancellaria........................... 344

Ellobiidae.................................... 429

Ellobiinae.................................... 429

Ellobium................................... 313,429

pellucens............................... 316,429

aff. E. pellucens............. 313, 316, 429, pi. 48

stagnalis-.----.-.------------.--------.--- 429

elocata Carinodrillia.......................... 376

(Carinodrillia) cf. C.......... 318, 375, pi. 58

elongata Ithycythara.......................... 392

Ithycythara cf. I.................. 320, 392, pi. 60

emendorferi Hannatoma...................... 303

Hannatoma cf. H....................... 303,304

Hannatoma? cf. H........................ 304

emersoni Conus............................ 346,353

empera Clathrodrillia......................... 383

Emperador limestone member of La Boca

formation ..-....-....--.--.- 311-317

enae, Pleurofusia.............................. 367

engonata, Cochlespira....................... 370,371

enneacyma enneacyma, Agladrittia (Agladrittia). 320,

386-386, pis. 59,64

subsp., Agladrillia (Agladrillia)......... 320, 386

entemna, Euclathurella........................ 397

Eocene or Oligocene series.................. 304-305

Eocene series....-.........-..-.-.......---- 303-304

epacta, Mitretta............................... 306

ephnidia, Solariella............................ 306

epicasta, Glyphostoma......................... 399

Epidromus testaceus........................... 432

epistomifera, Cancellaria..................... 336,337

dariena, Cancellaria (Cancellaria)......... 318,

323,325,335-337, pi. 52 

(Cancellaria) aft.C..... 322,325,336,337

epistomifera, Cancellaria..........-..--.- 336,337

lipara, Cancellaria (Cancellaria)........... 318,

323,337, pi. 52 

Epitonium scipio.............................. 325

sp........................................ 314

("Depressiscala") eucteanum.............. 325

epomis, Gonysycon............................ 306

Gonysyconct. G.........................-- 306

Eratoidea....- ............................ 330

Page 

estrellana, Agladrillia.......................... 386

euancycla, Marginella................... 306,307,330

eucharis, Euclathurella (Miraclathuretta)....... 320,

324,397, pis. 60,66 

Euchilodon morierei........................... 372

Euclathurella................................. 396

entemna.................................. 397

Euclathurella? liveoakensis............. ........ 396

Euclathurella (Euclathurella) vendryesiana... .. 320,

324,395, pi. 60

(Miraclathuretta) eucharis. 320,324,397, pis. 60,66 

Euclia.............................. 334,339,342,344

eucteanum, Epitonium ("Depressiscala")...... 325

eucteanus, Typhis (Talityphis)........ 318,434, pi. 63

Euglyphostoma.......................... 322,399,407

Eulima....................................... 326

acicularis............................... 326,327

acuta......................... 318,323,327, pi. 51

auricincta................................. 327

ischnon................................... 327

jacksonensis............................. 305,326

cf. E. jacksonensis............ 304,305,325, pi. 48

nobilis.................... 318,323,325-327,pi. 49

rectiuscula................................ 327

robusta................................... 328

sarissiformis.................. 318,323,327,pi. 49

scotti...................................... 327

towensendi................................ 327

sp........................................ 327

Eulimidae.................................... 326

Eumetadrillia. ........................... 387,401

euphanes, Drillia............................. 375

Eupleura thompsoni..... ---------------------- 323

eupora, Drillia................................ 379

euryhedra, Sinum............................. 323

eurys, Atys (Aliculastrum)....... 320,422-423, pi. 62

eurysoma, Drillia (Neodrillia) riogurabonis.... 318,

324,374-375, pis. 58,64 

"Drillia"................................. 374

eurytera, Valuta............................. 323,325

Valuta alfaroi........................... 324,325

euthynta, Sthenorytis toroensis................. 325

excolpa, Niso....----------------------------- 329

exilis,Acteon.... ....-..--...-.-..--.--.......- 416

F 

falconensis, Xancus........................... 323

Xancus validus........................... 324

Fascioliria gorgasiana......................... 323

subsp... -_ .-. .. -.- - 323

sp............................... ... 314

favosa, iepjdocydma.------------------------- 309

felis, Carinodrillia............................. 376

fenimorei, Pkurofusia................... 325,367,368

Pleurofusia (A. P................... 325,367,358

fergusoni, Conus............................ 346,351

Ficus carbasea ................................ 317

carbasea........................ 317,323,325

micronematica.............. 310,311,313,314

cf. F. carbasea micronematica.............. 316

mississippiensis........................... 305

cf. F. mississippiensis......-..-..-...-...- 305

pilsbryi................................... 307

cf. I. pilsbryi............................. 307

sp. (F. ventricosa group).-------- ------ 308

Fieldwork and acknowledgements. ----------- 300

fischeri, Globularia...................... 306,307,313

(Globularia) aff. G.......... 306,307,313,314

flammea, Terebra.............................. 406

Floribella.-............................ 317,420-427

aldrichi....................... 313,316, $1, pi. 50

floridana, Terebra..............-..----..-.---- 404

floridanus, Conus............................. 356

costaricensis, Conus....................... 356

Petaloconchus at. P.................... 324,430

fossilis, Retusa sulcata........................ 424

fraternalis, Philbertia (Nannodiella)........... 395

frisbeyae, Conus........................------ 357

fuegiensis, Pkurotoma (Surcula)............... 387
446 INDEX

Page 

fugax, Anachis (Costoanachis) rnira............ 323

fungia, Architectonica...-.--.. .............. 305

Architectonica cf. A....................... 305

funiculatum, Trigonostoma-................... 345

fusinella, Pleurofusia.......................... 368

Turns (Surcula).......................... 367

fusinus, Drillia............................... 369

Pleurofusia............................... 367

fusinus, Pleurofusia (Cruziturricula)...... 318,

324,368, 369-370, pis. 57,65

sanctidavidis, Pleurofusia (Cruztturricula). 368,

369 

Fusinus?sp-.......................... 310,314,316

Fusisyrinx.................................... 368

Fusiturricula........................... 360,367,368

latira..................................... 369

gabbi, Conus---------------------------------- 359

Ortfcawtoz. --------------- 313, 314, 432, pi. 49

Terebra................................... 408

gabrielensis, Pachycrommium?----------------- 305

0aZae, Brachycythara-.--.---.----------------- 393

Qaleodea nodosa... ._-----_--_ -----.--------- 304

Oaleodeaf cf. 0. nodosa..---------------------- 304

galvesia, Turritella-...-...-................... 305

Oamopleura..... .............................. 428

taurinensis................................ 428

garzaensis, Olobularia......................... 305

Gastropod faunal summaries and age of 

formations ............................. 303-326

Gatun formation...--_..-__-.--.--------.-- 317-324

Gatuncilloformation----------------------- 303-304

gatunense, Prunum (Microspira) -............. 318,

323, 332, pi. 51 

Strioterebrum............................. 409

Strioterebrum cL.--.-.-..............----- 409

kugleri, SMoterebrum--------------------- 413

gatunensis, Antillophos (AntUlophos) candei.... 311,

316,323,336

Antillophos? (Antillcphos?) cf.^4. candei... 310,

311,314,316 

Clathrodrillia...... 318, 324, 381-383, pis. 58, 65

Distorsio (Rhysema) decussata....... 322,323,325

Drillia-.................................. 381

Marginella................................ 332

Paziella (Panamurex)..................... 323

Pleurotoma (Drillia').-....--------........ 381

n. var., Pleurotoma (Drillia')............ 381,382

Strombus................................. 323

Terebra (Oxymeria)..................... 409,410

kugleri, Terebra (Strioterebrum).... ------ 409,410

Turbonilla.............................. 414-415

gatunensis, Turritella..................... 323

gatunensis?, Turritella-----..........-....--.- 317

gaudichaudi, Ectinochilus...................... 303

Ectinochilus cf. -E.... .... ..._....... 303

gausapata, Terebra............................ 411

(Strioterebrum)........................ 411

herviderana, Terebra....................... 412

gausapatum, Strioterebrum.-..-.-.-.-.--...... 320,

324,408, 411-41S, 413, pi. 62 

gavilanensis, Scobinella morieri.............. 372,373

ffaza, Conus................................... 356

ConMs multiliratus...................... 356,357

Oemmula..................................... 360

" Oemmula".................................. 307

Oemmula arnica............................. 307,360

cf. G. arnica......................... 306,307,360

hindsiana................................. 362

machapoorensis................. 318,324, 362-363

periscelida.............................. 360,362

vaningeni.-. 316,318,324, 361-362, 363, pis. 57, 65 

cf. 0. vaningeni..................... 314,316,360

sp...................................... 310,360

''Gemmula" sp...................... 306, 363, pi. 48

gertrudis, Pleurotoma (Genota)................ 396

Gibberula..................................... 333

i, ClathrodrUlia....................... 382,383

Page 

gigas, Lepidocyclina. -------------------------- 309

giraudi, Lepidocyclina- ------------------ 309,312,317

Olobularia----.......--.-------.-. 305,306,307,317

fischeri ........................... 306,307,313

garzaensis................................. 305

parisiensis. ------------------------------- 305

(AmpulellaT) warao..--...----------------- 305

(Ampulella) sp..........-.....-..---.--.. 305

(Olobularia) aff. 0. fischeri-..... 306,307,313,314

Glyphostoma...-.... ................ 322,398-399

amicta rintriada........................... 395

conradiana..........-.-........ --------- 399

epicasta................................... 399

guppyi.................................... 400

locklini..-............................... 399

partefilosa. ------------------------------- 401

airerao_.................................. 399

striata.................................... 305

typhon.............-..---.....-.--........ 402

woodringi................................. 400

eoater.---.---.--_--_..-.-.--___.---.-.---. 402 

mansfteldi............................. 402

sp........................................ 400

(Euglyphostoma)................ ........ 322

oZjgoraf...................... 320, 401, pi. 60

(Glyphostoma) dentiferum.. 320,324,399-400,401,

pis. 60, 66 

pyrgota..-....-.......... 320, 400-401, pi. 60

(Glyphostomops) hendersoni....----....... 399

(Rhiglyphostoma) allodapum.. 320, 40#, pis. 60, 66 

Olyphostomops............................. . 399

fffypta, Pleurofusia (Cruztturricula) ---. 368.3?" 

