Citation
Geological Survey professional papers

Material Information

Title:
Geological Survey professional papers
Abbreviated Title:
Geol. Surv. prof. pap.
Creator:
Geological Survey (U.S.)
Place of Publication:
Washington, D.C.
Publisher:
U.S. G.P.O.
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Frequency:
Irregular
completely irregular
Language:
English
Edition:
NO. 306 D-E
Physical Description:
1083 v. : ill., maps ; 29 cm.

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Subjects / Keywords:
Geology -- United States ( lcsh )
Aardwetenschappen ( gtt )
Geologie ( gtt )
Geology ( fast )
United States ( fast )
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serial ( sobekcm )
federal government publication ( marcgt )

Notes

Citation/Reference:
Chemical abstracts
Dates or Sequential Designation:
216-1298.
General Note:
Cataloged separately in LC after no. 688.
General Note:
Monthly Catalog Number: gp 81009497
General Note:
Most titles sold on an individual basis by the Supt. of Docs., U.S. G.P.O.

Record Information

Source Institution:
University of Florida
Holding Location:
Centers of Excellence at UF
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This item is a work of the U.S. federal government and not subject to copyright pursuant to 17 U.S.C. §105.
Resource Identifier:
023671617 ( ALEPH )
01445516 ( OCLC )
gs 14000289 ( LCCN )
0096-0446 ( ISSN )
Classification:
QE75 .P9 ( lcc )
551 ( ddc )
I 19.16: ( sudocs )

Related Items

Preceded by:
Professional paper (Geological Survey (U.S.))
Succeeded by:
U.S. Geological Survey professional paper

Aggregation Information

DLOC1:
Digital Library of the Caribbean
PCM:
Panama and the Canal
IUF:
University of Florida
IUFGOV:
Centers of Excellence at UF
UFPANCAN:
Documents of the Panama Canal

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Full Text




















UNIVERSITY
OF FLORIDA L I BR AR IES

















cl-,


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


SEP 197
%~$ ALL/4 41
-T
.S-


oe"


ttAT OF













Geology


and


Paleontology


of Canal Zone and



Adjoining Parts of Panama


Description of Tertiary Mollusks (Gastropods: Eulimidae, Marginellidae to H elminthoglyptidae)

By W. P. WOODRING


GEOLOGICAL


SURVEY


P ROF E S SIONA L


PAPER


A4 contribution to thie history of the Panarnai /and bridge


UNIEDSTAESGOVERNMENT PRINTING OFFICE, WASHINGTON 17


3 06 -D


UNITED STATES


: 19 70


ttAT OF



ww.
01A 3, Sh)

























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 2.25 (paper cover)

























CONTENTS


A bstract-- - - - - - - - -
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--E ocene series-- - - - - - -
Gatuncillo formation - - - - -
Eocene or Oligocene series- - - - - -
Marine member of Bohio(?) formation----Oligocene series-- - - - - - - -
Bohio formation - - - - - -
Caimito formation - - - - - -
Caraba formation-- - - - - -
PanamA formation - - - - -
M iocene series - - - - - - - -
Culebra formation - - - - -
Cucaracha formation --- --- -- -
La Boca formation, including Emperador limestone m ember - - - - - -
Alhajuela formation-- - - - -
Gatun formation-- - - - - -
Chagres sandstone, including Toro limestone
m em ber -- - - - - - -
Hlolocene 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


Description of Tertiary mollusks-contin ned from chapte r C - - - - - - - - - -
Gastropods-continued from chapter C -----Family Eulimidac - - - - - -
Family M~arginellidac - --- --- --Family Cancellariidae-- - - - -
Family Conidae - - - - - -
Family Turridae - - - - - -
Family Terebridae - - - - - -
Family Pyramidellidac ---------Family Acteonidae - - - - - -
Family Ringiculidac - - -- -
Family Scaphandridae ----------Family Philinidae-- - - - - -
Family Bullidae - - - - - -
Family Atyidne-- - - - - -
Family Retusidae-- - - - -
Family Spiratellidae--- - - - -
Family Cavolinidae - - - - -
Family Ellobiidae-- - - - - -
Family Hielmninthoglyptidae --------Additions and corrections to families covered in
preceding chapters - - - - -
R eferences cited - - - - - - - -
In d e x - - - - - - - - - - - -


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 Caimnito formations, and early Miocene miollusks 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 MAiocene mollusks from La Boca formation and Cliipola formation of Florida.
51. Early Miocene mollusk from Emiperador limestone member of La Boca formation and middle Miocene mollusks from Gatun formation.
52-6 1. Middle Miocene mollusks from Gatun formation.
62, 63. Middle Miocene mollusks from Gatun formation and late iMiocene mollusks from Chagres sandstone.
64, 65. Protoconchis of middle Miocene turrids from Gatun formation.
66. Protoconchis of middle Mliocene turrids and terebrids, and lateropod from Gatun formation.


Page

326 326 326 330
334
345 360
403
414 415 416 417 420 421
422 423 426
427 429 429

430 435 441




















GEOLOGY AND PALEONTOLOGY OF CANAL ZONE AND ADJOINING PARTS OF PANAMA


HELMINTHOGLYPTIDAE)



By W. P. WOODRING


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 Turridac, represented by 70 species and subspecies.
Only two of the species were found in the Gatuncillo formation, thc 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 Bohjo formation yielded ,Strio tcrcbrurn listro turn, the first unequivocal Striotcrebrum from the Oligocene of the Caribbean region. The fossils from the Caiinito formation, also of late Oligocene age, include Vagincila lophota, the first Oligocene American pteropod of the genus Vaginclia, and the earliest pteropod of the genus Cavolina: C. xcnica.
The name Carabra formation is adopted for strata~formerly assigned to the Caimito formation. The Panam6. formation is redefined to include strata in the Pacific coastal area formerly assigned to the Bohio and Caimito formations. Both the Caraba and Panarnd 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 froin the La Boca described in the present chapter include Floribella aldrichi, a remarkable philinid, and Ellobiurn aff. E. pellucens, the first American fossil representative of the salt-marsh pulmonate genus Ellobium. The Emperador limestone member of the La Boca yielded Cam panile cf. C. hcrculcanus. That form and C. hercitleanus itself are the last American species of the genus, younger than any in Europe. The fine-grained rocks in Madden basin, overlying the agglomierate of the pyroclastic-clay member of the Caimito formation of former usage, are now assigned to the La Boca.


The early Miocene Alhiajuela formation is defined to include the youngest strata in Madden basin. It includes a lower member (tme calcareous sandstone member of the Caimito formation of former usage) and an upper member (the Alhiajuela 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 conlds, 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 MAiocene age is adopted for the Chagres sandstone, instead of early Pliocene.
According to radiocarbon dates, the informally named Atlantic and Pacific inuck, 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 B~oca, and Gatnn 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 uiined 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 thle scaphopods, pelecypods, and cephialopods: an~ estimated 250 species.
As shown in the following table, tile gastropods so far described are unevenly distributed in the marine formations.
299







300 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Species of gastro pods in marine formations


Number
Of.
species


Formation
Eocene series:
Gatuncillo
Eocene or Oligocene series:
Marine member of Bohio(?)-Oligocene series:
Bohio
Caimito Caraba
Miocene series:
Culebra
La Boca, Gaillard Cut areaLa Boca, Madden basin-Alhnjuela - - - -
Gatun
Chagres-- - - - -


33

37

29 41
2

32 78
6
5
293 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 overlyinig Aihajuela formation consist chiefly of pelecypods.
Inasmuch as the gastropods make up about twothirds of the molluscan faunas, it is appropriate at this tie to reappraise the age of the fossiliferous formations. It is appropriate also to correct errors in stratigraphy and area]. 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 Panamd. that year and in 1949, 1954, 1959, 1965, and 1967. The work in 1965 was specially rewarding so far as the La B~oca formation is concerned. At that time, as a result of the prograii 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 amn specially indebted to R. H. Stewart, geologist of the Panama Canal Company, and his assistant Joanne Allen 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 B~oca, and Gatun formations. Mr. Stew~art also offered advice on subsurface geology based on his study of thousands of core holes. I have profited from advice from miy colleagues A. A. Olsson and Druid Wilson, and from members of the staff of the Division of Molliisk.s, of the U.S. National Museum.