Olyptostyla. ................................ 305,434

panamensis....... ... .. 305, $4-435, pi. 48

ffoZiatt, Olivella (Toroliva)...-.....-..-..-.. 323

Ooniopora cascadensis. ------------------------ 308-

cf. (?. cascadensis.. ....................... 308

Oonysycon.............. ............--....-. 306

epomis.................................... 306

cf. G. epomis ...... ..................... 306

gorgasiana, Fasciolaria.... .... .......... 323

subsp., yoscioZan'a..---------------------- 323

gracilenta, Pleurotoma........................ 396

gracilis,Atys.. ............................... 422

gracilissimus, Conus....... .---...........-- 359

granazonatoides, Conus.... .......... ...... 352

grandis, Niso......................... ...... 329

Granoturris..... .......................... 393

padolina.-- ............................... 393

granozonatus, Conus.. ........................ 359

guanabanum, Strioterebrum...... ......... ... 410

guppyi, Glyphostoma.......... ................ 400

Pachycrommium.................... 306,311,313

Pachycrommium aff. P.................... 306

Pachycrommiuml cf. P._...... 310,311,313,314

guppiana, Stigmaulax. ...................... 323,325

guttata, Valuta......-............. .......... 331

H 

fta&ra, Kurtziella (Cryoturris) ...... 320, 394, pi. 60

ftadra, Bathygalea (Miogalea) .................. 325

hadrum, Strioterebrum oresignum.............. 320,

408, 473-474, pi. 62 

haitensis, Pleuroliria......--........ ---..... 365

Pleurotoma- .............................. 364

Tere6ra................................... 404

halensis, Campanile-.......................... 431

haliostrephis, Carinodrillia............ .. 376

Hanetia .................................. 322,325

Hannatoma. .. ............................ 304

emendorferi. -.........-.--........ 303

cf. H. emendorferi..... .................. 303,304

Hannatoma"! cf. H. emendorferi................ 304

harpa,Acteocina.. .. .................... 418,419

harpiformis, Cancellaria....................... 340

harrisi, Conus............ . ................ 358

Conws burckhardti. ..... 318, 324, 358-359, pi. 57

haneyensis, Conus....--- ....... .... 358

---.--.--------------- 424

Page 

haytensis, Conus.. ...-.. -.-.-----.. -.... .. 351

hebetika, Microdrillia.--..-. ....... .......... 390

Heliacus (Astronacus) ........................ 322

stonemanae ............... ...... ...... 323

helicoides, Neverita (Hypterita) ...... ...... 322, 323

HeZiz coactiliatus. ....... .......... ........ 429

Helminthoglyptidae.. ------------- _ ... ... 429

hematUa, Marginella. ... .......... ........... 330

Hemipleurotoma bitropis. ........ _ ---------- 362

- ... . .................. 361

-------------------- _ ------- _ 430

amaraa--------------------- 318, 430-431, pi. 63

atriformis........ .... ...-.-- ... . .... ...... 306

sfeeref.. --.......-.-...-... ...... ......... 431

truncatus. ...... .... ...... .. ...... 431

(Longiverena) oeciscus.. ................ 306,313

aff. H.oeciscus. ................ 311,313,314

hendersoni, Glyphostoma (Glyphostomops) ..... 399

henekeni, Crassispira ........ ..... 377

leptalea, Crassispira (Crassispira) ... ...... 318,

377-378, pis. 58, 65 

Cypraea (Muracypraea) . .. ... . . -.. 324

Siphocypraea (Muracypraea) ...... .... 323

heptagona, Cythara ....... . ....... ... ... 390

Lepicythara .......... 320, 324, 390-397 , pis. 60, 64

AfawffeHa..- ------------------------------ 390

herculeanus, Campanile... .. .. ........ 316,431

Campanile cf. C..- -------- 312, 316, 437, pi. 51

hernandoensis Campanile .....---. . ..- -. 431 

blackwaterensis, Campanile ...... .. 431

herviderana, Terebra gausapata ...... ... ..... 412

(Sfrfofere&rztm) ----------------------- 412

Heterostegina antulea. ...--- ..-.. -. ----- 309 

israelskyi. ............ ........ 308

hettneri, Cancellaria.-..-...--.--.-.- 339,340

hillsboroughensis, Cryoturris? . . ... ... -. 394

hindsiana, Oemmula..---- - ----- .. . . 362

Hipponix cf. H. jjiZogua.... ------------------- 314

Hipponix? sp.-.-.-. -------------------------- 310

hispaniolae, Crassispira.-... . . .. .. 381 

hobsoni, Crassispira consors ...... ------------- 379

SarcaZa ------... --------------------- 378,379

hoerlei, Trigonostoma. ........ . . . .. ... 334

Holocene series. ...... ---------------------- 325,326

horrenda, Pleurotoma (Drillia) ................ 378

Hyalaea taurinensis. . .--.. .-...... . .---- 428

Hyalina. .. ..- .....-.. . 330

peZZwczda-- ------------------------------- 330

hypograpta, Ringicula... ...... ...... ..-.---- 417

imbricata, Persicula..... .......... .. .... 333

imitator, Conus.. ....................... 325,346,354

imitator"!, Conus.-............ ..... 322,325,356

imitator, Conus aff. C..... .. .......... 354,355

imitator, Conus ....... 318,324, 354-350,358, pi. 55

tew, Conus.... . -.-.- - .... 354,355 

Conus cf._----.----.--..----------.--- 354

subsp., Conus.......................... 355

inaegualis, Terebra.... ........................ 406

inaequistriata, Crassispira..................... 378

indocayapum, Strioterebrum-......... . . 320,

324, 408, 47JM73, pis. 62, 66

inflata elevata, Spiratella...--... 320, 324, 4#7, pi. 66 

Informally designated Atlantic and Pacific 

muck......-..---....-......-.---..--...-. 325-326

insulare, Trigonostoma.. ---------------------- 345

Trigonostoma cf. T..._..._____... 318,345 

intermedia, Semicassis....-.. . -..-. ...... 307

interrupta, Marginella............... .. - . 333

interruptolineata, Persicula.. ....-.-.-. .. 333

Introduction. -...---.-.-----.....-....... 299-300

isaacpetiti, Terebra (Oreoterebra)............. 320,

324, 405-406, pi. 61 

isabellae, "Cythara" ........................ 393

ischna, Strombiformis.......................... 326

ischnon, Eulima.....-------------------------- 327

Strombiformis-......... _____._.-..- 327
INDEX 447

Page 

ischnum, Strioterebrum...................... 306,408

islacolonis, Aphera............... 318, 323, 844, pi. 56

Cancellaria (Aphera)....--......---...--.. 344

islalindae, Clathrodrillia-...................... 384

Clathrodrittia cf. C........... 320, 38,$, pi. 58

israelskyi, Heterostegina....................... 308

isthmica, Agladtillia (Eumetadrillia).......... 320,

387-388, pis. 60, 64 

Drillia.................................... 387

Ithycythara.................................. 391,393

compsacosta............................. 392,393

defuniak........ 320, 324, S91-39S, 393, pis. 60, 64

elongata................................... 392

cf. I. elongata.---...-.--....---. 320,392, pi. 60

jacksonensis, Eulima........................ 305,326

Eulima d. E................. 304, 305, 326, pi. 48

Pachycrommiumf......................... 305

Scaphander............................. 305,420

(Scaphander) cf. S . 304, 305, 420, pi. 48

jacululum, Balds (Balds)-. 318, 323, 327-328, pi. 51 

Melanella(Eulima)........................ 327

jamaicensis, Balds............................ 328

Crassispira............................... 381

jaspideus, Conus.............................. 347

Josephina..................................... 364

K

karsteni, Architectonica (Architectonica) nobilis.. 325

QmceBaria -- .- . 339,340

Knefastia..................................... 369

kugleri, Strioterebrum gatunense............... 413

Terebra (Strioterebrum) gatunensis....... 409,410

Kurtzia....................................... 393

KurtzieUa. .............................. 393,396,397

(CrjroJwrris) habra................ 320,394, pi. 60

sp....................... 320,394-395, pi. 60

(Kurtzietta) pagella ...... 320,393,394, pis. 60,64

...... 320, 39S-S94, 397, pis. 60,64

La Boca formation, including Emperador

limestone member_........... 311-317

laevescens, Cancellaria........................ 335

laevifastiolum, Strioterebrum................. 409,413

laevigata sublaevigata, Sconsia.................. 323

lagua, Cancellaria... ................_....... 341

laricum, Campanile (Portoricia)............... 431

larvatus, Conus............................... 359

latilabris, Pterynotus. ......................... 434

latira, Fusiturricula........................... 369

Pleurofusia (Cruziturricula)............. 368,370

Latirustaurus................................ 325

(Polygona) anapetes........... ........... 325

latissima pilsbryi, Marginetta................... 332

Prunum................... ........... 332

latissimum, Prunum.......................... 332

lavelana, Cancellaria.......................... 341

Persicula venezuelana..................... 333

toeMeono, Mangilia........................... 399

lavillei, Conus............................... 352,353

lavinia, Turricula.. ..............__._..... 369

" Turricula" ..............._...._..... 369

Turricula (Surcula) ...................... 369

Zapinoides, TurricuZo......_................. 369

leander, MargineUa............................ 331

Volvarina........................ 318, SS1, pi. 51

Lecallia... ................................. 305,429

-Letostraca acuta... ............................ 327

ZeZoTidi.^lffZodriZZio.-.......................... 384

Clathrodrilla aff. C.. ... 320,384, pi. 59 

Zeoninus, COKUS.---.......................... 348

Lepicythara................................... 390

camaronensis.............................. 391

heptagona............ 320,324,390-391, pis. 60,64

terminula................................. 391

Page 

Lepidocyclina............................... 316,317

asterodisca................................ 308

favosa.................................... 309

gigas..................................... 309

giraudi.............................. 309,312,317

miraflorensis.............................. 317

parvula................................... 309

pustulosa tobleri.......................... 304

vaughani... ............................_ 309

waylandvaughani...-.-.............--..-- 309

yurnagunensis............................ 309

morganopsis.......................... 309

sp........................................ 308

Leptadrtllia................................... 374

Leptarius... .........................._..... 434

leptus.................................... 324

Leptegouana.---. ..........-..-.-.-..--.-....- 331

leptus, Leptarius... ........................... 324

Psilarius................................. 323

lessepsiana, Strombina (Strombina)............ 323

Leucosyrinx.................................. 370

Moris.................................... 370

nicoya.................................... 370

xenica............................ 318,370,pi. 57

Limacina elevata.............................. 427

limata, Cytharetta............................. 393

limonensis, Mitra (Pleioptygma?).............. 323

MitreUa.-. _._._.......................... 323

Lioglyphostoma......................... 360,397,399

adematum................................ 399

woodringi................................. 400

subsp...-. ...... ..-. ..--.-..... 402

Liotia sp..................................... 314

lipara, Balds (Balds)................ 318,328,pi. 51

Cancellaria (Cancellaria) epistomifera...... 318,

323, SSr.pl. 52 

Sulcoretusa sulcata........... 320,324, 424, pi. 62

Sulcularia.. .............................. 424

lissa, Atlanta (Atlantidea)..................... 323

Lissacteon.................................... 416

lissotropis, Drillia........................... 388,389

Pleurotoma (Mangilia).................. 388,389

Syntomodrillia............................ 389

listrota, Turritetta............................. 306

listrotum, Strioterebrum.. 305,306,307,408,409, pi. 48 

Littorina angulifera........................... 311

aff. L. angulifera........................ 310,311

lius, Conus cf. C.............................. 355

Conus imitator.......................... 354,355

Onus cf. imitator......................... 354

liveoakensis, Euclathurellaf.................... 396

locklini, Glyphostoma......................... 399

longa longa, Mitra (Candlla).................. 323

longirostropsis, Pleurofusia.................... 366

Pleurotoma............................. 366,367

Lophiotoma.................................... 364

lophota, Vaginella................ 306,307, 487, pi. 49

loroisi, Terebra...----........--..-..-......... 408

loroisii, Terebra...........-...----.-----.---.. 408

lucasensis, Colubraria....................... 432,433

lunata, Cymatosyrinx......................... 374

aclinica, Cymatosyrinx.................... 374

Pleurotoma............................... 374

lycopersicus, Echinolampas cf. E.............. 312

lymneiformis, Daphnella....................... 402

Pleurotoma............................... 402

Lyriafsp..................................... 314

M

machapoorensis, Drillia vaningeni.... .......... 362

Oemmula....................... 318,324,362-363

Tunis vaningeni.......... ................ 362

madlenta, Crassispira....................... 379,380

rectaiis, Drillia......................... 378,379

Pleurotoma sp. aff. P. aksidota.......... 378,379

macneili, Cancellaria.......................... 335

(Cancellaria) aS. C.......... 318, SS5, pi. 52

Page 

magna, Crassispira consors.................... 379

magne, Pleurotoma (Drillia) aksidota.......... 379

magnifica, Scobinella.... .................._. 373

mahogani, Conus.............................. 354

makista, Balds................................ 329

maldonadoi, Cancellaria (Euclid)............ 339,340

Malea......................................... 322

camura................................... 323

sp-_....--..- -- - - - 317

mandnella, Agaronia testacea.................. 323

Mangelia...................................... 392

"Mangelia" cryptopleura...................... 393

cymatias.................................. 394

Mangelia elevata............................... 393

heptagona................................. 390

Mangeliinae.--.-..-....-.---.------------ 360,390

Mangilia aguadillana.......................... 397

arteaga. ................................. 393

consentanea............................... 394

lavelleana................................. 399

melanitica................................. 395

torn.....-....--..---------------------- 392

mansfieldi, Glyphostoma zoster................. 402

manzanittaensis, Conus....................... 352

Conus aemulator..... .................. 351

inaoica, Balds............................. - 328

maonisriparum, Crassispira................... 381

mareana, Clathrodrillia........-..-..-.-------- 383

marginatus, Conus...........-.-----.------- 348,356

Conwscf. C........._-.--------------- 348

&OMSSOCZ, Coims..------------------------- 358

Marginetta.................................... 330

amygdala..............,.....------------- 331

a0ena.---__--.-..-.----------------------- 330

coniform is............ --------------------- 332

eburneola............---------------------- 330

----------------- 306,307,330

---. ----------------------- 332

------------------------- 330

intenupta..... -................. -.- 333

latissima pflgfirj/i..----------------------- 332

leander...................... . 331

mitrella..................... ..... 330

mollitor. .-...-.-..---.---.-----.-- 306,307,330

n«ido ------------------------------ 330

aff. M. nitida..._.._-._ _ - 331

serrate..-------------------------- 330

(Bullata) mindiensis.-------..----.-- 332

(Eratoidea) aff. M. mollitor................ 305.