CHANGES IN FORMATION AND AGE ASSIGNMENTS

Trhe geographic and geologic features mentioned in the following- summary are shown on plate 1, chapter A.
TjfJe name Caraba formation, defined on page 307, is adopted for strata in the Rio Mandinga area (south of Gamiboa), 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.
Thle Panamii 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 IBohio and Caimito formation are reassigned to the Panamd.
The agglomierate and tuff in the so-called pyroclasticclay member of the Caimito formation in Madden basic (p. 32) are identified by M.Stewart as the Las Cascadas agglomerate andl the overlying fine-grained nonvolcamce rocks as the La B~oca formation. The finegrained rocks include coralliferous limestone like the Emperador limestone member of the La Boca in the Gaillard Cut area.
The Aihajuela formation is defined on page 31T to include a lower member (the calcareous sandstone member of the Caimito formation of page 32) and an upper miem-ber (the Aliajuela 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 M iocen e.
Aihajuela sandstone member of Caimito formation, early M1i oeene.
Calcareous sandstone member of Caimito formation, early Mioceene.
Pyroelastie-elay member of Caimito formation, late Oligocene.
Caimito formation, Rio Mandinga area, late Oligoceene.
Panam~i formation, early Mviiocenre.
Bohio and Caimito formations, Pacific coastal area, late Oligocene.


Present usage

Chagres sandstone, late Miocene.
Upper part of Gatun formation, western area, middle M iocene.
Upper member of Albajuela formation, early Miocene.

Lower member of Aihajuela formation, early Miocene.

Las Caseadas agglomerate,
Oligoeene(?) and La Boca formation, early Miocene. Caraba formation, late
01ligocene.

Panaimi formation, late
Oligoeen e.
Panami formation, late
Oligocen e.


300







GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO HELMINTIIOGLYPTIDAE


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 curybathrodon, 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 pls.
Blake's species is briefly described on p. 438.
1955. White, E. I., On Lanina curybathrodon 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., Panamd: XX Cong. G6,ol. Internal., Coin.
Stratigraphique, Lexique Stratigrapbique International, v. 5, Am~rique Latine, fasc. 2a, Am~rique 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 C"4 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 inidl-Tertiarv
stratigraphical correlation, 163 p., 17 pls., 20 figs.,
Cambridge Univ. Press.
Gatuncillo, Bohio, and Caimito formations are of
Aquitanian (early Miocene) age (p. 36-37).
1963. Yokes, E. H., Cenozoic Murieide 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 Murcx 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 seawvays: 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 Muricidoe of the western Atlantic
region; pt. 2, Chicorcus sensu stricto and Chicoreus (Siratus): Tulane Studies Geology, v. 3, no. 4, p.
181-208, 4 pls.
Includes a species from Gatun formation.
1966. Woodring, W. P., Estratigrafia Terciaria de la Zona del
Canal y partes adyacentes de la Repilblica de Panamnd (Tertiary stratigraphy of Panama Canal Zone andl adjoining parts of the Republic of Panamd): Inst. Centroamericano In. y Teenologia 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 pls.,
1 fig.
Includes a species from Caimito formation and
one from Gatun formation.
1967. Bold, W. A. van den, Ostracoda of the Gattin formation,
Panamd: Idem, v. 13, no. 3, p. 306-318, 2_pls.
Twenty-eight species.
1969. Blacut, Gustavo, and Kicinpell, R. M., A stratigraphic
sequence of benthionic smaller foraminifera from the La Boca formation, Panama Canal Zone: Cushman Found. Foram. Research Contr., v. 20, pt. 1, p. 1-22,
pls. 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. 389390, 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. use(] ill


23b




37a


USGs
Cenozoic No.


24553 23648


Description of locality Gatuncillo formation


Upper course of Rio Palenque, 3.4 kmn in
dirct linc west of Nucvo San Juan and 1.3 km northwest of scttlcmcnt of Palenque, Col6n Province, Panami. Silty mudstone.
RI. H. Stewart, J. L. Allen, and Anselmo
Mena, 1968.
Top of Ccrro Pelado, 1 km north-northwest of
Gamboa, Canal Zone, altitude 223 mn.
Leached, soft sandstone. R. H. Stewart,
1964.


301







GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


No. USGS
used in Cenozoic Description of locality this No.
report

Culebra formation

112b 24502 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.
112c 24505 Panama Canal, east side of Culebra Reach, opposite Empire-Culebra slide, about Canal station 1730, Canal Zone. M. 1. Goldman, 1912.

Cucaracha formation

122c----------Panama Canal, west side of Cucaracha Reach, northeast slope of Cerro Escobar near Borinquin Highway, Canal Zone. Somewhat carbonaceous shale (coprolite bed) about 3 mn above ash flow. J. L. Allen, 1965.

La Boca formation

101a 23650 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. 101b 23651 Panama Canal, west side of Las Cascadas Reach, Canal station 1631 plus 15 m, Canal Zone. Basal echinoid-bearing limestone. W. P. Woodring, 1965. tlei 23656 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.
told 23668 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.
101e 23655 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. lOif 23653 Panama Canal, west side of Las Caseadas Reach, Canal station 1602, Canal Zone. Limestone at top of canal cliff. W. P. Woodring, 1965.
l0ig 23654 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. 101h 23652 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. Allen, 1965.
loui 23658 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. 11. Stewart, 1964.


No. Used in this report


1 17a 117b 11 7c


117d 118


USGS
Cenozoic No.


23662 23661 23660 23657 23659


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 1)1. 2, chap. A. Rt. H1.
Stewart, 1960.


Lower part of Gatun formation

138f 23663 South side of Transisthmian Highway, hillside excavation at Colehoneria Yero, about 450 m southwest of Cativa, Panamdi. R. H. Stewart, W. P. Woodring, and others, 1965. 138g 23664 South side of Transisthmian Highway, about 35 m east of 23663, Panamd. R. H. Stewart, 1965.

Middle part of Gatun formation

143a 24504 Panama Railroad, first cut southeast of Camp Totten Canal Zone. [Near locality 143.] M. 1. 4xoldman, 1912.
159d 24173 Gatun, Canal Zone. [Presumably Gatun Locks excavation.) M. I. Goldman and others, 1912. 160b 23665 South side of Rio Chagres, 175 mn below spillway of Gatun Dam, Canal Zone. Coralliferous conglomerate. W. P. Woodring, 1965. 160c 23666 Same locality as 23665. Silty sandstone underlying conglomerate. W. P. Woodring, 1965. 160d 24503 West end of Gatun Dam, Canal Zone. M. I.
Goldman, 1912.

Upper part of Gatun formation

177e 24174 Mount Hope borrow pits, Canal Zone. M. I.
Goldman, 1912.


302






GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE


NEW GENERIC AND SUBGENERIC NAMES

The following new generic and subgeneric names are proposed.
Dolostoina, Turridae, Mangeliinae.
Type: Dolostoma anorhepes Woodring, n1. sp., Gatun formation, Miocene, p. 398. Gender neuter. Euglyph ostoma, subgenus of Gly p/iostoma, Turridae, Mlangeliinae.
Type: Glypliostomna pairtefiosa Dali, Gulf of California and west coast of Baja California, p. 401.
Gender neuter.
Floribella, Philinidae.
Type: Dolabelda aldriclii Dali, Miocene, Florida,
Canal Zone, Cuba, p. 420. Gender fem-inine.
Lecailia, subgenus of Aceellia, J-ehninthoglyptidoe,
Xanthionycinlae.
Type: A'verelia (Lecallia) steiar~ti Woodring, n.
sp., marine member of Bohilo(?) formation, late Eocene or early Oligocene, p. 429. Gender neuter. Paleocavolivra, subgenus of Car olin a. Cavolinidae.
Type: Cavolina (Paleocaeolina) wenica Woodring, in. sp., Caimnito form-ation Oligocene, p. 428. Gender feminine.
Rlhiglyphos torna, subgrenus of Glyphostom a, Turridae,
Mangelinae.
Type: Glypltostoma xes ton 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-mieter 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 H-ill 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 Cuilebra form-ation'. Ruth Todd reports that the poorly preserved foram-inifera include Cassigerinella, indicating a latest Eocene or early Oligocene age. Another subsurface locality (Core Hole CI-J-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 layano district, east of the area covered by


plate 1, indicate that probably all of central Panama. was submerged in late Eocene timle.
F lat-lying limestone of the Gatuncillo, resting on an irregular surface of the basement complex, was found along and near a tributary of Rio Agna Sucia a kiloieter west of locality 27 on the Transisthimian 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. Thle 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: Pe)rsicida cf. P. semien and Conus cf. C. saurideins, both silicifled 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. A, bout 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:

Gastro pods from Gatuncillo formation and occurrence elsewhere of same or related species


Species fromn Gatuneillo formation

Vclates perversus (Gmelin), subsp.?
Turritella cf. T. carinata Lea_Turritella cf. T. sainanensis
Olssoli.
Dirocerithizan ante WoodrinoHannatoma cf. H. emendorferi
Olsson.'I

Calyptraea cf. C. aperta
(Solander).

Qostrombus aff. 0. ehiraensis
(Olsson).

Terebelium (Terebeltum)
procerum Merian.2
Tercbellum (Seraphs)
belemnitam (Palmer)?.
Ectinochilus cf. E. gaudichaudi
(d'Orbigny).
Cypraedia aff. C. suibelegans
(Trechmiann).
Pachycrommium? solenacumn
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 (Heiprin),
middle Eocene, southeastern U.S.
H. emendorferi, late Eocene, or
early Oligocene, Peril; late
Eocene, Columbia,
Venezuela.
C. aperta, Paleocene to
Oligocene, western Europe, southeastern U.S., Miocene,
Maryland.
0. chiraensis, late Eocene or
early Oligocene,, Peril; 0.
tournoucri (Bayan), middle
Eocene, Italy.
Middle Eocene, Jamaica,
Haiti, St. Bartholomew;
late Eocene, Trinidad. T. belemnitum, late Eoeene,
Florida.
E. gaudichaudi, late Eocene,
Peril.
C. subelegans, middle Eoeene,
Jamaica.


303








GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Gastro pods from Gatuncillo formation and occurrence elsewhere of
same or related specie s-C ontinued


Species from Gatuncillo formation


Galeodca? cf. G. nodosa
(Solander).
Yasila aff. Y. paytensis OlssonXancus cf. X. peruvianus
(Olsson).

Cteniiyria ctcnista WoodrinrrPersicula (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. paylensis, late Eocene', Peri. X. perusianus, middle Eocene to late Eoeene or early Oligocene, Peri.
C. coroni (Morlet), middle to
to late Eocene, western
Europe.
P. semen, middle to late
Eoeene, southeastern U.S. C. sauridens, middle Eocene to
early Oligocene, southeastern U.S.


I See following paragraph.
2 Described on 1). 192 as Terebeflum procerum?.


]Hnniatom~a cf. H. emnendoiferi wvas described on page 638 as liannatoma? cf. Hi. e?)en dor ,feri. H-undreds of specim-ens of this species from the new locality 23b shiow that it is indeed a fianriatonia, to be described in chapter E as a new species. Hiannatomza is classified as a cerithicl not, a thiarid. That it is a brackish grenus is doubtful.
The Gatuncillo 'gastropods have Tethyan (TVelates, ferebelliun and its subgenus Seraphs), western European, southeastern United States, West Indian, and Perumian affinities. Those with southeastern United States and Peri~ are preponderant. IVelates. Diroceritli mm. Han atom-a, 0 ostrombus, Seraphs, Ectinochilus, Cypramdia, Pae/iycrommiunt, Yatsila, 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. Sm-raller foraminifera from locality 21 h1a ve been claimed to be of early Oligocene age (ekn,1964), although at locality 23, practically along the strike, limestone yielded late Eocene larger foraminifera (p. 20). The Gatmicillo mnay 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. Withi somne 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 (Lamnes and others, 1962, p. 36). As a matter of fact, the basis for that allegation the occurrence of Lepidocycliva patidos a tobleri (P/jolcpidina of Eames and others) -has been repudiated (Eames and others, 1968, p. 302).


EOENE OR OLIGOCENE SERIES

MARINE MEMBER OF BOHIO(?) FORMATION

The terni "marine member of Bohio (?~) formation" is a poor desig-nation for this unit. Hill's casual designation "Vamnos d. Vamros [-Vamios Vamnos] beds" cannot b~e adopted, as it cannot be defined properly. 1His locality, on Rio Chagres northwest of what is now lBarro Colorado Island (locality 40), is now under the waters of Gatun Lake and the stratigraphic relations were uncertain before it was snl)inergred. (See discussion p. 122--23.) If mapping can be carried out in the peninsula ending ini Palenquilla Point (locality 41) and the peninsula to the west, it mnay be possible to define properly the strata containing this distinctive f auna.
Species in the failies covered in chapter D) are tabulated below. In that and other distribution tables the designation "sp." In the locality columns indicates ain incomplete, or poorly preserved species that may or miay not b~e the same as that in the species column, and thie designation "?sp." indicates that the species is questioned. Sym-bols for relative frequency are as follows:


Symbols used for relative frequency


Symbol
Rrare -F, f ew-C, common-. A, abundant-


Number
Of
specimens
1-2 3-5
6-20
>20


Gastropodls from marine member of Bohio(?) formation (Eu limidac,
Turridac to Helminthoglyptidac
[R, rare; F, few; C, common]


Eulima cf. E. jaclesonensis
Gregorio - - - -
Niso (Niso) 'umbilicata (Lea) ?- -- Zemacies? sp a-- - - - Turriculine? turrid ------Sea phander (Sea phander) cf. S. jacksonensis Palmer-----Retusa (Cylicitnina) aff. R.
adamsi (Palmer) ------Volvulella (Volvulella) aff. V.
conradiana (Gabb) -----Averellia (Lecallia) stewarti
Woodring, n. sp --------------


vamos Vamos 40a 40d


F


C


Localities

Palenquilla Point

41 411)


F


R


U


C

U


RU -- -


Trinidad Island

42


F







?sp. R


If Ze?,acies ? sp. a is indeed a species of Zeqnacies, it is a representative of a Paleocene to Pliocene New Zeal and-Austral ianl genuis heretofore unknown in America. Averellia steivairti, the oldest land snail in1 Central America, is associated with 40 marine species


304








GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE


at locality 41b. It is the type of the subgenus Lecalla.
Thiirty-seveii species are so far recorded from the marine member of the Bobjo (?) formiation-a represenitation comparable to that for the Gatuncillo formation. H-alf of then 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:

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


Species from marine member of
B ohio (?) formation

Tricolia calypta Woodring

Tsirritella adela Woodring

Turritella cf. T. caleta Olsson1 Architectonica (Stcllaxis) aff.
A. alveata (Conrad).
Arch itcctonica ef. A. fsungia
(Conrad).
Eulimta cf. E. jacksonensis
Greg-orlo.

Niso (Niso) umbilicata (Lea) ?Neverita (Glossan lax) bolivarensis tapina Woodririg.
Globularia (Am pulella) SI)Globularia (Am pulella?) nana
Woodring.
Bach ycrommium? proi .num
Woodring.


Moriom (Cancellomorum) cf.
AM. antiquin (Bayan) 1. Ficus cf. F. mississippiensis
Conrad.
Typhis (Laevityphis) aff. T.
recurvirostris Conrad.
Glyptostyla panamensis Dall-.