306, 307, 330, pi. 48 

sp--_ -- -- - 314

Marginellidae .- - --- 330

Marine member of Bohio(?) formation.---- 304-305

Mo*«irta .---.- -.---- .- 343

mataracana, TurriteWa-.- . ..... -.-.... 323

mazei, Conus................................. 359

mcgintyi, Conus............................... 359

medioamericana, Solenosteira dalli......-......- 323

Megatylotus.................... .......... - 307

Melanella astuta............................... 328

(Eulima) cercadica..........-....-.-...-.- 326

jacululum............................. 327

melanitica, Mangilia..-.........-....--....--- 395

Nannodiella cf. 2V - - ----- 320,395

meZonpena consors, MeZonpena .--. .--- -.- 323 

Melongena melongena consors.................. 323

sp-.-..- ............ -....-.-- 310,314

meroensis, Turritella.......................... 307

mesata, Mso (Niso)......... 318, 329-330, pis. 49, 51

metalium, Calliostoma (Calliostoma)........... 325

MeteZa. _.-... . . -- --- 322 

sp---- - - .-- -"- 310

metuloides, Phos........................ 359,361,410

mexicana, Pkurotoma (Glyphostoma)..-.------ 399

mexicanus, Antillophos (Autillophos).....-..-- 323

meyeri, Oligotoma...... . - ---- -------- 389

PZeuroZoma. _- . - .. ... . 389 

micratracta, Volvuletta (Volvulella). 320, 324, 425-4^5
448 INDEX

Page 

Microdrillia................................... 389

comatotropis.............................. 390

hebetika................................... 390

tersa...................................... 390

trim........... 320, 324, 374, 389-390, pis. 59, 64

Micromelo (Abderospira) chipolana............ 419

micronematica, Ficus carbasea....... 310,311,313,314

Ficus cf. F. carbasea...................... 316

Microspira.................................... 331

midiensis, Terebra spirifera.................... 413

muitaris, Crassispira........................ 379,380

Turns (Drillia)-------.................... 378

milleri, Trigonostoma......................... 334

mimeticum, Cerithium (Thericium)............ 306

mimys,Agladrillia............................ 385

mindiense, Prunum........................... 332

mindiensis, Marginella (Bullata).............. 332

Minioliva myrmecoon.......................... 326

minuta. Volvulella......................_..... 426

miocaenica, Uxia............................. 335

Miocene series.-..---...-.--..-...,.---.-.- 309-325 

Miogalea...................................... 325

Miogypsina antiUea........................... 309

mirafugaz.Anachis (Costoanachis)............ 323

mira,Anachis (Costoanachis).............. 323

Miraclathurella......................... 360,595-597

miraflorensis, Lepidocyclina................... 317

mmznda, Ancistrosyrinz....................... 371

Pkurotoma............................. 371,372

Mirascapha................................... 420

mississippiensis, Ficus........................ 305

Ficus cf.F............................... 305

Afftro (CbnriBa).. .__ ____ 322

dorfenafa............_---------_... 323

longalonga........................... 323

Mitraf (Camilla?) sp......................... 310

Af#ro (Pleioptygmaf) limonensis.............. 323

(Tiara) sp................................ 310

Mitrella acanthodes.. .......................... 306

epacta-------.-.-.-.-.--..-.....--.-....-- 306

limonensis................................ 323

aria...................................... 306

Mitrellal sp.................._.............. 314

mUrella, Marginalia........................... 330

Modulus sp.-...--...................._..._... 314

rooZis, ConMs. 318, 324, 346, 348, SSO-S51, 352, pi. 55 

Conus aff. C_............................. 351

bravoi, Conus............................. 348

mollitor, Marginella..................... 306,307,330

(Eratoidea) aff. Af-305, 306, 307, 530, pi. 48 

monida, Terebra (Strloterebrum)............... 414

monidum, Strioterebrum.... 320, 324, 408, 4/4, pi. 63

montserratensis, Cancellaria................... 340

morganopsis, Lepidocyclina yurnagunensis..... 309

morierei, Euchilodon. ......................... 372

Scobinella.......... 307, 318, 324, 57M75, pi. 58

Scobinella afl. S.................... 306,307,575

gavttanensis, Scobinella.................. 372,373

Morum antiguum............................. 305

chipolanum............................... 313

(Cancellomorum) cf. Af. antiguum ......... 305

cf. M. chipolanum................... 313,314

("Ontocidfa") cf. Af. antiguum............ 305

cf. Af. chipolanum..................... 316

multiliratus, Conus agassizi.................... 356

woKz -----------.------.....--........ 357

ffozo, Cbnua............................ 356,357

muttiliratus, Conus... 318,324,556-557,358,pi. 57 

spiekeri, Conus........................... 357

multistriata, Strepsidura...................... 435

muttistriatus, Conus........................... 356

AfMracj/proea..-.._..................._... 383

Murezpolynematicus........................ 311,316

rhyssus................................... 433

Pfrffo...................................... 363

(Murezl) cf. Af. polynematkus...... 310,311,316

(Afwrei) recMTO'rosMs..............__... 322

recurvirostris...................... 323

(Pteronoftw) ferfafs_.....______... 433

Murex Continued

(Siratus) cf. Af. polynematicus. - - 310,311,314,316 

Murexiella (Subpterynotus) textUis. ------------ 433

Muricidae-----..--.--.----.-----.-----.-..--- 433

Muricinae................................._ 433

musaensis, Conus............ 318,324, S47-S48, pi. 57

myakkanus, Rictazis.......................... 416

myrmecoon, Minioliva......................... 326

N

nonet, Olobularia (Ampulellal)................ 305

Nanarius..................................... 322

nancellaria, Cancellaria........... 318,54/-S42,pi. 53

Nannodiella.................................. S9B

amicta.................................... 395

at. N. melanitica........................ 320,595

rintriada................. 320,324,395, pis. 60,64

Narona................................. 334,542,344

Nassariidae.......-..............--.-.---.--. 434

Nassarius (Uztta) cercadensis................. 323

( Uzital) praeambiguus.................. 314,316

Natica sp., operculum.----------------------- 314

(Natical) bolus............................ 323

(Naticariusl) sp.......................... 310

nelsoni, Strioterebrum......................... 414

Nemocardium............................... 316,317

NeodrOlia.................................... 574

riogurabonis............................... 374

Neritasp..................................... 314

Neretina...................................... 311

(Fjffo?) sp...--. ..-- - 310,314 

Neverita bolivarensis bolivarensis............... 305

Neveritafsp................................ 310,314

Neverita (Olossaulax)......................... 322

bolivarensis tapina..................... 305

reclusiana xena....................... 322

(Hypterita)............................... 322

helicoides............................ 322,323

New generic and subgeneric names...... ..- 303

newcombi,Adrana.-- -- ---- -.-.......... 326

Nicema....................................... 322

nicobaricum, Cymatium....................... 307

nicoya, Leucosyrinz........................... 370

M*o._ ....... ...-.......... ............ 529

ezcoZpo... . ................. .. 329

grandis................................... 329

striatula.................................. 329

umbilicata................................ 305

CZVteo) 7ne«o<0 ......... 318, 529-350, pis. 49, 51

umbilicatal.............. 304, 305, 329, pi. 48

nitida, Marginella............................. 330

Marginetta aff. M......................... 331

nobilis, Architectonica (Architectonica)... 307,313,322 

(Architectonica) cf. A................. 316

karsteni, Architectonica (Architectonica).... 325

nobilis, Architectonica (Architectonica)...... 323

subsp., Architectonica (Architectonica)... 313.314 

Eulima................. 318, 323, 525-527, pi. 49

nodosa, Galeodea.............................. 304

Oaleodea? cf. (?.-. ...... ......... 304

Northianorthiae............................... 316

Northlat d.N.northiae.-................... 314,316

northiae, Northia.............................. 316

NorthiafcLN--------................. 314,316

Nummulitespanamensis .................... 308

O

S, TfypAw (Talityphis) alatus............. 323

obscura, Colubraria...... 318, 322, 323, 452-453, pi. 63

obscurus, Triton.............................. 432

odopoia, Terebra............................ 316,404

oeciscus, Hemisinus (Longiverena)........... 306,313

(Longiverena) afl. H............ 311,313,314

oedemata, Atys................................ 423

ogygium, Cymatium (Septa)................... 307

Oligocene series..__-..._..._.__... 305-309 

OligotomameyerL............................. 389

Page 

Oliva......................................... 434

dimidiata................................. 434

(Oteo) sp................................. 314

(Strephonella)............................ 322

colpotus............................. 323,454

plicata...........--..--..........--. 324,434

OliveUa sp.................................... 314

(Niteoliva)terryi.......................... 323

(Toroliva) goliath.......................... 323

Olivia, Pleurofusia..... ___.-.-..__.-.-.-. 367

Olividae..-.. ... .................... 434

OJivinsa...................................... 434

olssoni, Glyphostoma (Euglyphostoma). 320,401, pi. 60 

Turritetta.......... ........._._...._...... 305

onzola, Clathrodrittia.......................... 382

Scobinella... ............................ 373

Oostrombus.... ........_..-__._ ... 304

chiraensis......... ........................ 303

aff. 0. chiraensis..........-.--.-.-..-...- 303

tournoueri... __...-_...................... 303

Oreoterebra...................... 313,322,405-404,406

oresignum, Strioterebrum...-.--.----...--.--.- 413

hadrum, Strioterebrum...-- 320,408, 418-414, pi.62

oresignum, Strioterebrum.. 320,324,408,4/5, pi. 62

Orthaulaz....................... 306,307,311,317,432

cf. 0. aguadiUensis........................ 432

gabbi.......................... 313,314,452,pi.49

pugnaz................................. 306,307

cf. O.pugnax....................... 306,307,432

sp.. 306,307,316,432

Orthaulaz? sp......... . ..... 310

oryza, Actaeonidea............................. 416

Rictazis............................. 320,324,4/5

ozia, Mitrella.................................. 306

ozytata, BuUa (Volvula) cf ......... 425

Volvula................................... 425

Volvulella (Volvulella)......... 320,324, J&S, pi.62

ozytropis, Pleuroliria.......................... 365

ecuadoriana, Polystira..................... 365

Pachycrommium.................... 304,305,306,311

Pachycrommium f............................. 317

gabrielensis.................... . - 305

Pachycrommium guppyi...........-.---. 306,311,313

aff. P. guppyi____..... .. - -- 306

Pachycrommium? cf. P. guppyi...... 310,311,313,314

jacksonensis............................... 305

proinum....... ......... . ... - 305

solenaeum..............................-- 303

trtnttateruta...........----- . 308,313

afl. P. ftrinitatensis...................... 308

cf. P. Itrinitatensis..................... 313,314

padolina, Granoturris......------------------ 393

pagella, Kurtziella (Kurtziella)................ 320,

595,394, pis. 60,64 

pagera,Daphnella................... 320, 405, pi. 61

Paleocavolina...... . -------------- 306,307,408

Panama formation___ ..--.. -- 308-309 

panamella, Clathurella.............-..-..---.- 385

panamensis, Olyptostyla......... 305, lfilf-lflt>, pi. 48

Nummulites.........................- 308

Panamurex.. ...--. ------------------------- 322

Panaterebra... 322,407

panthea, Pleurofusia (Cruzlturricula)....--.-- 368

Turricula (Surcula)...................... 369

papulosus, SerpzitorM*-. -------------------- 323

Paraborsonia..........................--.--- 307,375

brassoensis......................-------- 307,374

afl. P. brassoensis. ------------ 306,307,575-574

Paraclathuretta.... ------------------------ 396

paraensis, Colubraria.....--------------------- 433

parallela, Volvula.. ..___ - --- 426

Volvulella (Volvuklla) cylindrica-......-..- 320,

322, 324, 425, pi. 62 

Paraterebra............................. 322,406,407

Paravolvuklla... ...... .- -..- -------- 426

pori«en*J«, Olobularia..............----------- 305
INDEX 449

Page 

parkeri, Turns (Surcula) ...._.........._... 374

partefilosa, Glyphostoma-...................... 401

parvula, Lepidocyclina-....................... 309

patricius, Conus............................ 346,347

paupercula, Bulla........................... 421,422

BvMaria.................................. 421

pavonia, Terebra (Strioterebrum)............... 412

paytensis, Yasila.............................. 304

Yasila aff. Y.............................. 304

paziana, Sulcoretusa-..--.-................... 424

Paziella (Panamurex) gatunensis.............. 323

pellucens, Ellobium......................... 316,429

Ellobium &S. E............. 313, 316, 429, pi. 48

pellucida, Hyalina........-.--...----..---.... 330

Trigona................................... 345

pellucidus, Pterynotus......................... 433

pennae. Crassispira corners......._.......... 379

pentagonus, Cyclostremiscus (Ponocyclus)... ... 323

perdiciana, Cancellaria-.-..-_................. 343

periscelida, Gemmula.-.-. ................... 360,362

permutabile, Bittium.......................... 313

pernobilisf, Sthenorytis........................ 325

Perplicaria................................... 334

cZarfci ---.. .-.__._._.-. ....-I 334

perprotracta, Adrana.......................... 326

Persicula..................................... 38%

cercadensis................................ 333

cowtfjana couviana......................... 334

imbricate.................................. 333

interruptolineata.......................... 333

propeooesa__................._....... 334

»eme»................................... 304,333

venezuelana............................... 333

lavelana ............................ 333

venezuelana..................... 311,316,333

(Gibberula) cf. P. semen.---.. 303,304,333, pi. 48 

(Rabicea) couviana stenygra.... 318,333-334, pl.51

venezuelana amydra................... 310,

311,314,316,333, pi. 48 

persimilis, Volvula. ...........__............ 425

VbZroZeZZa................................. 425

perspirata, Crassispira............ ........__ 379

Peruflcus.. ____........._............... 435

peruvianus, Conus..... .................._ 307,359

Conus cf. C......................... 306,307,359

Xancus........ ..._.......____...... 304

Xancuscf. X............................. 304

perversus, Velates............................. 303

sabsp.?, Velates........................... 303

Petaloconchus..... ........-...-.... ..-. .-. 430

domingensis... ..........-...-.---...--.... 430

afl. P. floridanus....................... 324,430

sculpturatus.. ............... 318,323,430, pi. 63

petiti, Terebra..-.........--...-...-..--....- 405

phengoides,AgladriUia (Agladrillia)............ 320,

386-387, pis. 59,64 

Philbertia (NannodieUa) fraternalis............ 395

Philine corrugata............................. 421

PhUinidae... ................................. 420

phoinicoides, Volvula.......---.---............ 426

Volvulella (Volvulella)........ 320,324,426, pi. 62

Phosmetuloides.. ....................... 359,361,410

picta, PleuToliTia............................. 365

pilosus, Hipponix cf. H....................... 314

pilsbryi, Ficus................................ 307

Ficus cf. F................................ 307

Marginella latissima.. ..................... 332

Prunum latissima.... ...---..-.....-.--.-- 332

Pihbrytyphis.................................. 322

Pinguigemmula............................... 361

pinguis, Ancilla (Ebuma)... ............ . 323

plana, Crepidula..... ......................... 323

pZareiceps, Conws.....__............. 316,350

Conwscf. C.-................--.--- 314,316,350

planiliTatus, Conus.......................... 346,356

Pleurofusia............................. 366-367,375

acra..................... 318,367-368, pis. 57, 65

enae...................................... 367

Pleurofusia Continued

fenimorei............................ 3,25,367,368

cf. P. fenimorei..................... 3*25,367,368

fusinella.................................. 368

fusinus................................... 367

longi Tostrops is............................. 366

Olivia.. ................................... 367

servata.................................. 367,370

sp............................... 306, 367, pi. 48

Pleurofusiaf sp....... ...................... 314,367

Pleurofusia (Cruziturricula)................... 322

arcuata... ............................ 368

CTuziana.. .......................... 368,370

fusinus fusinus....................... 318,

324,368, 369-370, pis. 57, 65 

sanctidavidis.. .................. 368,369

ffJj/pto.............................. 368,370

Za«ra....____.___.____.__.____._____ 368,370

panthea... ............................ 368

tficfcs6wr0ensis.___................... 368,370

PZewroZirio.................................... 363

albida.... ................................. 365

albidoides................................. 363

ared ersoni................................. 365

6arrettt..._.._.___._._._.___._.___...._.._. 365

haitensis.......... ........................ 365

ozjrtropis___............................ 365

pJcta..................................... 365

sp........................................ 365

(Polystira) ecuadoriana..... 322 325, 365, pi. 62

tenagos .......... 318, 324, 364-866, pis. 57,65

sp.................................... 318

(Polystiraf) sp......-.--....-...--.......- 314

Pleurotoma abundans......................... 378

acula..................................... 364

acutirostra..... ........................... 360

albida.. ................................ 363,364

tellea... .............................. 364

sp. afl. P. alesidota macilenta... .......... 378,379

arcuata... ............._......__..... 369

cedo-nulli................................. 371

chttdreni.................................. 360

cochleaTis................................. 363

cottaris...... ..........................---- 367

consors.-.-..--.-...---..--.-.--.---.----- 378

CTistata.. ............................... 370,371

dam..... ................................. 378

gracilenta................................. 396

ftattensis.................................. 364

longiTOStTOpsis........................... 366,367

lunata.. ................................ 374

lymneiformis....-..---....--..-...--...--- 402

meyeri....................... ............. 389

miranda............................... 371,372

pontonensis...--..-...-...---....--.-...-- 361

servata.................................... 366

supramirifica.......... ....-..---..-...--- 363

tigrina-................................... 364

vicksburgensis.......... ....--.-..----.---- 370

(Andstrosyrinx) aff. radiata. .........-..-- 372

(DTillia) alesidota magne................... 379

dalli................................ 378,379

ffatonensis...--.--..---.-------------- 381

gatunensis n. var.................... 381,382

homnda.............................. 378

(Gemmula) vaningeni.........-.--......-- 361

(Genota) gertTudis......................... 396

(Glyphostoma) mexicana................... 399

(Mangelia) tiara........................... 390

(Mangilia) lissotropis.................... 388,389

(Surcula) fuegiensis....................... 387

plicata, Oliva (StrephoneUa).. ............... a24,434

Pliolepidina.................................. 304

PoZinices 6rwnnewss«6cZaMSw».......-----....- 323

stanislasmeunieri.......................... 323

PoKntcesPsp................................ 310,314

polynematicus, Murex.......... ............. 311,316

(Murexf) cf. M.................. 310,311,316

(Siratus) cf. M.............. 310,311,314,316

Page 

._.__.._._..-.__...._._________. 360,363

oxytropis ecuadoriana.... ............_... 365

ttfrffo..--....-...-.--...-..._............ 364

(Pleuroliria) tenagos...................... 364

pontonensis, Pleurotoma..... .............._. 361

Porites........................................ 312

PoTtOTicia.....-.-............................. 431

portoricoensis, Drillia censors...__....._... 378

PotamidessuprasuJcatu*............ 310,311,313,314

praeambiguus, Nassarius (Uzitaf). .......... 314,316

praecellens, Turritella (Bactrospira) altiUra...- 323 

praeconsors, Crassispira...... ......._....... 379

praelubrica, Strombiformis.....--..--......--.. 326

precursor, Prunum............................ 332

Tricofta.._________________________________ 305

pnmws, Scaphander............ ............_ 420

procerum, Terebellum (TerebeUum)............ 303

procerwm?, Terebellum............... ......... 304

proJnwm, Pachycrommiumf.................... 305

propeobesa, Persicula________._....... 334

propevenusta, Cancettaria..................... 343

prosulcata, Sulcularia......................... 424

protews, Conus..............__....._.__ 848

aemulator, Conus......................... 361

Protocardia____......._...._..._.__ 317

Prunum...................................... 331

apalachee.......... .....__......___. 316

coniforme...... ........................... 332

coniformis................................. 332

latissima pilsbryi.......................... 332

latissimum................................ 332

mindiense................................. 332

precursor....._...................._ 332

sforeriwm.-.--.-..----.-..-...---.---.--.. 332

(Microspira) aff. P. opoZocftee. 314,316,332, pi. 48 

gatunense............... 318, 323, 332, pi. 51

sp.................................... 314

pseudoleroyi, Scalina.......................... 323

PstZorfws................................... 322,4*4

leptus..................................... 323

pterinus, Typhis.............................. 434

Pteropwrpwra.--....--.-..-..-....--.----.-..- 433

rhyssa.................................... 433

vokesae.................................... 433

Pterynotus.................................... 433

latUabris. ................................. 434

pellucidus____......._........--....- 433

tripterus.................................. 433

(Subpterynotus) textilis... 318,323,433-434, pi. 63 

Ptychosyrinx...--..----.-.----.-------.-----.- 361

puertocolombiana, Clathrodrittia............. 382,383

Drittia.................................... 381

pwertoricensis, TypAis......................... 434

pugnax, Orthaulax.. ........................ 306,307

Orthaulaxct. O. .................... 306,307,432

punctata, Tornatella.......................... 415

punctatus,Acteon.......................... ... 415

punctostriatus,Actaeon. ..-...-.---.....-...-- 415

Acteon(Acteon}............ 320, 324, 415, pi. 62

puncto-striata, Tornatetta..................... 415

pupiforme, Strioterebrum...................... 413

Purpura cossmanni..............------------- 389

pustulosa tobleri, Lepidocyclina... ............. 304

pycnum, Teinostoma (Pseudorotetta)........... 323

pycta, Cancellaria............................. 338

pygmaeus, Conus................ 347

Pyramidellldae............................. 414-415

pyrgoma, Crassispira (Hindsiclava).. 325, 380, pi. 62 

pyrgota, Glyphostoma (Glyphostoma)........... 320,

400-401, pi. 60 

pyriformis, Conus........................... 346,347

Pyruclia... ......................... 334,338,342,344

Q 

quercinensis, Retusa.......................... 424

biforis, Retusa (Cylichnina)................ 320,

324,423-424, pi. 62 

quirosana, Strombina.......................... 316

Strombina cf. S......................... 314,316
450 INDEX

R Page 

Rabicea... .................................... 883

radiata, Cochlespira. . ....................... 371,372

Pleurotoma (Ancistrosyrinx) aff........... 372

radula, Terebra ............................... 410

raptum, Strioterebrum. ........................ 414

Strioterebrum aff. S.......... 320, 408, 414, pi. 61

reclusa, Semicassis ( Tylocassis) ................ 323

redusiana xena, Neverita (Glossaulax).. ....... 322

recognitus, Conus.. ........... 318,324,846-847, pi. 55

recta, Volvulella. .............................. 426

rectaxis, Drittia macilenta ................... 378, 379

rectiuscula, Eulima. .......................... 327

recurvirostris, Murex (Murex) ................. 322

recurvirostris, Murex (Murex) ............. 323

Typhis... ................................ 305

(Laevityphis) aff. r......... .......... 305

reevi, Ancistrosyrinx cedonutti.. ............... 371

References cited. . .......................... 435-440

reticulata, CanceUaria.. ....................... 335

Betusa. .................................... 417,423

adamsi... .............................. 305,423

anthera. .................................. 424

biforis...... .............................. 423

decapitata ................................ 424

quercinensis .............................. 424

sulcata fossilis . ........................... 424

venillii... ................................ 424

J?e£wsa?sp_ ............ _ ................... 423

Retusa (Cylichnina) aff.R. adamsi...... 304,305,^3

quercinensis biforis.... 320, 324, 428-484, pl.62

Retusa? (Cylichnina?) sp._ ................... 316

Retusldae.. .................................. 423

rez, Xancus cf. A'_ ..................... 310,311,316

rhadina, Atys (Roxaniella) . . . . 305, 306, 4##, pi. 48

rhicna, Architectonica (Architectonica) .......... 307

Rhiglyphostoma. ................ 322, 399, 400, 40/ -40?

Rhinoclavis (Ochetoclava) ...................... 322

Rhipophos. ................................... 322

Rhizorus.. .......... ......................... 424

adelaidis. ................................. 424

rhyssa, Pteropurpura ......................... 433

rhyssus, Murex. .............................. 433

Rictaxis. ................................... 322,4/5

myakkanus ............................... 416

oryza.. ............................. 320,324,4/5

Ringicula.. ................................... 4/6

hypograpta . . . ............................ 417

(Ringiculella) semistriata.320, 324, 418-417, pi. 62 

tridentata. .......................... 416,417

(Ringiculella?) sp....................... 306,4/5

Ringiculella.... .............................. 4/5

Ringiculidae...... ........................... 416

rintriada, Glyphostoma amicta ................. 395

Nannodiella.. .......... 320, 324, 395, pis. 60, 64

riogurabonis, Drillla ... ........................ 375

.DnKm afl. D.............................. 375

eurysoma, Drittia (Neodrillia) ............. 318,

324, 374-375, pis. 58, 64 

Neodrillia. . .............................. 374

riomaensis, Acteon ............................ 415

riosantiagensis, Conus... .................... 348, 349

Rissoina (Zebinella?) sp. ..................... 314

ritanida, Crassispira.. ........................ 381

robusta, Eulima ... ............................ 328

Terebra... ....... ......................... 408

rowelli, CanceUaria... ....................... 335,338

chipolana.. ............ 320, 324, 419-420, pi. 62

Roxaniella .................................... 488

runchaena, CanceUaria... ..................... 343

rasa, ^4c<eocma. ........... 320, 324, 4/8-4/9, pi. 62

saavedrai, Clathrodrillia...... 318, 383, 384, pis. 59, 65

Clathrodrillia aS. C.......... 320, 383-384, Pi- 58

Saccharoturris............................... 360,394

samanensis, Turritdla......................... 303

Turritella cf. T............... ......... 303

Page 

sanctiandreae, Drittia vaningeni................ 362

sanctidavidis, Pleurofusia (Cruziturricula)

fusinus......................... 368,369

Turris albida.............................. 369

sarahberlinerae, BuUa......................... 422

sarissiformis, Eulima............ 318, 323, 327, pi. 49

Strombiformis............................. 327

sauridens, Conus........................... 304,349

Corawscf. C......................... 303,304,349

scalatetta, Trigonostoma....................... 345

Trigonostoma cf. 71.........-..-..---.-- 318,345

scaliae, Conus................................. 353

Scalina pseudoleroyi.......................... 323

weigandi.................................. 825

Scaphander................................... 480

jacksonensis............................ 305,420

pn'm«8.................-...-.....-.-.- . 420

(Scaphander) cryus...... 305, 306, 316, 4#0, pi. 48

cryusl................................ 316

cf. S. jacksonensis...... 304, 305, #0, pi. 48

Scaphandrldae............................... 417

scheibei, Cancellaria........................... 339

scipio, Epitonium............................. 325

Scobinella.... .................. 307,322,325,363,372

ecuadoriana...................... 325,373, pl.62

magnifica. .--.......-..--...-.--..----.--- 373

morierei............. 307,318,324,372-373, pi. 58

aff. S.morierei...................... 306,307,373

morierigavilinensis..................... 372,373

omola...................................... 373

Sconsia....................................... 383

laevigata sublaevigata...........--...---... 323

seott/, Bittium.............................. 313,314

Eulima................................... 327

Strombiformis............................. 327

sculpturatus, Petaloconchus...... 318,323,430, pi. 63

smew, Persicula............................ 304,333

(Oibberula) cf. P......... 303,304,333, pi. 48

Semicassis aldrichi............................ 311

intermedia................................ 307

(Echinophoria) apenes.................... 307

sp.................................. 307,314

Semicassis? (Tylocassis?) cf. S. aldrichi...... 310,311

Semicassis (Tylocassis) reclusa........-------- 323

sp...................-.... .-. ... 314

s, Conus symmetrkus............ 353,354

«, Echinolampas...................... 317

semistriata, Ringicula (Ringiculella) .......... 320,

324,4/5-4/7, pi. 62 

Seraphs...................................... 304

Serpulorbispapulosus...----------............ 323

sp........................................ 314

Serrata....................................... 330

serrata, Marginella............................ 330

servata. Pleurofusia......................... 367,370

Pleurotoma............................... 366

airrcafa, Cancellaria.......................... 337,340

Sinum euryhedra.............................. 323

sp...................................... 310,314

Siphocypraea (Muracypraea) henekeni......... 323

Siphogenerina................................ 313

sirena, Glyphostoma........................... 399

Solariella altiuscula........................... 306

depressa.................................. 306

ephnidia.................................. 306

Solariorbis (HapaZcrft/sl..--------------------- 322

solenaeum, Pachycrommiumf-................. 303

Solenosteira................................... 322

dallidalli................................. 323

medioamericana....................... 323

subsp................................. 325

soHda, Bulla..----........................... 422

Cancellaria............................... 338

solidus, Conus.............................. 346,347

sophus, Conus................................ 355

soverbii, Colubraria............................ 432

spez, Strioterebrum............................ 326

spenceri, Amputtinopsis... -------------------- 307

spiekeri, Conusmultiliratus................... 357

Page 

spirata, Strepsidura........................... 435

Trochita.................................. 307

spirata?, Trochita........................... 322,323

spirata, Trochita cf. 71........................ 307

Spiratella................................... 426-42?