Scaphandcr (Sea phander) ef.
S. jaclesonensis Palmer. Retusa (Cylichnina) aff. Rt.
adamnsi (Palmer).
Volvilella (Volvulella) aff. V.
conradiana (Gabb).
Averellia (Lecallia) stewarti
Woodring, n.sp


Occurrence elsewhere of related species


T. precursor (Dali), early
Miocene, Florida.
T. qalvesia Olsson, Oligocene,
Perdt.
T. caleta, Oligocene, Peil A. alveata, middle to late
Eocene, southeastern U.S. A. fun gia, 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 Yokes, middle
Eoene, California.
P.? gabrielensis (Clark), late
Eocene, Colombia; P.?
jaeksoncnsis (Harris), late Eocene, southeastern U.S. M1'. ant iquini, late Eocene,
Italy.
F. mzississippiensis, Oligocene,
southeastern U.S.
T. reen rvirostris, Oligocene,
southeastern U.S.
G. striata (Newton), middle
or late Eocene, Nigeria. S. jacksoncnsis, late Eocene,
southeastern U.S.
Rt. adamisi, middle Eocene,
southeastern U.S.
V. conradiana, middle Eocene,
southeastern U.S.
A. coactiliata (F6rrmisac),
Holocene, northern Mkxieo
to Trinidad.


1 identified as Turritella oissoni Clark, a late Eocene Colomibian species, by Allison (in Allison and Adegoke, 1969, p. 1254).
2 Described on page 203 as Aloruma ("Oniscidia") cf. ill. ant igum.


Age.-The gastropods from the marine miem-ber 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 pamanensis, which is redescribed on page 434 and reillustrated on plate 48, figure 23. As a result of reappraisal, Glyptostyla is


305


monotypic so far as America is concerned. A closely related species, however, G. striata (described as a species of S5trepsidura), occurs in the Eocene of Nigeria. Newvton (1922, p. 109) favored a middle Eocene age for the Nigerian deposits, but Eameis (1957, p. 30) preferred late Eocene. Of the genera and subgenera in the preceding lists, Stellaxi8. Glob idaria, Pachycroinrniuin, Clyptostyla. Zemacies?, 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 form-ation. It is possible that this Bohilo ( ?) f auna represents an unusual biofacies in the upper part of the Gatuncillo. Stratigraphic relations, however, are unknown. A late Eocene agre would agree with Cole's identification of the larger foraminifera, quoted on page 23.

OLIGOCENE SERIES

1BOHIO FORMATION

The only miollusks from the lBohio 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:
Gastro pods from upper part of Bohio formation on Barro Colorado
Island (Mlar ginellidae to Atyidae)
[R, rare; C, common; A, abundant] Locality 49d
Marginelta (Eratoidea) aff. 31. mnoilitor Dali ---------------R
Conus ef. C. sulculus Dali----------------------------- R
Conus aff. C. chipolanus Dali ---------------------------C
Drillia? (Neodrillia?) SI)---------------------------------- R
Strioterebrum listrotunt Woodring, nl. SI)------------------- R
Acteon (Acteon) aff. A. tainpae 1)all ----------------------R
Acteocina cf. A. bullata (Kicner) ------------------------A
Scaphander (Scaphander) cryus Woodring, n. sp -----------C
Atys (Roxaniella) rhadina Woodring, n. sp ---------------R

8trio tere b7ri?? listrotmr is the first unequivocal Strioteirebrion, from the Oligocene of the Caribbean region.
Of the 29 species from the upper part of the Bohio formation 10 are not inmed specifically. Seven of the eioht unequivocally inmed 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 (A tys ni adivna) is found in the Oligocene Mint Spring mi-arl member of the Marianna llimiestone in southeastern United States, which is genorally assigned to the middle Oligocene. On the face of the table Miocene affinities far outweigh Oligocene. In the Caribbean region, however, Oligocene molluscanl faunas are meager conipared with those of Miocene age and for the most part consist of large, robust forms.








GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Gastro pods 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 lBohio formation


Solariella ephnidia WoodringTurritella of. T. altilira
Conrad.

Turritella listrota Woodring


Hemisinus (Longiverena)
oeciscus Woodring.
Cerithium (Thericium) mimeticum Woodring.
Orthaulax sp I1- - -


Clobularia (Clobularia) aff.
G. fischeri (Dali).

Pachycrommium aff. P. guppyi
(Gabb).


Gonysycon epomis Woodring. Gonysycon cf. G. epomis
Woodring.
Mitrella epacta WoodringMar ginella (Eratoidea) aff.
M. mollitor Dali.


Conus cf. C. sulculus Dall~Conus aff. C. chipolanus DaIl_Strioterebrum listrotum
Woodring, n. sp.
Acteon (Acteon) aff. A. tampae
Dali.
Acteocina of. 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
Mioccne, Jamaica; S.
depressa Dali, Hlolocene.
Subspecies of T. altilira range
from late Oligocene to late
Miocene.
7'. venezuelana Hodson, early
Miocene, Venezuela, Canal
Zone.
H. atriformis Cooke, late
Oligocene, Antigua.


Possibly related to 0. pugnax
(I-Ieilprin), late Oligocene,
Georgia, early Miocene,
Florida.
C. ischeri, 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; M1. 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. tam pae, early Miocene,
Florida.
A. bullata, early Miocene to
Holocene.
Doubtfully also La Boca
formation.
Middle Oligocene, Mississippi.


I Described on page 191 as Orthaulax cf. 0. pugnax (I1cilprin).



The Suwannee limestone of Florida, of late Oligocene age, offers a more suitable biofacies, but its fauna shows little affinity with that from the lBohio formation. Orthaulax, Glb lularia. Pachycroin'tium, and Gonysycon are extinct.
The larger foraminifera from- locality 42d (Cole, 1.957) 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 at moderately deep-water facies on Barro Colorado Island (localities 54g to 54n) and Palo I-Jorqueto 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 shiallow-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 (Archlitectonica and Conus) occur in both. (The Architeetonica of the moderately deep-water facies is an unidentified species.) Chapter D covers the species in the next two tables.

Gastro pods from shallow-water facies of Caimito formation (Marginellidae to Turridae)
[11, rare; F, few; C, common]


Marginella (Eratoidea) aff. M. mollitor DallConus sp--------------------------------Conus of. C. sulculus Dall -------Conus aff. C. chipolanus Dali -----Cemmula of. C. amica Casey ------Pleurofusia sp - - - - - - -


Localities
55b 56 57

*---------RC
-R--- R
--- -- RF
-F--- R -


54g


54hi RI



RI Rij


Rt


54i 54k


C Rt


R

Rt Rt


541


Rt


54m


Rt


54n 55a


--- - -


The turrids and pteropods in the preceding list are noteworthy. Carolina xenica, the oldest species of the genus, Is the type of the subgenus Paleocavolina.
Forty-one species of gastropods are recorded from the Calinito formation. H-alf of them are not named at the specific level. Five of the seven unequivocally identified species arc, endemic. The next two tables show


306


Gastro pods from moderately deep-water facies of Caimito formation (Conidae to Cavolinidae)
[R, rare; C, common]

Localities


Conus cf. C. peruvienus 0Olsson "Gemmula" sp -------Zemaries? sp. 1) -------Corhiespira? sop-------Clavino turrid ------:--Scobinella aff. S. morierei (cossmann)----- ---Paraborsorvia all. P. brasseenst .s (Mlansfield)------ -Sirioterebrum sp-------Rinqicula (Ringiculellai') sp Bulla? sp -- - - - -
Atys (Aliculastrum) sp----Veginella lophota Woodring, ni. sp----------------------Cavolina (Paleocavolina) xenica Woodring, n. sp----


K --- F







GASTROPODS: EULIMIDAE, MARGINELLIDAE TO IHELMINTIHOGLYPTIDAE


the distribution elsewhere and in other Canal Zone formations of the same or related species.

Gastro pods from shallow-water facies of Caimito formation and occurrence elsewhere and in other Canal Zone formations of the same or related species


Species from caimito formation


Turritelia meroensis Olsson-Turritella (Bactrospira)
altilira Conrad, subsp.1

Architectonica (Architectonica)
rhicna Woodring.
Trochita of. T. spirata Forbes ~ Orthaulax sp I - - -


Cypraea of. C. chilona D allGlob'ularia (Globularia) aff.
G. fischeri (Dall).

Ampullinopsis spenceri
(Cooke).

Semicassis (Echinophoria)
apenes Woodring.
Semicassis (Echinophoria) spCymatium (Scepta) ogygiuin
Woodring.
Ficus of. F. pilsbryi (B.
Smith).

Mar ginella (Eratoidea) aff.
M. mollitor Dall.


Conus of. C. sulculus Dall


Conus aff. C. chipolanus --Gemmula of. G. amica Casey-_


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

Late Oligocene, Santiago
area, Panamdt, Ecuador,
Perd.
Subspecies of T. altilira
range from late Oligocene
to late Miocene.
A. nobilis R16ding, early
Miocene to Holocene.
T. spirata, possibly middle
Miocene to Holocene.
Possibly related to 0. pugnax
(I-Ieilprin), late Oligocene,
Georgia, early Miocene,
Florida.
C. chilona, early Miocene,
Florida. Also Culebra and
La Boca Formations. G. fischeri, early Mliocene,
Florida. Also Bohio and
La Boca Formations. Late Oligocene, Antigua;
possibly Puerto Rico,
Santiago area, Panamd,
Ecuador, Perd.


S. intermedia (Brocehi)
Miocene to Pliocene,
Italy.
C. nicobaricum (Rd6ding),
Holocene.
F. pilsbryi, middle Miocene,
Jamaica, Dominican
Republic.
M. mollitor, early Miocene,
Florida; M. cuancycla
Gardner, early Miocene,
Florida. Also Bohio formation.
C. sulculus, early Miocene,
Florida. Also Bobjo
formation.
C. chipolanus, early M~iocene,
Florida. Also Bohio,
Culebra, and La Boca
formations.
G. amica, early Oligocene,
Mississippi.


I Described on page 104 as Turritella (Torcufa) altilira Con~rad, subsp.
2 Recorded on page 81 as Trochita cf. T. trochifornnis (Born).
3 Described on page 191 as Orthaulax cf. 0. pugna. (Ho-lprin).


Aqe.-The Caim-ito gastropods, like those from the lBohio formation, have Oligocene and Miocene affinities. Numerically Miocene outweighs Oligocene. Nevertheless Oligocene affinities are more pronounced for the Caimito than for the underlying lBohio and the percentage of extinct genera is almost twice as high for the Caimito. The two unequivocally identified species that are not endem-ic (Tuinitefla mneroensis and Ainpullinopsis spenceiri) occur elsewhere in deposits of


307


Gastro pods from moderately deep-water facies of Caimito formation
and occurrence elsewhere and in other Canal Zone formations
of related species


Occurrence elsewhere and in other Canal
Zone formations of related species

C. peruvianus, late Eocene,
Perd.
S. morierei, early to late
Miocene.'
P. brassoensis, middle Miocene,
Trinidad.
S. listrotum Woodring, Bohio
formation.
V. depressa Daudin, Miocene,
western Europe; V. chipolana
Dall, early Miocene,
Florida.


1 See page 373 for localities.


Oligocene age. Though Agnpullinopsis (Mle gatylotus 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 Staiplin) of western Europe, in southeastern United States, and in the Tertiary Caribbean province. The following genera and subgenierat are extinct: 0 vt/ianlax, Globularia, Agnpullinopsis, Echinophoria, "Genvua," Zernacies?, Scobivella, Parab ors onia and Paleocavolina.
The beautifully preserved planktonic and benthonic foram-inifera in the moderately deep-water facies of the Calimito on Barro Colorado Island were identified by Bolli and assigned by him to the Globorotalia kutgleri zone (lBolli, in Woodring, 1958, p. 22-23, 27). The smaller foramini fera, larger foram-inifera (p. 29-30; Cole, 1957), and miollusks indicate a late Oligocene age.
As part of the sweeping allegation that no marine Oligrocene is known in Amierica, except in the Tampico area of Mexico and Cuba, the Bohio and Caimito formiations have been alleged to be of Aquitanian (early Miocene) age (Eames and others, 1962, p. 36-37). The sweeping allegation has been tacitly repudiated (Eaml-es and others, 1968, p. 292-295).

CARABA FORMATION

On the recom-mendation of Mr. Stewart the name Caraba formation is adopted for strata formerly aissigned to the Caimlito formation. The name, in the formn Caraba facies of the Cainnito 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 kiloimeters southwest of Gamiboa and about 750 meters


Species from caimito formation


Conus cf. C. peruvianus
Olsson.
Scobinella aff. S. morierei
(Cossmann).
Paraborsonia aff P. brassoensis
(Mansfield).
Strioterebrum sp ------Vaginella lophota Woodring,,
n. sp.


Cavolina (Paleocavolina) zenica
Woodring, n. sp.







GEOLOGY AND PALEONTOLOGY OF CANAL ZO-NE


cast 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
metersr)


8. A ndesitie lava -- - - - - - - -
7. Agglom erate-- - - - - - - -
6. Sandy siltstone - - - - - - - -
5. Hard, dense, buff limestone; Cly peaster fragments -4. Sandy siltstone and calcareous, pebbly sandstone;
Clypeaster concavus and few mnollusks (locality 60)-3. Hard, dense, buff limestone; Clypeaster fragments---2. Sandy siltstone and poorly sorted, silty sandstone;
many Heterostegina issraelskyi, also Lepidocyclina
asterodisca (locality 59) - - - - - -
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 snore common than boulders; dacite prophyry
conspicuous among clasts --- --- ---- --


Approximate thickness of section.


15 60
8
39

18
8


18




70


200


These strata, characterized by the exceptional thickness of conglomerate andl conglomeratic sandstone, presumably interfinger with the Calinito 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 (pl. 1) as part of the Pedro Miguel aggoeaan in the area straddling the part of Madden H-ighway South of the Transisthimian (or Boyd-Roosevelt) Highway overpass as part of the Caimito formation. Exposures may be seen at the falls on the east side of Madden H-ighiway at the monument site four kilometers Soth of the overpass and on abandoned Army roads east of the highway. Lenses of dense marine limestone, containing calcareous atlgw and larger foraminifera, have been found at locality 97, off Madden H-ighway, and six kilometers north-northwest of Pedro Miguel.
The fossils so far recorded from the Caraba form-ation 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 foranmnifera, 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
Cuishmnan, near locality 97.1
Heterostegina israeiskyi Gravell
and Hlanna, locality 59.2

Lepiclocyclina asterodisca
Nuttall, locality 509.

Lepidocyclina sp., near
locality 97.
Coral:
Goniopora cf. G. cascadensis
Vaughan, locality 61.3


Gastropods:
Bach ycrommium? 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, M-'6xieo. Also
Caisnito formation.
Early Oligocene, _M(xico;
late Oligocene, Texas,
Trinidad.


G. cascadensis, La Boca
formation; late Oligocene,
Antigua; early Miocene,
Auguilla.

P.? trinitatens is, early
Miocene, Trinidad. Also
La Boea formation. Species of F. ventricosa
,group range from late Oligocene to Holocene.
Possibly also Caimito
formation.


Late Oligocene, Antigua;
early Miocene, Anguilla.


I Across Madden Highwxay from locality 97; identification by K. N. Sachs, Jr. 2See p. 30; identifications fly W. S. Cole.
3See p. 30; identification by J. W. Wells.
4 Sec p. 31; identification by C. W. Cooke.