inflata ekvata................ 320, 324, 4#7, pi. 66

Spiratellidae-..---.---.---.-.-.---........-.- 426

spirifera, Terebra........................... 409,413

midiensis, Terebra......................... 413

Terebra (Acus) bipartita................ 409, 410

(Strioterebrum)........................ 409

spiriferum, Strioterebrum---....-----.-.-....-- 320,

324, 408, 409-411, pis. 62, 66 

Splendrillia................................... 389

springvaleense, Crucibulum (Disp^taea)........ 323

springvalensis, Conus......................... 355

spurius, Conus......... 318, 324, 346, 34S, 350, pi. 55

atlanticus, Conus......................... 348

group, CcraMS----.---..................... 347

stagnalis, Ettobium............................ 429

stanislasmeunieri, Polinices....---------------- 323

steerei, Hemisinus............................. 431

-SteHozfg.___.__._.....____._.__...___..__.__.. 305

Sthenorytis pernobilis?......................... 325

toroensis euthynta......................... 325

toroensis.............................. 325

stenotella, Kurtziella (Kurtziella).............. 320,

393-394, 397, pis. 60, 64 

stenygra, Persicula (Rabicea) couviana......... 318,

333-334, Pi- 51 

stewarti, Averellia (Lecallia)................... 304,

305, 429-4W, pi. 48 

stibara, Cylichnella atacata....... 320, 324, 4/9, pi. 62

stibarus, Conus. .............................. 359

Stigmaulaxguppiana....................... 323, 325

stimpsoni, Conus.............................. 346

stonemanae, Heliacus (Astronacus)............ 323

storerium, Prunum........................... 332

Slrephonella.................................. 434

Strepsidura................................. 305, 435

multistriata............................... 435

spirata.................................... 435

striata.................................... 435

Strepsiduropsis............................... 435

rfrfoia, BuBa.--..---- --.---- --. --- 422

Glyphostoma.............................. 305

Strepsidura............................... 435

striatula,Niso--.............................. 329

Strioterebrum................................. 408

armillatum...................-..-----.-.-- 414

asperum...................--.-----....... 409

clethra.................................. 316,409

cf. -S. etoftra---------------------- 316,409, pi. 50

coZm'.-- --.--. . -.-- ----- - 413

colombianum.............................. 410

dislocatum.............................. 410,413

gatunense.......................-----...-- 409

cf. gatunense...............----.-.--....-- 409

gatunense kugleri.... ---------------------- 413

gausapatum....... 320,324,408, 411-412, 413, pi. 62

guanabanum.............................. 410

indocayapum..... 320,324,408, 418-418, pis. 62,66

ischnum................................ 306,408

laevifasciolum. -------------------------- 409,413

listrotum............. 305,306,307,408,409,pi. 48

monidum................. 320,324,408,4/4,pi- 63

nelsoni.................................... 414

oresignum..... .........-...-..-.....-..--- 413

hadrum............... 320,408,4/3-4/4,pi- 62

oresignum............ 320,324,408,4/3, pi. 62

pupiforme................................ 413

rajrfwTO---------...-.-..--.--------------- 414

afl. S.raptUTB-- - ------ 320,408,4?4, pi- 61

spd__............................. -. 326

spiriferum........ 320,324,408,409-4//,pls. 62,66

ZeZe mbiense................................ 409

waltonense.............................. 409,411

wolfgangi......... 320,324,408,4//,412,pis. 62,66

sp...-..-...-------- 306,307,310,320,325,408,409
INDEX 451

Page 

Strombidae_ ................................ 432

Strombiformisacuta-.......................... 327

ischna.................................... 326

ischnon.--------.....-.......-- ..-.-.... 327

praelubrica..... ........................... 326

sarissiformis....-....-......... -........ 327

scotti. ---- ---..-.... --.-.... 327

Strombina quirosana,......................... 316

cf. S. quirosana-.-...................... 314,316

(Sincola)---................--..-....... 322

chiriquiensis.......................... 323

(Strombina) lessepsiana................... 323

StrombineUa .............................. 322

Strombinophos............................ 322

Strombus gatunensis.... ....................... 323

sp.------------_-..__._.._.._.__.-.. 310,314,316

subbullata, Acteocina-.-................--.-. 417,418

costaricana, Acteocina..................... 417

subdausus, Polinices brunneus................ 323

subelegans, Cypraedia......................... 303

Cypraedia aft. C.......................... 303

subgrundifera, Turritella.................... 313,314

Turritella cl. T........................... 316

sublaevigata, Sconsia laevigata_.---.....--... 323

subperpolita, Drillia........................... 375

subsemicostatus, Cymatophos.................. 323

subsulcifera cembra, Terebra (Oreoterebra)...... 320,

324,405, pi. 61 

subsulcifera, Terebra (Oreoterebra)......... 320,

324, 404-400. pi. 61 

Subterynotus- .-. ...............-....-.... 322,433

subvaricosa, Clathrodrillia..................... 383

sulcatafossilis, Retusa......................... 424

Sulcoretusa............................... 424

Ziporo, Sulcoretusa........... 320, 324,424, pi. 62

harveyensis, Sulcularia.................... 424

sulcifera, Terebra.............................. 406

Sulcoretusa................................... 424

paziana................................... 424

sulcata................_..._............... 424

lipara... ................ 320, 324, 424, pi 62

Sulcularia lipara.............................. 424

prosulcata................................ 424

sulcata harveyensis..--.......--....-----.. 424

sulculorum, Atys.............................. 423

sulculus, Conus-.--.................... 306,307,350

COJMW cf. C................. 305,306,307, 349-350

supramirifica, Pleurotoma..................... 363

suprasulcatus, Potamides............ 310,311,313,314

Surculahobsoni.-.--_._..................... 378,379

Symbols used for relative frequency -...--.... 304

symmetricus, Conus......... 318, 324, 353-354, P1 - 57

domingensis, Conus..................... 353,354

semiobsoletus, Conus.................... 353,354

Syntomodrttlia.............................. 388-889

lissotropis................................. 389

woodringi................................. 389

taeniolata, Volvarina........ ........ ....... 331

tainoa, Carinodrillia (Buchema) --------------- 375

TalUypMs.................................... 434

tampae, Acteon.............................. 306,415

(Acteon) aS.A.................. 305,306,4?5

tapeina, Cancellaria (Cancellaria)-.....- -..- 318,

335,339, pi. 51 

taphrus, Conus.....----------------- 318, 3,54, Pi- 57

tapina, Nevertta (CHossaulax) bolivarensis...... 305

tarri, Mangttia................................ 392

taurina, Terebra (Paraterebra) aff. T.......... 320,

408-407, pi. 61 

taurinensis, Gamopleura....................... 428

Hyalaea.................................. 428

taurinum, Buccinum- _______ . ___ 406

taurus, Latirus.... . __ - - _ 325

Teinostoma (Aepystoma)................. .... 322

(Pseudorotella) pycnum................... 323

telembiense, Strioterebrum..................... 409

tellea, Pleurotoma albida....................... 364

Page 

tenagos, Pleuroliria (Polystira) _______ ____ 318,

324,364-^65, pis. 57,65 

Polystira (Pleuroliria).... .__-__ -____ 364

Terebettum ................................. 304

belemnitum............................... 303

procerumf. ___- - _ - --- 304

(Seraphs) belemnitum'!-................... 303

( Terebettum) procerum. ................... 303

Terebra-................................... 322,403

acuaria................................. 411,412

bipartita.................................. 404

cratera.................................... 408

dislocata .__--__ ---- -- __ - __ 409

dumbauldi...-....... --.-............... 408

flammea.................................. 406

floridana.................................. 404

gabbi...................................... 408

gausapata...--....-.-....---......--.--..- 411

herviderana........................... 412

haitensis... --.-..........-.....--....... 404

inaequalis................................. 406

ZorowL -- - 408

loroisii.................................... 408

odopoia................................. 316,404

petiti..................................... 405

radula............-.-.....-...-.-..-.-.... 410

ro6M*ta.---------------------------------- 408

apm/era.-----.- -- ------ 409,413

midiensis............................. 413

sulcifera-................................. 406

toawa..____..___.._-.--_-_-__.---------__ 406

wolfgangi..--.........--....-..-........ 410,411

sp. ind--_-------------------------------- 404

(4cus) bipartita bipartita.................. 411

spirifera........................ 409,410

(Oreoterebra) dicheres......... 313,316,404, pi. 50

isaacpetiti. _-.. 320,324, 405-406, pi. 61

subsulcifera cembra...--.. 320,324,405, pi. 61

subsuldfera....... 320,324,404-405, pi. 61

(Oxymeria) gatunensis................... 409,410

(Panaterebra)-.------------............... 322

cucurrupiensis........ 320,324, 407-408, pi. 61

(Paraterebra) cembra.....----------------- 405

cf. T. cMcarrMpiewsia. - -------- 407

aff. T. taurina............ 320, 408-407, pi. 61

sp--_-_------------------------------ 310

(Strioterebrum) gatunensis kugleri........ 409,410

gausapata.... .........--....---...-.-- 411

herviderana............................ 412

monida.... -------------------------- 414

pavonia - .-..........-...-----.------ 412

spirifera............................. 409

sp.A.. . -- -- ------ 412

sp. b....... 412

Terebralia dentilabris---...............----..-- 311

dentilabrist..........-.... . ....- 310,311

Terebridae--. . - -- -- 403

terminula, "Cythara" (Brachycytharaf) cf. 391

costaricensis, Cythara......---------------- 390

Lepicythara.------------------------------ 391

ierrj/i, Cancellaria (Charcolleria)............... 318,

323, S4S-S44, pi. 54 

OZWZa (Niteoliva)....................... 323

tersa, Microdrillia............................. 390

tessellata, Aphera...... ----------------------- 344

Cancellaria........................... .... - 344

testacea mancinella, Agaronia...-.....___._-.-- 323

testaceus, Epidromus....... ------------------- 432

Triton-.--...----. ---------------- 432

texana, Terebra............-...........--..-.-. 406

texasiana, Volvulella (Paravolvulella) ---------- 426

textilis, Murex (Pteronotus).... ---------------- 433

Murexiella (Subpterynotus)................ 433

Pterynotus (Subpterynotus).... . -- -- . 318,

323,4SS-4S4, pi. 63 

Thais (Vascula).....--------------.------- 322

Thiaridae----- - ----------------- 430

Thiarinae_.....--------.------.--- ------- 430

thompsoni, Eupleura...........--------------- 323

Page 

Tiara......................................... 322

tiara, Pleurotoma (Mangelia).................. 390

tigrina, Pleurotoma.-.......................... 364

tobleri, Lepidocyclina pustulosa ---------- - 304

Tornatella bullata....______ _--...... . . 417

punctata...........................------- 415

puncto-striata...-..........-.....--...-.-. 415

(Actaeon) cubensis........................ 415

Tornatina bullata- ..__-..----.-.------------ 417

tornatus, Conus............................... 358

Toro limestone member of Chagres sandstone-.-..--.------------------------- 322,325

toroense, Trigonostoma....... ....-------------. 345

toroensis, Conus...--....-.---- -------------- 353

euthynta, Sthenorytis, . ---------------- 325

toroensis, Sthenoytis.....---.------------ 325

tortuosopunctatus, Conus............ 346,357,358,359

tortuosostriatus, Conus.....-------------------- 318

322,324,325,352,359, pi. 57 

tournoueri, Oostrombus..---------------------- 303

towensendi, Eulima.-.-...-...-......-.--..--- 327

trachyostraca, Cancellaria dariena.------------- 336

triaspis, Cavolina (Cavolina).. ...-. 320, 428, pi. 63 

Trichodiscina- -.--.-..............-.---------- 429

Trichodiscus....----........---..------------- 429

Tricolia calypta..-.............---.----------- 305

precursor-...--.....---------------------- 305

tridentata, Ringicula (Ringiculella).--------- 416,417

Trigona pellucida.................-.---------- 345

Trigonostoma...-......--...-.-.-.---.-. 334, 344-84$

funiculatum..... ..... _..-..------------- 345

hoerlei---.------- ------ ---------------- 334

insulare............,..-.-.-.-------------- 345

cf. T.insulare.--...._--..-.--- ------- 318,345

miUeri.. ..-.-. - --- ~ 334 

scalatella.... ------------------------------ 345

cf. T. scalatella....-------------------- 318,345

toroense... . _____-_. ----------------- 345

woodringi- ---...-.....-------------------- 345

n. sp ------------------------ 322,345, pi. 63

trina, Microdrillia .... 320,324,374,389-390, pis. 59, 64

trinitatensis, Crassispira consors......... -.- 379

Drillia consors.....................------- 378

Pachycrommiumf. ---------------------- 308,313

Pachycrommium? aff. P.?----------------- 308

Pachycrommiuml cf. P.?. --------------- 313,314

trinitatis, Cavolina audeninoi. ----------------- 428

tripterus, Pterynotus............--- .- ---- 433

triticumtritonis, Cylichnella.- ------------------ 419

Triton obscurus.-.-.-------------------------- 432

testaceus- ---------------------- ---- 432

trochiformis, Trochita.. ------------------- 322,324

Trochitacf. T....------------------ 307,316

Troc/iito-.------------------------------------ 322

spirata ...... ...-. ---- 307

spiratal ..... - - 322,323

cf. T. spirata....................- -- - 307

trochiformis---..-----.---.-------------- 322,324

cf. T. trochiformis.-.- --------------- 307,316

sp -- 314

truncatus, Hemisinus_... ._---------------- 431

turbinopsis, Conus.-.--.--------------------- 357

Turbonilla bartschiana. ----------------------- 414

gatunensis.- ---------------------------- 414-415

"Turricula" arcuata..----------------------- 369

Turricula lavinia... --------------------------- 369

"Turricula" lavinia.-------------------------- 369

Turricula lavinoides.-------------------------- 369

(Surcula) lavinia-.. --------------------- 369

panthea_---------------------------- 369

Turriculinae ----------------------------- 360,366

Turriculine? turrid. ------------------- 304

Turridae.------------------------------------ 360

Turrinae------------------------------------- 36°

Tunis albida ----------------------- 363,364

sanctidavidis-------------------------- 369

brassoensis.------------------------------- 362

limachapoorensis -------------- 362
452 INDEX

Tunis Continued

(Bela) dominicensis....................... 394

(Crassispira) consors...................... 378

(Drillia) militaris.. ....................... 378

(Qemmula) vaningeni..................... 361

(Surcula) fusinella........................ 367

parkeri............................... 374

turrita, Brachycytha.ro.......................... 391

Turritella..................................... 430

abrupta.... ................. 318, 323, 430, pi. 63

adela..................................... 305

cf. T.attiHra ----------------------- 306

berjadinensis cocoditana................... 313

cf. T. berjadinensis cocoditana........... 313,314

caleta..................................... 305

cf. T. caleta............................... 305

caparonis............................... 311,313

carinata................................... 303

cf. T. carinata...........____......... 303

collazica.................................. 313

cf. T. collazica.................. 313,314,316,431

galvesia....... .........__.............. 305

gatunensisl...................... ......... 317

gatunensis gatunensis..................... 323

listrota.................................... 306

matarucana-..-.-.-....................... 323

meroensis............--...-..--.-..-...... 307

olttoni.................................... 305

samanensis............................... 303

cf. T. samanensis......................... 303

subgrundifera.--. ...................... 313,314

cf. T. subgrundifera............ ........... 316

venezuelana................. 306,310,311,313,314

(Bactrospira) altUiraaltilira......... 317,323,325

praecellens........................ 323

subsp............... 306,307,313,314,316

(Bactrospira?) amaras....... 310,311,313,314,316

(Torcula) altilira altilira .................. 317

subsp...- .................. 307,316

(Torculal) amaras.................. 310,311,316

Turritellldae................................. 430

tyloessa, Crassispira (Crassispirella).. 318, S81, pi. 58 

Typhinae........ ......................... 434

Typhis....................................... 434

alatus..................................... 434

pterinus.................................. 434

puertoricensis............................. 434

recurvirostris.............................. 305

(Laevityphis) aff. T. recurvirostris......... 305

(Talityphis) alatus obesus................. 323

eucteanus.................... 318,484, pi. 63

typhon, Glyphostoma.......................... 402

U

uaquala, Cancellariaagria..................... 342

ultimus, Conus consobrinus..... -... - 352,353

umbilicata, small form, Bulla. 320,324, 481-488, pi. 62 

waltonensis Bulla......................... 422

Niso...................................... 305

umbilicata?, Niso (Niso)......... 304,305,320, pi. 48

undulata, Vagihella................. 320,324, 487-488

undulatum, Balantium........................ 427

Utriculastra ................................ 417

Uzia miocaenica.............................. 336

Vaginella................ ..................... 48?

caribbeana....... ..........__.....__. 427

Vaginella Continued

chipolana........ ....................... 307,427

depressa ................................ 307,427

lophota......... ............. 306,307,427, pi. 49

undulata....................... 320,324,427-428

validus, Xancus............................. 311,316

Xancusct. X................... 310,311,314,316

falconensis, Xancus....................... 324

vaningeni machapoorensis, Drillia... .. _ 362 

sanctiandreae, Drillia..................... 362

Oemmula.... 316,318,324, 861-368, 363, pis. 57, 65

OemmulacL G..................... 314,316,360

Pleurotoma (Qemmula)................... 361

machapoorensis, Tunis................... 362

Tunis (Oemmula)........................ 361

vaughani, Campanile.......................... 431

Lepidocyclina............................. 309

veatcfii, Conus................................. 351

Cymatophos. ........................... 311,316

Cymatophos f cf. C... ........... 310,311,314,316

veatchi, Cymatophos....................... 323

Velates.... .................................... 304

perversus................................. 303

subsp.?............................... 303

vendryesiana, Clathurella...................... 396

Euclathwella (Euclathuretta)............. 320,

324, 396, pi. 60

venezuelana, Cancellaria ...................... 340

Persicula................................. 333

lavelana, Persicula........................ 333

venezuelana, Persicula............... 311,316,333

amydra, Persicula (Rabicea).............. 310,

311,314,316,333, pi. 48 

Turritella................... 306,310,311,313,314

ventricosa, Cavolina........................... 429

(Cavolina) cf. C.............. 320, 489, pi. 63

venusta, Cancellaria........................... 343

Clathrodrittia.... ......................... 383

Drillia........ ............................ 381

Vermetid?.................................... 317

Vermetidae..-..__.__.-_.................... 430

vernllii,Retusa.......... ..................... 424

versivestita, Asperdapfine. .........-...-...-.-- 403

vicksburgensis, Pleurofusia (Cruziturricula).. 368,370 

Pleurotoma.. ............................. 370

virgo, Murex.................................. 363

Polystira.... .............................. 364

vokesae, Pteropurpura......................... 433

Volutaalfaroi............. .................... 325

eurytera.......-.-.....---.---. . 324,325

eurytera...... .............. .... .- - 323,325

gattata.................................... 331

Volsartna................................... SSO-SS1

avena..................................... 331

collina.................................... 331

leander........................... 318, SS1, pi. 51

taeniolata................................. 331

Volvula cercadensis............................ 425

cylindrica.-----.---------.---.-.---------- 425

oxytata.................................... 425

parallela........................--.--.---- 426

persimilis................................. 425

phoinicoides............................... 426

Volvuletta................................... 4*4~4*s

conradiana.............................. 305,425

minuta................................... 426

persimilis................................. 425

recta.. _____.-...__- _-.---.----- -- - 426

Page 

Volvulettal sp................................. 425

Volvuletta (Paravolvulella) texasiana........... 426

(Volvuletta) aff. V. conradiana....... 304,305,425

cylindrica parallela.... 320,322,324,420, pi. 62

micratracta.................. 320,324, 485-486

oxytata................... 320,324,425,pi. 62

phoinicoides.............. 320,324, 486, pi. 62

W

walli, Conus multiliratus...................... 357

waltonense, Strioterebrum.................... 409,411

waltonensis, Bulla umbilicata.. . ............. 422

waylandvaughani, Lepidocyclina............... 309

weigandi, Scalina............................. 325

Weinkauffia................................... 423

werenfelsi, Cancellaria......................... 340

wetheritti, Acteon.............................. 417

whitfieldi, Dirocerithium....................... 303

wigginsi,Aphera............. .......__.._ 344

williamgabbi, Conus........................... 347

winchesterae, Carinodrillia..................... 376

wolfgangi, Strioterebrum. ----- _- .---- -- 320,

3 24,408, 411, 412, pis. 62,66 

Terebra................................. 410,411

woodringi, Glyphostoma...... ................. 400

Lioglyphostoma..........................- 400

subsp., Lioglyphostoma.................... 402

Syntomodrillia......................-~... 389

Trigonostoma............................. 345

Xancus........................................ 383

falconensis................................ 323

peruvianus..... ........................... 304

cf. A", peruvianus.......... ................ 304

cf. X. rex........................... 310,311,316

validus.... _____._ ...__._- .- _-- __ 311,316

cf. X. validus.................... 310,311,314,316

validus falconensis......................... 324

sp..._.__ __..__.._-_._-_-_- _ -. ._ 316 

Xanthonycinae__.._- _- -- -- -- _. - 429 

xena, Neverita (Glossaulax) reclusiana......... 322

xenica, Cavolina (Paleocavolina).. 306,307,428, pi. 49 

Leucosyrinx...... ................ 318, 370, pi. 57

Xenophora delecta.... ...............__ _. 323

sp........................................ 314

Y 

Yasila........................................ 304

paytensis.................................. 304

aff. Y.paytensis....... ......---.-..-...-.- 304

yurnagunensis, Lepidocyclina........-----..--- 309

morganopsis, Lepidocyclina..........----.- 309

Zemades...................................... 366

Zemacies?............................... 305,307,866

sp. a____.----...-... -..__ 304, S66, pi. 48

sp. b............................. 306, 866, pi. 48

zooki, Carinodrillia (Carinodrillia)........-.... 318,

37S-376, pis. 58, 65

Drillia.................................... 375

(Crassispira).......................... 375

zoster, Glyphostoma............................ 402

mansfieldi, Glyphostoma................... 402
PLATES 48-66
PLATE 48

FIGURES 1, 2, 12. Scaphander (Scaphander) cf. S. jacksonensis Palmer (p. 420).

Locality 41b. Marine member of Bohio(?) formation. 

1, 2. Height 9.5 mm, diameter 5.3 mm. USNM 646086. 12. Height (incomplete) 11 mm, diameter 7.5 mm.

USNM 646087. 

3. Persicula (Gibberula) cf. P. semen (Lea) (p. 333).

Height 6.5 mm, diameter 4 mm. Locality 38. Gatuncillo formation. USNM 645853. 

4, 6. Acteocina cf. A. bullata (Kiener) (p. 417).

Locality 42d. Bohio formation. 

4. Height 4 mm, diameter 1.7 mm. USNM 646092. 

6. Height 5.3 mm, diameter 2.4 mm. USNM 646091. 

5, 10, 11. Scaphander (Scaphander) cryus Woodring, n. sp. (p. 420).

Locality 42d. Bohio formation.

5. Paratype. Height 9 mm, diameter 5 mm. USNM 646094. 

10, 11. Type. Height 13.5 mm, diameter 7.8 mm. USNM 646093.

7. Niso (Niso) umbilicata (Lea)? (p. 329).

Height (incomplete) 11.2 mm, diameter 5.5 mm. Locality 41. Marine member of Bohio(?) formation. USNM 

646090.

8. Eulima cf. E. jacksonensis Gregorio (p. 326).

Height (incomplete) 3.1 mm, diameter 1.1 mm. Locality 42. Marine member of Bohio(?) formation. USNM

646089. 

9, 14, 18. Averellia (Lecallia) stewarti Woodring, n. sp. (p. 429).

Type. Height 6 mm, diameter 15.5 mm. Locality 41b. Marine member of Bohio(?) formation. USNM 646088. 

13. ZemaciesC?) sp. a (p. 366).

Height (incomplete) 19.3 mm, diameter 8.5 mm. Locality 40d. Marine member of Bohio(?) formation. USNM

645854. 

15, 16. Marignella (Eratoidea) aff. M. mollitor Dall (p. 330).

Estimated restored height 10.5 mm, diameter 5.7 mm. Locality 42d. Bohio formation. USNM 645855. 

17. Atys (Roxaniella) rhadina Woodring, n. sp. (p. 422).

Type. Height 6.8 mm, diameter 2.6 mm. Locality 42d. Bohio formation. USNM 646095. 

19 ,20. "Gemmula" sp. (p. 363).

Height (almost complete) 20.7 mm, diameter 18.2 mm. Locality 54j. Caimito formation. USNM 645856.

21. Zemaciesl sp. b (p. 366).

Height (incomplete) 32 mm, diameter (incomplete) 11.6 mm. Locality 54n. Caimito formation USNM 645857.

22. Strioterebrum listrotum Woodring, n. sp. (p. 408).

Type. Height 10 mm, diameter 2.3 mm. Locality 42d. Bohio formation. USNM 645869.

23. Glyptostyla panamensis Dall (p. 434).

Height (incomplete) 28.7 mm, diameter 20.8 mm. Locality 40a. Marine member of Bohio(?) formation. USNM 

135199.

24. Ellobium aff. E. pellucens (Menke) (p. 429).

Height 17.5 mm, diameter 8 mm. Locality 116a. La Boca formation. USNM 646100.

25. Prunum (Microspira) aff. P. apalachee (Gardner) (p. 332).

Height 10.2 mm, diameter 5.9 mm. Locality 116a. La Boca formation. USNM 645959.

26. Persicula (Rabicea) venezuelana amydra Woodring, n. subsp. (p. 333).

Type. Height 7.6 mm, diameter 4.9 mm. Locality 116a. La Boca formation. USNM 645860.

27. Pleurofusia sp. (p. 367).

Height (incomplete) 18.7 mm, diameter 9 mm. Locality 56. Caimito formation. USNM 645858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D PLATE

MIDDLE EOCENE MOLLUSK FROM GATUNCILLO FORMATION, LATE EOCENE OR

EARLY OLIGOCENE MOLLUSKS FROM MARINE MEMBER OF BOHIO(?) FORMATION,

LATE OLIGOCENE MOLLUSKS FROM BOHIO AND CAIMITO FORMATIONS,

AND EARLY MIOCENE MOLLUSKS FROM LA BOCA FORMATION

368-278 O - 70 - 11
PLATE 49

FIGURE 1. Eulima nobilis Guppy (p. 326).

Height 5.9 mm, diameter 1.7 mm. Locality 147b. Middle part of Gatun

formation. USNM 646072. 

2. Eulima sarissiformis (Pilsbry and Johnson) (p. 327).

Height (incomplete) 11.5 mm, diameter 2.7 mm. Locality 138c. Lower part of

Gatun formation. USNM 646074. 

3, 4, 6, 9. Carolina (Paleocavolina) xenica Woodring, n. sp. (p. 428).

3, 4. Type, ventral side. Height 5.5 mm, width 6.3 mm, diameter 3.5 mm. 

Locality 54j, Caimito formation. USNM 646097.

3. Right side.

4. Exterior. 

6, 9. Paratype, dorsal side. Height 6.4 mm, estimated restored width 6.5 mm,

diameter 1.8 mm. Locality 54h. Caimito formation. USNM 646098. 

6. Exterior. 

9. Right side. 

5. Niso (Niso) mesata Woodring, n. sp. (p. 329).

Paratype. Height (not quite complete) 21.1 mm, diameter (not quite complete) 8 mm. Locality 142. Middle part of Gatun formation. USNM 646080. 

7, 12. Orthaulax gabbi Ball (p. 432).

Height (incomplete) 69 mm, maximum shoulder diameter 51 mm, minimum 

shoulder diameter 39.5 mm. Locality lOlh. La Boca formation. USGS 

646101. 

8. Balcis (Balcis) cetia Woodring, n. sp. (p. 328).

Type. Height (not quite complete) 28.3 mm, diameter 8.7 mm. Locality 138f.

Lower part of Gatun formation. USNM 646078. 

10, 11. Vaginella lophota Woodring, n. sp. (p. 427).

Type. Height (not quite complete) 5.1 mm, width 2.3 mm, diameter 1.4 mm. 

Locality 54j, Caimito formation. USNM 646096.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D PLATE 49

LATE OLIGOCENE MOLLUSKS FROM CAIMITO FORMATION, EARLY MIOCENE MOLLUSK FROM 

LA BOCA FORMATION, AND MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION
PLATE 50

FIGURES 1, 2, 5-8. Floribella aldrichi (Dall) (p. 421).

1, 2. Height (incomplete) 33.5 mm, diameter (incomplete) 31 mm.

Locality Il5a, La Boca formation. USNM 646099. 

5-8. Topotypes. McClelland farm, Chipola River, Calhoun County, 

Fla. Chipola formation.

5. 7. Height (incomplete) 61 mm, diameter (incomplete) 64 mm. 

USNM 645998.

6. 8. Height (incomplete) 43 mm, diameter (practically complete) 

51.5 mm. USNM 645999.

3. Terebra (Oreoterebra) dicheres Woodring, n. sp. (p. 404).

Type. Height (incomplete) 35 mm, diameter 11.5 mm. Locality 115a. 

La Boca formation. USNM 645870.

4. Strioterebrum cf. S. celethra (Maury) (p. 409).

Height (incomplete) 25.8 mm, diameter 9 mm. Locality 115b. La 

Boca formation. USNM 645871.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D PLATE 50

X2 '

EARLY MIOCENE MOLLUSKS FROM LA BOCA FORMATION 

AND CHIPOLA FORMATION OF FLORIDA
PLATE 51

FIGURES 1, 2. Balcis (Balcis) jacululum (Maury) (p. 327).

Height (not quite complete) 3.3 mm, diameter 1.1 mm. Locality 48b.

Middle part of Gatun formation. USNM 646075. 

3. Niso (Niso) mesata Woodring, n. sp. (p. 329).

Type. Height (incomplete) 14 mm, diameter (incomplete) 6.4 mm. Locality

138c. Lower part of Gatun formation. USNM 646079. 

4-6. Volvarina leander (Brown and Pilsbry) (p. 331).

4. Height 11.4 mm, diameter 5.2 mm. Locality 177b. Upper part of Gatun

formation, eastern area. USNM 645708. 

5. Height 9 mm, diameter 4.4 mm. Locality 155c. Middle part of Gatun

formation. USNM 645707. 

6. Height 8 mm, diameter 3.7 mm. Locality 147b. Middle part of Gatun

formation. USNM 645706. 

7. Persicula (Rabiced) couviana stenygra Woodring, n. subsp. (p. 333).

Type. Height 10 mm, diameter 6.8 mm. Locality 138c. Lower part of 

Gatun formation. USNM 645711. 

8, 9. Cancellaria (Cancellaria) tapeina Woodring, n. sp. (p. 335).

Type. Height (almost complete) 34.7 mm, diameter 20.5 mm. Locality 182a.

Upper part of Gatun formation, western area. USNM 645714. 

10, 11. Prunum (Microspira) gatunense (Brown and Pilsbry) (p. 332).

10. Height 14.5 mm, diameter 8 mm. Locality 138c. Lower part of Gatun 

formation. USNM 645710.

11. Height 17.8 mm, diameter 12 mm. Locality 139c. Middle part of 

Gatun formation. USNM 645709.

12. Balcis (Balcis) aulaca Woodring, n. sp. (p. 328).

Type. Height (not quite complete) 7.8 mm, diameter (not quite complete) 

2.2 mm. Locality 138. Lower part of Gatun formation. USNM 646077.

13. Eulima acuta Sowerby (p. 327).

Height 7.3 mm, diameter 1.2 mm. Locality 159d. Middle part of Gatun 

formation. USNM 646073.

14. Balcis (Balcis) lipara Woodring, n. sp. (p. 328).

Type. Height (not quite complete) 8.3 mm, diameter 2.5 mm. Locality 138. 

Lower part of Gatun formation. USNM 646076.

15. Campanile cf. C. herculeanus (Cooke) (p. 431).

Height (incomplete) 87.5 mm, diameter 54.5 mm. Locality Il7c. Emperador 

limestone member of La Boca formation. USNM 646102.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D PLATE 51

*!-*;.«r»c--^4-L

12

EARLY MIOCENE MOLLUSK FROM EMPERADOR LIMESTONE MEMBER OF LA BOCA FORMATION 

AND MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION
PLATE 52

FIGURES 1, 2. Cancellaria (Cancellarid) anomoia Woodring, n. sp. (p. 334).

Type. Height 19.1 mm, diameter 11.7 mm. Locality 138c. Lower part of

Gatun formation. USNM 645712. 

3, 4. Cancellaria (Cancellaria) aff. C. macneili Mansfield (p. 335).

Height 19 mm, diameter 12 mm. Locality 137. Lower part of Gatun

formation. USNM 645713. 

5, 6. Cancellaria (Cancellaria) apimela Woodring, n. sp. (p. 337).

Type. Height 22.6 mm, diameter 13 mm. Locality 182a. Upper part of

Gatun formation, western area. USNM 645720. 

7, 8. Cancellaria (Cancellaria) epistomifera lipara Woodring, n. subsp. (p. 337).

Type. Height 28 mm, diameter 17.5 mm. Locality 183. Upper part of

Gatun formation, western area. USNM 645719. 

9, 10. Cancellaria (Pyruclia) cibarcola cibarcola Anderson (p. 338).

Immature specimen showing early sculpture. Height 24 mm, diameter 15.9

mm. Locality 138c. Lower part of Gatun formation. USNM 645723. 

11-18. Cancellaria (Cancellaria) epistomifera dariena Toula (p. 335).

Lower part of Gatun formation. 

11, 12. Strongly turreted specimen. Height 31.5 mm, diameter 20 mm.

Locality 136a. USNM 645718. 

13, 14. Distinctly turreted specimen. Height 27.7 mm, diameter 17.6 mm.

Locality 138c. USNM 645717. 

15, 16. Slender specimen. Height 31 mm, diameter 16.5 mm. Locality I38c.

USNM 645716.

17, 18. Inflated specimen. Height 34 mm, diameter 21.5 mm. Locality 138c. 

USNM 645715.
PROFESSIONAL PAPER 306-D PLATE 52

16 17 

MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION
PLATE 53

FIGUBE 1, 2. Cancellaria acalypta Woodring, n. sp. (p. 341).

Type. Height 15.6 mm, diameter 10.2 mm. Locality 136a. Lower part of

Gatun formation. USNM 645730. 

3, 4. Cancellaria nancellaria Woodring, n. sp. (p. 341).

Type. Height 7.6 mm, diameter 4.5 mm. Locality 147b. Middle part of

Gatun formation. USNM 645731. 

5, 6. Cancellaria (Narona) barystoma Woodring, n. sp. (p. 342).

Type. Height 16.3 mm, diameter 7.8 mm. Locality 138d. Lower part of

Gatun formation. USNM 645732. 

7, 9. Cancellaria (Pyruclia) diadela Woodring, n. sp. (p. 338).

Type. Height (practically complete) 39.5 mm, diameter 33 mm. Locality

182. Upper part of Gatun formation, western area. USNM 645724. 

8, 10-12. Cancellaria (Pyruclia) cibarcola cibarcola Anderson (p. 338).

Locality 138c. Lower part of Gatun formation. 

8, 11. Height (practically complete) 35.7 mm, diameter (incomplete) 25 mm.

USNM 645722.

10, 12. Exceptionally large, high-spired specimen. Height (practically 

complete) 49.2 mm, diameter (incomplete) 30 mm. USNM 645721.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D PLATE 53

X2

10 11 ~ 12 

MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION
PLATE 54

FIGURE 1, 2. Cancellaria (Euclid) dinota Woodring, n. sp. (p. 340).

Type. Height (practically complete) 29.9 mm, diameter 18 mm. Locality

I38c. Lower part of Gatun formation. USNM 645728. 

3, 4, 7, 8, 11, 12. Cancellaria (Euclia) codazzii Anderson (p. 339).

Lower part of Gatun formation.

3, 4. Slightly angulated, slightly spinose specimen. Height (practically complete) 33 mm, diameter 20.7 mm. Locality 138c. USNM

645725.

7, 8. Moderately angulated, moderately spinose specimen. Height (practically complete) 35.6 mm, diameter 24 mm. Locality 138c. USNM

645726.

11, 12. Strongly angulated, strongly spinose specimen. Height (practically complete) 32 mm, diameter 21.5 mm. Locality 138e. USNM

645727. 

5, 6, 9, 10. Cancellaria (Charcolleria) terryi Olsson (p. 343).

5, 6. Height 42.3 mm, diameter 19 mm. Locality 138c. Lower part of

Gatun formation. USNM 645734. 

9, 10. Height (incomplete) 53.5 mm, diameter 24 mm. Locality 155.

Middle part of Gatun formation. USNM 645733.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D PLATE 54

X2

12

MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION
PLATE 55

FIGURE 1, 2. Conus imitator imitator Brown and Pilsbry (p. 354).

1. Height 45.4 mm, diameter 22 mm. Locality 177d. Upper part of Gatun 

formation, eastern area. USNM 645749.

2. Height 33 mm, diameter 16 mm. Locality I38c. Lower part of Gatun 

formation. USNM 645750.

3. Conus acolus Woodring, n. sp. (p. 349).

Type. Height 35 mm, diameter 20.8 mm. Locality 136a. Lower part of 

Gatun formation. USNM 645741.

4. Conus recognitus Guppy (p. 346).

Height (almost complete) 39 mm, diameter (body whorl broken back) 

20.8 mm. Locality 182. Upper part of Gatun formation, western area. 

USNM 645736. 

5, 6. Conus aemulator aemulator Brown and Pilsbry (p. 351).

Type. Height 22.7 mm, diameter 12.8 mm. Gatun Locks excavation. Middle 

part of Gatun formation. Acad. Nat. Sci. Phila. 1961. Photograph by 

A. A. Olsson. 

7. Conus spurius Gmelin (p. 348).

Height 45.8 mm, diameter 28.5 mm. Locality 155. Middle part of Gatun

formation. USNM 645738. Photograph by A. A. Olsson. 

8-10. Conus mohs Brown and Pilsbry (p. 350).

8. Slightly reduced. Height 160 mm, diameter 90 mm. Locality 185.Upper 

part of Gatun formation, western area. USNM 645742.

9. 10. Immature specimen. Height 54.5 mm, diameter 27.7 mm. Locality 

155b Middle part of Gatun formation, eastern area. USNM 645743.
PROFESSIONAL PAPER 306-D PLATE 55

MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION
PLATE 56

FIGURE 1, 2. Aphera islacolonis (Maury) (p. 344).

Height 16 mm, diameter 10 mm. Locality 138f. Middle part of Gatun formation. USNM 645735. 

3, 7, 9. Conus consobrinus consobrinus Sowerby (p. 352).

3, 7. Height 47.7 mm, diameter 30.3 mm. Locality 138c. Lower part of 

Gatun formation. USNM 645745.

9. Height 61.7 mm, diameter 30.3 mm. Locality 161b. Middle part of 

Gatun formation. USNM 645746.

4. 8. Conus aemulator aemulator Brown and Pilsbry (p. 351).

Height 45 mm, diameter 28 mm. Locality 155a. Middle part of Gatun 

formation. USNM 645744.

5. 6. Cancellaria (Euclia) dinota Woodring, n. sp. (p. 340).