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

PANAMA FORMATION

The Panamnd formation is redefined to include strata formerly assigned not only to the Paniania itself, but also to the lBohio and Cainmito formations, and the P~edro Miguel agglomlerate. As redlefinedi the Panamni' consists chiefly of agglomierate and tuff, extending from the Miratlores Lake area to Panamni! City, and also northeastward across the continental divide and east ward in the Pacific, coastal area to and beyond the limit of plate 1. Tjh formation also includes tuffaceous sandStonle, tuffaceous siltstone, lenses of stream deposits, andl lenses of marine lim-estone.
Mi'. Stewart showed mnany outcrop~s of agglomnerate, including iany new exposures resulting from highway construction and suburban development northeast of the main part of Panania City and along the Transistlimnian. Highwvay. Trle agglomerate consists of anguilar to suibroundedl blocks, mostly anclesitic, generally widely scattered in a matrix of fine-grained tuff. The


308






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE


proportion of angular and subrouncled blocks aci the amount, of tuff interbeddeci with, or ov-erlying, agglomerate changes from place to place.
Streamn deposits, made up of crudely bedded, tuffaceous sandstone containing scattered rounded, subrounded, andc subangular boulders and cobbles, formerly assigned to the Bohio formation, are shown onl plate 6. In the Pacific coastal area eastward from Rio Abajo tufiaceous sandstone and fine-grained tuft make up a progressively laroe-r part of the formation anci in the Tocii'nen Airport area farther east only tinegraineci tuft was seen in excavations during construction.
The thickest lens of I'liestone was observed at locaiity 44, albouit 150 meters north of the Transisthmlian Highway, where almost cylidrical stacks of algal limestone( rise above the general surface to a miaximumiii hieight of 6 meters. The lowest exposeci part of the limiestone contains scattered pebbles anci sand grains.
Pectinicis were found in algal limestone at, localities 44 and 45, but no other m-ollusks. Larger foraminifera froml algral limestone anid their (distribution elsewhere and in other Canial Zone formations are as follows:
Larger foramiinifera fromn Panarnid form a/ion and their occur rrene
elsewhere and in other Canal Zone formations
[Identifications by W. S. Cole]


Species from Panamdi formation


Heterostegina antilica
Cushman, localities 43,1
45,1 95.2
Lepidocyclina giraudi R.
Douvill6,5 localities 43, 95.


Lepidocyclina waylandvaughani
Cole, locality 45.

Lepidocyclina yurnaguncnsis
Vaughan,' localities 43, 45, 95.
Lepidocyclina van ghani
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
Caisnito formations.
Widespread in Oligocene,
especially late Oligocene, of
Caribbean region. Also Bohio, Caimito, and La
Boca formations.
Late Oligocene, Antigua,
Trinidad, M\6xico. Also
Bohio and Caimito formations.
Widespread ia late Oligocene
of Caribbean region. Also
Caimito formation.
Late Oligocene, Antigua. Also
Bohlo 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, Cairnito, Cuilebra, and La Boca formations.


309


Age.-The. larger foraminifera in the preceding list aire typical for a late Oligocene age in the Caribbean region. All except two occur also in the Cain-ito formation. Two range upward into the La Boca formation. Like the Caraba form-ation, the Panalii form-ation is inferred to be the equivalent of part, or all, of the Caimito. The early Miloceiie age formerly assigned to the Panamad was based onl supposed stratigyraphic relations to the La Boca, which hiave turned out to be erroneous.
Between Madden H-ighway and the Chiva Chivat roaci (the road extending from Miraflores Lake northeast ward to the Transisthmlian 11Iighway)te ara anid Panaii formations presumably interfinger. Much fieldwork, however, remiais to be clone in that heavily forested and other areas before the stratigraphic relations of the agglonerates of those formations and of the Las Cascadas agglomeirate are properly known.

IOCENE SERIES
CULEBRA FORMATION
As restricted oni pagre 9-44, 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 onl page 1244, 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 onl the Gatincillo 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, Bloca formation. In the Culebra Reach area the transgression was interruptecd by dieposition of a northwestward thinning wedge of nonmarine tuff, later altered to bentonitic clay, of the Cucaracha, formation. The Culebra itself wedges out north-westward, as along the Empire and Las Cascaclas reaches the La, Boca overlies the Las Cascadas agglomerate; that is, as the marine tranlsg1'ression continued, it, extended farther northward. During La B~oca time doubtless the entire central Panaina, area, was submerged.
The present chapter covers five species from the Culebra formation, but none is specially noteworthy. Onl account of the transfer of fossil localities, the table onl page 3010 includes all of the Culebra gastropods: 32 species, about half of which are not named at the specific level.


I See p. 27.
2 See p. 33.
3Recorded on p. 27 and 33 as Lcpidocyclia percuae Cushman.
4 Includes Lepidecyclina yurnaguncnsis mnorganepsis Vaughan of p. 27 and 33.








310 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE1

Gastro pods from Culebra formation (Neritidae to Terebridae) and occurrence in La Boca formation
[II, rare; F, few; C, common; A, abundant]

Localities La
____ ____________ ____________ -___ _______ -_____ ___ Boca for
102 103 104a 104b 106 107 108b 108c 110 110a lila 1l1b 112 112a mation

Neritina (Vitta?) sp ------------------------------ ---- ---- ---- ---- ---- ---- ---- F -- ---- ------- ---- ----Littorina aff. L. angulif era (Lamarck) ------------------------------ ---- -------------R ---------------Turritella (Bactrospira?') amaras Woodring I--------- ---- ---- ---- ---- -----A R F A R C F C F X
Turritella venezuelana Hodson --------------------- ---- ---- ---- ---- ---- R-----------F ---C ---F C X
Pot amides suprasulcatus (Gabb) ------------------- ----B R U -- R --- C C ---- -------------- -----X
Terebralia, dentilabris (Gabb)? ------------------------ ------------------- 1--------------------- ----Hipponix? sp ---------------------------------------------------R-----------------------------Calyptraea cf. C. centratis (Conrad) ---------------- ---- ---- -------- ---- ---- -----R ---R-------------- ------X
Crucibutum sp ------------------------------------------------ --1 ---- ---- ---- ------- ---- ----Strombus sp ----------------------------------------------------R----- ---- ---- --------- ---- ----?R ----Orthaulax? sp -------------------------------- ---- ---- -------- ---- ---- ---- ---- ---- R---- ---- ----- ---- ----Cypraea cf. C. chilona Dali ----------------------- ------------------------ ---------U----- ---- ---------- -----X
Natica (Naticarius?) sp --------------------------- --------------------------------- ----R---R---- ----- -Polinices? sp ---------------------------------------- ---- ---- ---- R ----------------------- ---- ----Neverita? sp--------------------------------------- ---- ------------ R---------------------------- R---- ----Sinum sp------------------------------------------- ------------R-U------- ---- -------------------- ----- R----Pachycrommium? cf. P. guppyi (Gabb) ------------- ------------------------ --------------------------R ---X
Semicassis? (Tylocassis?) cf. S. aidrichi (Dali) ------- -------- ---- ---- ---- ---- ---- ------R----------- ---U-- -Ficus carbasea micronematica (Brown and Pilsbry)~ -R-------- -------- -------- ------------ -----R-----------X
Murex (Siratus) cf. M. pot ynematicus Brown and
Pilsbry2--------------------------------------------------------------- -----U--- R --------------------- X
Metuta sp ------------------------------------- -------- -------- ---- ---- ---- -----R----- ---------------U --Cymatophos? of. C. veatchi (Olsson) ---------------- ------------------------ ----------F---------------RU -- X
Antillophos? (Antillophos?) cf. A candei gatunensis
(Toula) -------------------------------------U- --------------R-----------------------F ---R -- X
Melongena sp----------------------------------- ---- ---- -------- ---- ---- ---- R RR---- ---- ---------R R--Fusinus? sp------------------------------------- ---- ---- ---- ---- ---- ---- ---- -------- ---- R---- ----- ---- ----Mitra? (Cancilla?) sp. .----------------------------------------------- ---- ---- ------Xancus of. X. vatidus (Sowcrby)4 ------------------ ---- ----UR R----------- --------------------------------- X
Persicuta (Rabicea venezuetana amydra Woodring,
n. subsp ----------------------------------------------------- R---- ---- ---- ---- -------------- ----- ---- X
Conus aff. C. chipotanus Dali --------------------- ------------U- ---- ---- ---- --------- ---- ---- ---- ----- ------X
Gemmua sp-----------s-------------------------- ---- ---- ---- ---- ---- ---- -- ---Terebra (Praterbra)------------------------------ -F -------------------- ----- ----------------- ----- --
Strioterebrum sp--------------------------------- ---- ---- ---- ---- ---- ---- ---- ---- ---- ---- R---- ----- ---- ----I Described on page 101 as Turritella (Torcula?) amaras.
2 Described on page 215 as Murex (Murex?') cf. M. polynematirus.
3 Recorded on page 283 as Mitre (Tiara) sp.
4 Described on page 286 as Xancus cf. X. reX Pilsbry and Johnson.







GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE


Though most of the species listed in the table lived in marine waters of normal salinity, Neretina, Littorina aff. L. angulifera, and Terebralia den tilabris? indicate brackish water, and Potamides sup vasulcatuts 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:

Gastro pods from Culebra formation and occurrence elsewhere and
in other Canal Zone formations of same or related species


Species from Culebra formation



Littorina aff. L. angulif era--(Lamarck).
Turritella (Bactrospira?)
amaras Woodring.1

Turritella venezuelana HIodson-

Potamides suprasulcatus
(Gabb).

Terebralia dentilabris (Gabb)?...

Calyptraea cf. C. centralis
(Conrad).

Cypraca of. C. chilona Dall---Pachyerommium? cf. P. guppyi
(Gabb).


Semicassis? (Tylocassis?) cf.
S. aldrichi (Dali).
Ficus carbasca micronematica
(Brown and Pilsbry).
Murex (Siratus) cf. M. polynematicus Brown and
Pilsbry.2
Cymatophos? of. C. veatchi
(Gisson).


Antillophos? (Antillophos?) of.
A. candei gatunensis (Toula).

Xancus cf. X. validus
(Sowerby) .3

Persicula (Rabicca) venezuelanaamydra Woodring, n. subsp.

Conus aff. C. chipolanus Dall--


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

L. angutif era, 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, D)ominican
Republic. Also Bobjo and
La Boca formations.
S. aldrichi, early Miocene,
Florida.
Early Miocene, Peril. Also
La Boca formation.
Ml. polynernaticus, Gatun
formation. Also La Boca
formation.
C. veatchi, Gatun formation;
middle Miocene, Costa
Rica. Also La Boca formnation.
A. candci gatunensis, Gatun
formation. Also La Boca
format ion.
X. validus, early to Middle
Miocene, Dominican
Republic. Also La Boca formation.
P. venezuelana venezuclana,
early Miocene, Venezuela.
Also La Boca formation. C. chipolanus, early Miocene,
Florida. Also Caisnito and
La Boca formations.


368-278 0 '70 2


Age.-Though the Culebra fauna is small, it has an unmistakable Miocene stamp-the earliest of the Miocene f aunas 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 Pack yerontlnium are extinct, but both genera are doubtfully identified.

CUCARACHA FORMATION

F resh-water snails of the genus Hernisinus (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 laud mammals recently found in the Cucaracha are considered to be of middle Miocene age (Whitmore and Ste-wart, 1965, p. 182).

LA BOCA FORMATION, INCLUDING EMPERADOR LIMESTONE MEMBER

Gaillard Cut area.-Isolated outcrops of fine-grained rocks of the La Boca formation have been accessible along the Canal, and also outcrops of sandstone somne 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 pro,gram 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 Bo0ca was exposed on the west side of the reach in a strip about 12,5 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. rjlje 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 IBeach at Canal Station 1630
Approxinlate
thickness
(vieters)


6. Emperador limestone member:
e. Noncoralliferomz limest one; Aequipecten canalis,
Antusium, Spondylmes, Clypeaster concavus
(locality 117b) --------------------------d. Poorly exposed sandy siltstone -------c. Coralliferous limestone; corals, small Spondylu -b. Sandy siltstone - - -- - - - -


1. 5
3
1.5 1. 5


311


I Described on p. 101 as Turr/ite (Torcita?) aniaras. Described on p. 215 as ilhcrcx (MorezIX) cf. 1. pet ijeematicas.
3 Described on p). 286 as Xaneus cf. X. rex 1Pilsbry and Johnson.







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. Emperadlor limestone member-Continued
a. Coralliferous limestone; corals, Turritella amaras
and other mollusks (locality 117a)-----5. Greenish silty clay-- - - - - - -
4. Carbonaceous shale-- - - - - - -
3. Siltstone, cla 'y, silty sandstone, poorly sorted sonmcwhat calcareous sandstone, and conglomeratic sandstone at foot of steep slope; mollusks in somewhat calcareous sandstone :30 mecters to northwest
(locality 101e) -- - - - - - - -
2. Siltstone and clay, poorly exposed on bench----1. Lens of disintegrating echinoid-bearing limestone just
beyond station 1632; mollusks and echinoids (Echinolampas ef. E. lycopersicus, Clypeaster concacus, identifications by' P. M. Kier; locality l0ib)
Las Cascadas agglomerate.
Approximate thickness of section--------


1. 5 1. 5






10. 15
12






36. 3


At station 1627 another len-s 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 101a). The echinoids, identified by P. M. Kier, are as follows: Eckivolainpa.s cf. E. lycopers icis and A gassioia clevei.
The Einperador limestone member was exposed on a bench at station 1626 (locality 117h). Corals, especially Acropora and Poi tes. and a few mollusks are strewn on the bench. The mollusks include five specimens of a bi- ceritid, Campanile cf. C. hercutleanits.
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 101h). This fossiliferous bed, characterized by the abundance of a small Aequipecten and a small Min ya, was not seen elsewhere. It is about 25 meters above the Emperador limestone member. The (011lection 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 aggl omerate, 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 foranunifeal samples, which have becen studied by IBlacut 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 siltstonc; 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 11--------------------------------------- 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 siltstonc; 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 in ---------------------- 9. 2
9. Claystone and mudstone, lower half dark gray, upper half chocolate brown ---------------------------1. 8
8. Soft mnudstone 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 branching0 corals -------------------------------- 2. 1
c. Thin-bedded, calcareous siltstone --------------- 2
b. Thin-bedded limestone, mat of broken branching
corals (locality 117d) ------------------------6
a. Thin-bedded, calcareous, sandy siltstonc ---------1. 5
6. Sandstone, mudstone, and carbonaceous shiale --------- 4
5. Mudstone and sandy siltstone; many Bittium scotti and molds of other sniall mollusks (locality 101d);
sample 410-1, 1.1 in above base ------------------3. 3
4. Sandy siltstone; mollusks, for most part fragments: Cypraea, Anadara, Trachycardium, Tagelus (locality
l0le)------------------------------------------.5
3. Carbonaceous shale------------------------------.6
2. M_1udstonc, black calcareous nodules; ostracodes, fragmentarv mollusks------------------------------ 1. 2
1. Bentonitic clay---------------------------------- 1. 3
Las Cascadas agglomerate.

Thickness of section--------------------------- 77. 2


H7al~f a kilometer farther northwest at lens of dense limiestonie crops out in the upper part of exposed strata, as shown in the following sections

Section of La Boca formation on west side o f Las Cascadas Reach
from Canal station 1605 to 1601
Approximnate
thiknmess
(mnete rs)
8. Silty sandstone and sandy* siltstone; scattered foraminifera and mollusks --- ------------------------- 15
7. Lens of white vitric tuff--------------------------- 1
6. Silty sandstone and sandy siltstonc ------------------6
5. Lens of dense limestone; Pachycrommiumn? cf. P.?
trissitatensis, Aequipecten canalis, Amusium, echinoids (EA-hinotamnpas semtiorbis, Clypeaster concavus, identilied by P. -\I. Kier; locality 101g)------------------4. 5
4. Sandy ,-siltstonie and silty sand(stonie, fewthin calcareouis
beds------------------------------------------ 20


312








GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO IHELMINTHOGLYPTIDAE


Section of La Boca formation on west side of Las Cascadas Reach
from Canal station 1605 to 1601-Continued
Approximate thickness metersr)


3. Poorly exposed sandy and silty beds -------2. Emperador limestone member:
e. Lenticular limestone, many Amusium -----d. Alternating siltstone and limestone in beds 30 to
60 cm thick -- - - - - - -
e. Lim estone - - - - - - - -
b. Siltstone------------------------------
a. Coralliferous limestone neair level of Canal --1. Carbonaceous clay, and tuffaceous siltstone ----Approximate thickness of section--------