Exceptionally large specimen. Height 34.9 mm, diameter 21 mm. Locality

138d. Lower part of Gatun formation. USNM 645729. 

10, 11. Conus bravoi Spieker (p. 348;.

Lower part of Gatun formation.

10. Height 59.4 mm, diameter 36.6 mm. Locality 138c. USNM 645739.

11. Height (incomplete) 64 mm, diameter 44 mm. Locality 137a. USNM 

645740.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D PLATE 56

11

MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION
PLATE 57

FIGURES 1, 7. Conus taphrus Woodring, n. sp. (p. 354).

Type. Height (almost complete) 28.2 mm, diameter 15 mm. Locality 161d. Middle part of Gatun formation.

USNM 645748. 

2. Conus musaensis Olsson (p. 347).

Height 20.6 mm, diameter 10 mm. Locality 170. Middle part of Gatun formation. USNM 645737. 

3, 4. Conus multiliratus muUiliratus Bose (p. 356).

Height (almost complete) 27.2 mm, diameter 14.4 mm. Locality 175. Upper part of Gatun formation, eastern

area. USNM 645751. 

5, 6. Conus tortuosostriatus Toula (p. 359).

Locality 175. Upper p?rt of Gatun formation, eastern area.

5. Slender specimen. Height (not qu^te complete) 26.3 mm, diameter 11.7 mm. USNM 645756.

6. Inflated specimen. Height (incomplete) 28.2 mm, diameter 15.7 mm. USNM 645755. 

8. Pkurofusia acra Woodring, n. sp. (p. 367).

Type. Height (practically complete) 22.4 mm, diameter 7.5 mm. Locality 155c. Middle part of Gatun formation.

USNM 645764. 

9, 10. Cochlespira (Ancistrosyrinx) cedonulli (Reeve) (p. 371).

Height (incomplete) 19.3 mm, diameter (spines broken) 10.2 mm. Locality 177b. Upper part of Gatun formation. 

USNM 645769.

11. 18, 22. Gemmula vaningeni (Brown and Pilsbry) (p. 361).

Locality 175. Upper part of Gatun formation, eastern area.

11, 18. Malformed specimen. Height (incomplete) 25.9 mm, diameter 10 mm. USNM 645758. 

22. Height (not quite complete) 38.8 mm, diameter (incomplete) 12.3 mm. USNM 645757.

12. 23, 24. Pleuroliria (Polystira) tenagos (Gardner) (p. 364).

12. Profile modified by peripheral carina. Height (incomplete) 42.3 mm, diameter 16.6 mm. Locality 138c. Lower 

part of Gatun formation. USNM 645760.

23. Profile slightly modified by peripheral carina. Height 58.8 mm, diameter 16.6 mm. Locality 173. Upper part 

of Gatua formation, eastern area. USNM 645762.

24. Profile somewhat modified by peripheral carina. Height 51.4 mm, diameter 14.4 mm. Locality 157. Middle

part of Gatun formation. USNM 645761. 

13, 14. Conus symmetricus Sowerby (p. 353).

Height 20.4 mm, diameter 11.8 mm. Locality 155a. Middle part of Gatun formation. USNM 645747. 

15. Leucosyrinx xenica Woodring, n. sp. (p. 370).

Type. Height 15.4 mm, diameter 4.9 mm. Locality 185. Upper part of Gatun formation, western area. USNM

645768. 

16, 21. Pleurofusia (Cruziturricula) fusinus fusinus (Brown and Pilsbry) (p. 369).

Locality 138c. Lower part of Gatun formation.

16. Height (not quite complete) 33.5 mm, diameter (incomplete) 9 mm. USNM 645766. 

21. Height (incomplete) 45.7 mm, diameter (incomplete) 13.5 mm. USNM 645765. 

17. Conus burckhardti harrisi Olsson (p. 358).

Height 37.7 mm, diameter 13.5 mm. Locality 138a. Lower part of Gatun formation. USNM 645754. 

19, 20. Conus burckhardti burckhardti Bose (p. 357).

Locality 155. Middle part of Gatun formation.

19. Height (practically complete) 42.3 mm, diameter 17.7 mm. USNM 645752.

20. Height (practically complete) 33.7 mm, diameter 14.8 mm. USNM 645753.
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MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION
PLATE 58

FIGURES 1, 22. Crassispira (Hindsidava) consors consors (Sowerby) (p. 378).

I. Height (not quite complete) 32 mm, diameter 9.2 mm. Locality 155a. Middle part of Gatun formation.

USNM 645779. 

22. Height (not quite complete) 42.9 mm, diameter 12 mm. Locality 175. Upper part of Gatun formation,

eastern area. USNM 645778. 

2, 3. Crassispira (Crassispirella) tyloessa Woodring, n. sp. (p. 381).

Type. Height 13.4 mm, diameter 5.2 mm. Locality 139c. Middle part of Gatun formation. USNM 645783. 

4, 5. Crassispira (Crassispirelld) cymation Woodring, n. sp. (p. 381). 

Locality 161c. Middle part of Gatun formation.

4. Paratype. Height (incomplete) 7.5 mm, diameter 4 mm. USNM 645782.

5. Type. Height (not quite complete) 8 mm, diameter (incomplete) 3.3 mm. USNM 645781. 

6, 7. Clathrodrillia aff. C. saavedrai Woodring, n. sp. (p. 383).

Height 10.4 mm, diameter 3.5 mm. Locality 161c. Middle part of Gatun formation. USNM 645791. 

8, 20, 21, 23 -25. Clathrodrillia gatunensis (Toula) (p. 381).

8. Height (almost complete) 34 mm, diameter 13.5 mm. Locality 161a. Middle part of Gatun formation. USNM

645786.

20, 21. Height (almost complete) 33.6 mm, diameter 8.8 mm. Locality 155c. Middle part of Gatun formation. 

USNM 645788.

23. Height (almost complete) 40.5 mm, diameter 12.5 mm. Locality 159b. Middle part of Gatun formation. 

USNM 645787.

24. 25. Height (almost complete) 52.4 mm, diameter 17.5 mm. Locality 138c. Lower part of Gatun formation.

USNM 645785. 

9-12. Carinodrillia (Carinodrillia) zooki (Brown and Piisbry) (p. 375).

9, 10. Height 21.2 mm, diameter 6.8 mm. Locality 138a. Lower part of Gatun formation. USNM 645774.

II, 12. Height 19.8 mm, diameter 6 mm. Locality 175. Upper part of Gatan formation, eastern area. USNM

645773. 

13, 14. Carinodrillia (Carinodrillia) cf. C. elocata (Piisbry and Johnson) (p. 376).

Height (almost complete) 26.8 mm, diameter 9.8 mm. Locality 139c. Middle part of Gatun formation. USNM

645776. 

15, 16. Clathrodrillia cf. C. islalindae (Maury) (p. 384).

Height (not quite complete) 15.8 mm, diameter 5.5 mm. Locality 138f. Lower part of Gatun formation. USNM

645793. 

17, 18. Drillia (Neodrillia) riogurabonis eurysoma Gardner (p. 374).

Height (practically complete) 11.8 mm, diameter 4.6 mm. Locality 138c. Lower part of Gatun formation. USNM

645771. 

19. Crassispira (Crassispira) henekeni leptalea Woodring, n. subsp. (p. 377).

Type. Height (practically complete) 30.6 mm, diameter 10.5 mm. Locality 147h. Middle part of Gatun formation.

USNM 645777. 

26, 27. Scobinella morierei (Cossmann) (p. 372).

Height (almost complete) 37 mm, diameter (incomplete) 12.5 mm. Locality I38a. Lower part of Gatun formation. 

USNM 645770.
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MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION
PLATE 59

FIGURE 1, 2. Agladrillia (Agladrillid) phengoides Woodring, n. sp. (p. 386).

Type. Height 6.8 mm, diameter 2.2 mm. Locality 147b. Middle part of

Gatun formation. USNM 645799. 

3, 4. Agladrillia (Agladrillid) characta Woodring, n. sp. (p. 385).

Type. Height 12.8 mm, diameter 4.3 mm. Locality 147g. Middle part of

Gatun formation. USNM 645794. 

5, 6. Clathrodrillia aff. C. lelandi (Olsson) (p. 384).

Height (incomplete) 21.8 mm, diameter 7.5 mm. Locality 136a. Lower part

of Gatun formation. USNM 645792. 

7-9. Microdrillia trina Mansfield (p. 389). 

Middle part of Gatun formation.

7. Flat-sided specimen. Height 6.3 mm, diameter 1.9 mm. Locality 147g. 

USNM 645809.

8. Subequal peripheral and lower spiral threads. Height 5.6 mm, diameter 

2 mm. Locality 147b. USNM 645807.

9. Peripheral spiral thread stronger than lower threads. Locality 147b.

USNM 645808.

10, 11, 15, 16. Agladrillia (Agladrillia) enneacyma enneacyma (Brown and Pilsbry) (p. 385). 

Locality 147b. Middle part of Gatun formation.

10. 11. Inflated specimen. Height 7.2 mm, diameter 2.8 mm. USNM 645797. 

15, 16. Slender specimen. Height 7.7 mm, diameter 2.5 mm. USNM 645796.

12. 17. Agladrillia (Agladrillia) acaria ectypha Woodring n. subsp. (p. 387).

Type. Height 6.5 mm, diameter 2.3 mm. Locality 138. Middle part of Gatun 

formation. USNM 645801.

13. 14. Clathrodrillia saavedrai Woodring, n. sp. (p. 383).

Type. Height 16.5 mm, diameter 6.2 mm. Locality 147b. Middle part of 

Gatun formation. USNM 645790.
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MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION
PLATE 60

FIGURE 1. Glyphosioma (Rhiglyphostoma) allodapum Woodring, n. sp. (p. 402).

Type. Height (incomplete) 7 mm, diameter 3.3 mm. Locality 138d. Lower part of Gatun formation. USNM 645835. 

2, 3. Euclathurella (Euclathurella) vendryesiana (Dall) (p. 396).

Type of Pleurotoma (Genota) gertrudis Toula. Gatun Locks excavation. Middle parts of Gatun formation. Techniscbe 

Hochschule, Vienna.

4. Lepicyihara heptagona (Gabb) (p. 390).

Height 11.5 mm, diameter 5.7 mm. Locality 157. Middle part of Gatun formation. USNM 645811.

5. Kuftziella (Kurtziella) pagella Woodring, n. sp. (p. 393).

Type. Height 4 mm, diameter 1.8 mm. Locality 147b. Middle part of Gatun formation. USNM 645816.

6. Kurtziella (Kurtziella) stenotella Woodring, n. sp. (p. 393).

Type. Height 4.5 mm, diameter 1.9 mm. Locality 147b. Middle part of Gatun formation. USNM 645818.

7. Glyphostoma (Euglyphostoma) olssoni Woodring, n. sp. (p. 401).

Type. Height 13.6 mm, diameter 5.7 mm. Locality 176. Upper part of Gatun formation. USNM 645834.

8. Kurtziella (Cryoturris) sp. (p. 394).

Height 2.5 mm, diameter 1.2 mm. Locality 138d. Lower part of Gatun formation. USNM 645821. 

9, 20. Dolostoma anorhepes Woodring, n. sp (p. 398).

9. Immature specimen. Height 5 mm, diameter 1.9 mm. Locality 147b. Middle part of Gatun formation. USNM

645823. 

20. Type. Height (almost complete) 15.5 mm, diameter 6 mm. Locality 138. Lower part of Gatun formation. USNM

645822. 

10. Nannodiella rintriada (Mansfield) (p. 395).

Height 4.5 mm, diameter 1.9 mm. Locality 147b. Middle part of Gatun formation. USNM 645827. 

11, 18, 19. Agladrillia (Eumetadrillia) isthmica (Brown and Pilsbry) (p. 387). 

Lower part of Gatun formation.

11. Height 13.9 mm, diameter 5.6 mm. Locality 138d. USNM 645804.

18, 19. Height 19.9 mm, diameter (incomplete) 6.8 mm. Locality 138c. USNM 645803.

12. Cytharella? cf. C. compsacosta (Gardner) (p. 392).

Height (not quite complete) 4.2 mm, diameter 1.7 mm. Locality 138c. Lower part of Gatun formation. USNM 645815.

13. Ithycythara defuniak Gardner (p. 391).

Height 4 mm, diameter 1.5 mm. Locality 147b. Middle part of Gatun formation. USNM 645812.

14. Ithycythara cf. /. elongala (-Gabb) (p. 392).

Height (not quite complete) 6.2 mm, diameter 2.2 mm. Locality 138c. Lower part of Gatun formation. USNM 645814.

15. Kurtziella (Cryoturris) habra Woodring, n. sp. (p. 394).

Type. Height 5.2 mm, diameter 1.9 mm. Locality 138a. Lower part of Gatun formation. USNM 645820. 

16, 17. Agladrillia (Eumetadrillia) acidna Woodring, n. sp. (p. 388).

Type. Height 18 mm, diameter 5.9 mm. Locality 172. Upper part of Gatun formation, eastern area. USNM 645806. 

21. Euclathurella (Miraclathurella) eucharis Woodring, n. sp. (p. 397).

Type. Height 9 mm, diameter 3.6 mm. Locality 138c. Lower part of Gatun formation. USNM 645825.

22. 25. Glyphostoma (Glyphostoma) dentiferum Gabb (p. 399).

Middle part of Gatun formation.

22. Height (incomplete) 38.2 mm, diameter 15.2 mm. Locality 155a. USNM 645830. 

25. Immature specimen. Height 32.2 mm, diameter 12.2 mm. Locality 157. USNM 645831.

23. 24. Glyphostoma (Glyphostoma) pyrgota Woodring, n. sp. (p. 400).

Type. Height (incomplete) 37.3 mm, diameter 15 mm. Locality 157. Middle part of Gatun formation. USNM 645833
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MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION

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PLATE 61

FIGURE 1, 14. Terebra (Paraterebra) aff. T. taurina (Lightfoot) (p. 406).

1. Height (incomplete) 48 mm, diameter 15.5 mm. Locality 179. Upper part of Gatun formation, western area.

USNM 645844. 

14. Finely sculptured specimen. Height (incomplete) 43 mm, diameter 16.5 mm. Locality 155. Middle part of

Gatun formation. USNM 645843. 

2. Daphnella pagera Woodring, n. sp. (p. 403).

Type. Height (almost complete) 12.3 mm, diameter 4.7 mm. Locality 173a. Upper part of Gatun formation, eastern

area. USNM 135580. 

3, 7, 8, 9. Terebra (Oreoterebrd) subsulcifera subsulcifera Brown and Pilsbry (p. 404).

Middle part of Gatun formation. 

3. Coarsely sculptured specimen. Height (incomplete) 39.8 mm, diameter 14 mm. Locality 139c. USNM 645839.

7. Height (incomplete) 51 mm, diameter 14.5 mm. Locality 155b. USNM 645838.

8. 9. Height (almost complete) 68.5 mm, diameter 12.8 mm. Locality 155. USNM 645837.

4. Dolostoma dinota Woodring. n. sp. (p. 398).

Type. Height (practically complete) 12.9 mm, diameter 5.3 mm. Locality 175. Upper part of Gatun formation, 

eastern area. USNM 645829.

5. Terebra (Oreoterebra) subsulcifera cembra Olsson (p. 405).

Height (almost complete) 48 mm, diameter 10.8 mm. Locality 136. Lower part of Gatun formation. USNM 645840.

6. Strioterebrum aff. S. raptum (Gardner) (p. 414).

Height (almost complete) 10 mm, diameter 3.5 mm. Locality 138c. Lower part of Gatun formation. USNM 646048. 

10, 11, 15, 16. Terebra (Panaterebra) cucurrupiensis Oinomikado (p. 407).

10. Height (almost complete) 66.4 mm, diameter 19.5 mm. Locality 138c. Lower part of Gatun formation. USNM 

645847.

11. Slender specimen. Height (almost complete) 65.5 mm, diameter 15.5 mm. Locality 139e. Middle part of Gatun 

formation. USNM 645848.

15. Strongly inflated specimen. Height (incomplete) 51.8 mm, diameter 21.5 mm. Locality 138f. Lower part of 

Gatun formation. USNM 645845.

16. Height (incomplete) 75.5 mm, diameter 22 mm. Locality 138c. Lower part of Gatun formation. USNM 645846. 

12, 13. Terebra (Oreoterebra) isaacpetiti Maury (p. 405).

12. Height (almost complete) 99.5 mm, diameter 17.5 mm. Locality 182. Upper part of Gatun formation, western 

area. USNM 645842.

13. Height (incomplete) 63.5 mm, diameter 14.7 mm. Locality 139c. Middle part of Gatun formation. USNM 

645841.
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MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION

16
PLATE 62

FIGURE 1, 2. Acteocina elachista Woodring, n. sp. (p. 418).

Type. Height 2.3 mm, diameter 1.2 mm. Locality 147b. Middle part of Gatun formation. USNM 646052. 

3, 4. Acteocina rusa Gardner (p. 418).

Height 2.8 mm, diameter 1.3 mm. Locality 139b. Middle part of Gatun formation. USNM 646053.

5. Sulcoretusa sulcata lipara (Woodring) (p. 424).

Height 2.7 mm, basal diameter 1.3 mm. Locality 147b. Middle part of Gatun formation. USNM 646061.

6. Retusa (Cylichnina) quercinensis biforis Pilsbry and Johnson (p. 423).

Height 2.8 mm, diameter 1.5 mm. Locality 147g. Middle part of Gatun formation. USNM 646060. 

7, 9. Acteocina bullata (Kiener), small form (p. 417).

Height 4.9 mm, diameter 2.2 mm. Locality 139b. Middle part of Gatun formation. USNM 646051. 

8, 16. Atys (Weinkauffia) cadus Woodring, n. sp. (p. 423).

Type. Height 2.1 mm, diameter 1.4 mm. Locality 139b. Middle part of Gatun formation. USNM 646058.

10. Atys (Aliculastrum) eurys Woodring, n. sp. (p. 422).

Type. Height 7.3 mm, diameter 3.6 mm. Locality 159d. Middle part of Gatun formation. USNM 646057.

11. Acteon (Acteon) punctostriatus (C. B. Adams) (p. 415).

Height 3.7 mm, diameter 2 mm. Locality 138c. Lower part of Gatun formation. USNM 646049.

12. Ringicula (Ringiculelld) semistriata d'Orbigny (p. 416).

Height 3 mm, diameter 2 mm. Locality 159d. Middle part of Gatun formation. USNM 646050.

13. Cyclichnella atacata stigara Woodring, n. subsp. (p. 419).

Type. Height 3.2 mm, diameter 1.8 mm. Locality 147b. Middle part of Gatun formation. USNM 646054.

14. Roxania chipolana (Dall) (p. 419).

Height 3.5 mm, diameter 2.2 mm. Locality 147b. Middle part of Gatun formation. USNM 646055.

15. Volvulella (Volvulella) phoinicoides (Gardner) (p. 426).

Height 3.3 mm, diameter 1.1 mm. Locality 138c. Lower part of Gatun formation. USNM 646065.

17. Volvulella (Volvulella) cylindrica parallela (Pilsbry and Johnson) (p. 426).

Height 4.7 mm, diameter 1.6 mm. Locality 159d. Middle part of Gatun formation. USNM 646084.

18. Bulla umbilicata Roding, small form'(p- 421).

Height 8.5 mm, diameter 5.2 mm. Locality 161c. Middle part of Gatun formation. USNM 646056. 

19, 20. Volvulella (Volvulella) oxytata (Bush) (p. 425).

Locality 138c. Lower part of Gatun formation.

19. Moderately slender specimen; apical part of outer lip abnormal. Height 3.6 mm, diameter 1.4 mm. USNM 646063.

20. Slender specimen. Height 3.4 mm, diameter 1.2 mm. USNM 646062.

21. 27. Crassispira (Hindsiclava) pyrgoma Woodring, n. sp. (p. 380). 

Locality 208. Chagres sandstone.

21. Paratype. Height (incomplete) 26.3 mm, diameter 9.6 mm. USNM 645866. 

27. Type. Height (incomplete) 35.5 mm, diameter 10.5 mm. USNM 645867.

22. 35. Strioterebrum oresignum hadrum Woodring, n. subsp. (p. 413).

Locality 136. Lower part of Gatun formation.

22. Paratype. Height (incomplete) 24.2 mm, diameter 8.2 mm. USNM 646046. 

35. Type. Height (incomplete) 46.4 mm, diameter 12.8 mm. USNM 646045.

23. 24. Strioterebrum gausapatum (Brown and Pilsbry) (p. 411). 

Middle part of Gatun formation.

23. Height (incomplete) 33.6 mm, diameter 7.5 mm. Locality 155b. USNM 646040.

24. Topotype. Height (almost complete) 27.3 mm, diameter 6.7 mm. Locality 159d. USNM 646041. 

25, 31, 32, 36. Strioterebrum spiriferum (Dall) (p. 409).

25. Lectotype, an immature shell. Height (almost complete) 30.1 mm, diameter 7.8 mm. Mount Hope, Canal Zone. 

Upper part of Gatun formation, eastern area. USNM 113654.

31. Height (almost complete) 61.7 mm, diameter 13 mm. Locality 138d. Lower part of Gatun formation. USNM 

645850.

32. Topotype of Terebra gatunensis Toula. Height (incomplete) 58.5 mm, diameter 14.5 mm. Locality 159. Middle

part of Gatun formation, eastern area. USNM 645849.

36. Height (incomplete) 44.5 mm, diameter 12 mm. Locality 138e. Lower part of Gatun formation. USNM 645851. 

26. Pleuroliria (Polystira) ecuadoriana (Olsson) (p. 365).

Height (incomplete) 39.5 mm, diameter (incomplete) 22.5 mm. Locality 208. Chagres sandstone. USNM 645867. 

28. Strioterebrum indocayapum Olsson (p. 412).

Height (almost complete) 36.8 mm, diameter 8.7 mm. Locality 138c. Lower part of Gatun formation. USNM 646042. 

20. 30. fftrinterebrum wolfgangi (Toula) (p. 411).

Upper part of Gatun formation, eastern area.

29. Slender specimen. Height (incomplete) 37 mm, diameter 8.2 mm. Locality 175. USNM 646039.

30. Moderately slender specimen. Height (almost complete) 38.8 mm., diameter 9.5 mm. Locality 177b. USNM

646038. 

33, 34. Scobinella ecuadoriana Olsson (p. 373).

Height (incomplete) 43.4 mm, diameter 18.5 mm. Locality 208. Chagres sandstone. USNM 645863. 

37. Strioterebrum oresignum oresignum Olsson (p. 413).

Height (incomplete) 43.5 mm, diameter 10.2 mm. Locality 139c. Middle part of Gatun formation. USNM 646044.
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LATE MIOCENE MOLLUSKS FROM CHAGRES SANDSTONE
PLATE 63

FIGUEE 1, 2. Clathrodrilliaf sp. (p. 384).

Height (incomplete) 25.9 mm, diameter 9.7 mm. Locality 208. Chagres

sandstone. USNM 645868. 

3, 4. Strioterebrum monidum (Woodring) (p. 414).

Height (almost complete) 23.5 mm, diameter 5.8 mm. Locality 138c. Lower

part of Gatun formation. USNM 646047. 

5. Carinodrillia (Carinodrillia) sp. (p. 376).

Height (almost complete) 27.6 mm, diameter 10.3 mm. Locality 208.

Chagres sandstone. USNM 645864. 

6, 7. Darbya? (Buridrillia?) sp. (p. 377).

Height (incomplete) 15 mm, diameter 6 mm. Locality 208. Chagres sandstone. USNM 645865.

8. 13. Hemisinus amaras Woodring, n. sp. (p. 430).

Type. Height (incomplete) 20.5 mm, diameter 12.5 mm. Locality 161d. 

Middle part of Gatun formation. USNM 646071.

9. 10. Typhis (Talityphis) eucteanus Woodring, n. sp. (p. 434).

Type. Height (practically complete) 19 mm, diameter 9.7 mm. Locality 138f.

Lower part of Gatun formation. USNM 646083. 

11, 12. Cavolina (Cavolina') cf. C. ventricosa (Guppy) (p. 429).

Height 3.9 mm, width 4.3 mm, diameter 3.3 mm. Locality 177. Upper 

part of Gatun formation, eastern area. USNM 113855.

11. Ventral side.

12. Dorsal side. 

14, 15. Trigonostoma n. sp. (p. 345).

Height (almost complete) 20.2 mm, diameter 14.7 mm. Locality 208. Chagres

sandstone. USNM 645861. 

16, 17. Cavolina (Cavolina) triaspis Woodring, n. sp. (p. 428).

Type. Height (not quite complete) 4.2 mm, width 4.5 mm. Locality 177d. 

Upper part of Gatun formation, eastern area. USNM 646067.

16. Dorsal side.

17. Ventral side. 

18, 19. Pterynotus (Subpterynotus) textilis (Gabb) (p. 433).

Height (incomplete) 24.5 mm, diameter 19.5 mm. Locality 138c. Lower

part of Gatun formation. USNM 646082. 

20, 21. Colubraria obscura (Reeve) (p. 432).

Height (almost complete) 40.5 mm, diameter 15.5 mm. Locality 160c.

Middle part of Gatun formation. USNM 646081. 

22, 24. Turritella abrupta Spieker (p. 430).

Locality 138e. Lower part of Gatun formation. 

22. Height (incomplete) 68 mm, diameter 31 mm. USNM 646068. 

24. Height (incomplete) 37 mm, diameter 19 mm. USNM 646069. 

23. Petatoconchus sculpturatus H. C. Lea (p. 430).

Height of cylinder 28 mm, diameter 14 mm. Locality 139c. Middle part of 

Gatun formation. USNM 646070.
PROFESSIONAL PAPER 306-D PLATE 63

MIDDLE MIOCENE MOLLUSKS FROM GATUN FORMATION AND 

LATE MIOCENE MOLLUSKS FROM CHAGRES SANDSTONE
PLATE 64

[All figures X 66]

FIGURE 1. Agladrillia (Agladrillia) characta Woodring, n. sp. (p. 385).

Locality 147b. Middle part of Gatun formation. USNM 645795.

2. Drillia (Neodrillia) riogurabonis eurysoma Gardner (p. 374).

Locality 139b. Middle part of Gatun formation. USNM 645772.

3. Agladrillia (Agladrillia) phengoides Woodring, n. sp. (p. 386).

Locality 147b. Middle part of Gatun formation. USNM 645800.

4. Agladrillia (Agladrillia) enneacyma enneacyma (Brown and Pilsbry (p. 385). 

Locality 147b. Middle part of Gatun formation. USNM 645798.

5. Microdrillia trina Mansfield (p. 389).

Locality 147b. Middle part of Gatun formation. USNM 645810.

6. Kurtziella (Kurtziella) pagella Woodring, n. sp. (p. 393).

Locality 147b. Middle part of Gatun formation. USNM 645817.

7. Nannodiella rintriada (Mansfield) (p. 395).

Locality 147b. Middle part of Gatun formation. USNM 645828.

8. Agladrillia (Agladrillia) acaria ectypha Woodring, n. subsp. (p. 387). 

Locality 138. Lower part of Gatun formation. USNM 645802.

9. Kurtziella (Kurtziella) stenotella Woodring, n. sp. (p. 393).

Locality 147b. Middle part of Gatun formation. USNM 645819.

10. Agladrillia (Eumetadrillia) isthmica (Brown and Pilsbry) (p. 387). 

Locality 138. Lower part of Gatun formation. USNM 645805.

11. Lepicythara heptagona (Gabb) (p. 390).

Locality 157. Middle part of Gatun formation. USNM 645811.

12. Ithycythara defuniak Gardner (p. 391).

Locality 138c. Lower part of Gatun formation. USNM 645813.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D PLATE 64

12

PROTOCONCHS OF MIDDLE MIOCENE TURRIDS FROM GATUN FORMATION

368-278 O - 70 - 12
PLATE 65

[All figures X 56]

FIGTJKE 1. Gemmula vaningeni (Brown and Pilsbry) (p. 381).

Locality I47b. Middle part of Gatun formation. USNM 645759.

2. Pleuroliria (Poly stir a) tenagos (Gardner) (p. 364).

Locality 147b. Middle part of Gatun formation. USNM 645763.

3. Pleurofusia acra Woodring, n. sp. (p. 367).

Type. Locality 155c. Middle part of Gatun formation. USNM 645764.

4. Pleurofusia (Cruziturricula) fusinus fusinus (Brown and Pilsbry) (p. 369). 

Locality 146. Middle part of Gatun formation. USNM 645767.

5. Crassispira (Crassispira) henekeni leptalea Woodring, n. subsp. (p. 377). 

Type. Locality 147h. Middle part of Gatun formation. USNM 645777.

6. Crassispira (Hindsiclava) consors consors (Sowerby) (p. 378).

Locality 138c. Lower part of Gatun formation. USNM 645780.

7. Clathrodrillia gatunensis (Toula) (p. 381).

Locality 138c. Lower part of Gatun formation. USNM 645789.

8. Clathrodrillia saavedrai Woodring, n. sp. (p. 383).

Type. Locality 147b. Middle part of Gatun formation. USNM 645790.

9. Carinodrillia (Carinodrillid) zooki (Brown and Pilsbry) (p. 375). 

Locality 147g. Middle part of Gatun formation. USNM 645775.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D PLATE 65

 f/

a»r ..***

,'

PROTOCONCHS OF MIDDLE MIOCENE TURRIDS FROM GATUN FORMATION
PLATE 66

[All figures X 56]

FIGUEE 1. Strioterebrum wolfgangi (Toula) (p. 411).

Locality 174. Upper part of Gatun formation, eastern area. USNM 135599.

2. Dolostoma anorhepes Woodring, n. sp. (p. 398).

Locality 147b. Middle part of Gatun formation. USNM 645824.

3. Glyphostoma (Glyphostoma) dentiferum Gabb (p. 399).

Locality 138c. Lower part of Gatun formation. USNM 645852.

4. Strioterebrum spiriferum (Dall) (p. 409).

Locality 138c. Lower part of Gatun formation. USNM 645852. 

5, 7, 9. Spiratella inflata ekvata (Collins) (p. 427).

Locality 136a. Lower part of Gatun formation. USNM 646066. 

6. Glyphostoma (Rhiglyphostoma) allodapum Woodring, n. sp. (p. 402).

Paratype. Locality 138d. Lower part of Gatun formation. USNM 645836. 

8. Euclathurella (Miraclathurella) eucharis Woodring, n. sp. (p. 397).

Locality 147b. Middle part of Gatun formation. USNM 645826. 

10. Strioterebrum indocayapum Olsson (p. 412).

Locality 147b. Middle part of Gatun formation. USNM 646043.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 306-D PLATE 66

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PROTOCONCHS OF MIDDLE MIOCENE TURRIDS AND TEREBRIDS, 

AND PTEROPOD FROM GATUN FORMATION

U. S. GOVERNMENT PRINTING OFFICE : 1970 O - 368-278