7. 5


6

2
3

3


3


.5

70. S


Immediately South of the major fault at station 1598 the base of the La B~oca, overlyling the Las Cascadas agglomierate, consists of variegated clay overlain by fine-grained vitric tuft.
Core hole LU111 149, located 2,50 meters west of the top of the measured section at station 1622, penetrated strata higher than the measured section. Onl the assumiption that a 2.3-meter bed of vitric tuff penietratedl in the core hole corresponds to unit 13 of the measured section, the combined outcrop and stubsurface thickness of the La B~oca is 137 meters, and the top of the formation is not represented.
Madden basin.-As mentioned on page 300, the agyglomierate on the south side of Madiden basin, exposed on the Transisthmian H-ighway imluiedliately north of the intersection with Madden Highway, is identified iy Mr. Stewart as the Las Cascadas agglomierate and the overlying fine-grained rocks as the La Boca formi-atlin. Both were forinerly grouped as the pyroclastic-clay memi-ber of the Caimito formation (p. 32). The finie-grained rocks are poorly exposed or unexposed. Mol ds of foramnini fera, including Sipto gene rin a, were observed in calcareous mudstone on a tributary of Rio Chilibre (Quebrada A ncha, not, shown on plate 1) about 500 meters east-northeast of the Transisthimian TI-ighw-ay bridge across Rio Cillibre. 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 oni plate 1 evidently represent outcrops of different parts of ai limestone unit, now identified as the Emperador limestone nemi-ber of the La, B~oca. In unpublished reports it has been designated informally as the Chilibre limiestone.
Gastro pods, Gaillard (Cut area.-Seveuteen species from the La, Boca formation in the Gaillardl Cut, area are described or recorded in the present chapter. Porib5ella aidriehi is the most noteworthy of these species. It is considered to be a remarkable plilnid, by far the largest fossil or living species. Ellobiinn aft. E. pellu-


313


cens is the first Ame-irican fossil representative of the salt-miarsh pulm-onate genus Fllobium. Terebra diclle)es is the earliest known species of the subgenus Oreotere bra.
The La Boca, fauna has been greatly enlarged by the collections froln localities 101h and 116a (sole 60 and somec 90 species of mi-ollusks respectively). As for the Culebra formation, all the gastro pods from the La I3oca formation of the Gaillard Cut area are listed in the, table on pages 314-315. The table lists 7(8 species, half of which are not named at the specific level. Three of those not so named, chiefly froln localities 101hi and 116a, are to be described in chapter E. No mollusks have been recovered in Madden basin, except from the Emperador lunestone member. (See, table, p. 314.)
The occurrence elsewhere and in other Canal Zone formations of the same or related species is as follows: Gastro pods 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
I \'Iaury.
Turritella (Bactrospis'a?)
am eras Woodring.1

Turritella (Baclros pira) altilira
Conrad, subsp.2

T'urritelia subgr undifera Dall 3. Tns'rritella venezuelana HodsonT1mirritella cf. T. berjadincnsis
cocoditana H-odson.
Hem isinus (Lonqiverena) aff.
H. oeciscus Woodring. Potamides suprasulcalus
(Gabb).

Bitlium scoili Brown and
Pilsbry.
Architectonica (Architectonica)
nobilis Rd6ding, subsp.4 Calyptraea cf. C. centralis
(Conrad).

Orthaulax gabbi,1al C'ypraea ef. C. chilona Dall -Globularia (Globular ja) aff.
G. fischeri (Dall).

Pachycrommiium? ef. P.?
trinitatensis (Mansfield). Pachycrommium? cf. P.
guppyi (Gabb).


Miorum (Cancellomorunt) ef.
All. chipolanumn !Maury.5 Ficus carbaseanmicronematica
(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 Oligocene to
middle -Miocene. Also
Culebra formation.
B. permutabile, early Miocene,
Florida.
A. nobilis, early Miocene to
HIolocene.
C. centralis, early Miocene to
H-olocene. Also Culebra
formation.
Early Miocene, Florida. C. chilona, early Miocene,
Florida. Also Culebra, and
Caimnito 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. Ml. chipolanumn, early
M\'iocene, Florida.
Early Miocene, Perd. Also
Culebra formation.


Continued on page 316, with footnotes.







314 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Gastro pods from La Boca formation of Gaillard
[R, rare; F, few;

Localities

98 99a 99b 99c 99d 99e 99f 99g 99h 100 100a

A cm a ea ? sp - - - - - - - - - - - - - - - - - - - - - - - - -
L io tia s p - - - - - - - - - - - - - - - - - - -
N e r ita s p - - - - - - - - - - - - - - - - - - - - - - - - - -
N eritin a (V itta ?) sp - - - - - - - - - - - - - - - - - - -
C yolophorid, genus? - - - - - - - - - - - - -- - - -- - - -- -
Rissoina (Zebinella?') sp ------------------------------- ------- R --------- -------.-- -----Turritella cf. T. colt azica M aury -- - - - - - - - - -- - - -- - - - -- -
Turritella (Bactrospira?) amara8 Woodring I--------------------- 13- -------A --- ------ ------ ------ --Turritella (Bactrospira) altilira Conrad,' subsp-- - - - -- -- - -- - -- -- -- -- -- -- -
Turritella subgrundifera Dali I --------------------------------------R --------------------- -----Turritella venezuelana Hodson -------------------------- ------R -- ------ ----- --------------Turritella cf. T. berjadinensis cocoditana Hodson-- - -- - - - - - -- -- - -- - - -
Serpulorbis sp ------------------------------------ ---- ------ ------ ---- ?R---- ----??R-------- -----Hemisinus (Longiverena) aff. H. oeciscus W oodring-- - -- - - - -- - -- - -- -- -
Potam ides suprasulcatus (G abb) - - - - - - - - - - - - - -
C erithium (T hericium ) sp -- - - - - - - - - -- - - - -- - - - -- -
C a m p a n ile ?' sp -- - - - - - - - - - - - - - - - - - - - - - - - - -
Bittium scotti Brown and Pilsbry---------------------------- ---- -C----- ---- ------ ---- ---- ------ ---- C. A
M o d u lu s sp - - - - - - - - - - - - - - - - - - - - - - - - - -
A rchitectonica (Pseudotorinia?) sp -- - - - - - - -- - - - - - -- - - -
Architectonica (Architectonica) nobilis R16ding, subsp --------- -----------R R------- ---- ---- ------------R --Epitonium sp ----------------------------------------------------------R------------------- -----C irso trem a sp - - - - - - - - - - - - - - - - - - - - -
H ipponix of. H pilosus (D eshayes)-- - - - - -- - - -- - -- - - -- - -
Calyptraea cf. C. centrals (Conrad) ----------------------- ---------- C F------- --------- ------ -----R ---T ro ch ita sp .4 - - - - - - - - - - - - - - - - - - - - - - -
Crepidula sp ------------------------------------------ ---- -------RP ---------- ---- ---- ------ ---- -----Xenophora sp------------------------------------------ ---- ------------ ---- ------ ---- ---- R -------Strombus sp ----------------------------------------------- ------ ------P. C -- R F F -----Orthaulax gabbi Dali --------------------------------------------- ---------- ?sp.R --------- sp.R------------ sp.R
Cypraea cf. C. chilona Dali ------------------------------ ---?sp.P.------- ---- ------ ---- ------ -------P--N atica sp., operoulumn - - - - - - - - - - - - - - - - - -
Polinices?' sp ---------------------------------------------- ------ -----------P.------------------ -----NeveritP? sp ------------------------------------------- ----P. A F R ---P ---.-- -----Sinum sp ------------------------------------------------- ------ ------ F F ----------------Globularia (Globularia) aff. G. ftscheri (Dali) -------------- -R-------------P.-- --- ----- --- --Pachycrommium? cf. P.? trinitatensis (Mansfield) ------------ ------- ---- --------- ---- ---- ------ R-----------Pachycrommium? cf. P. guppyi (Gabb)-- - - - - - -- -- - -- -- - - -
C a s s is s p - - - - - - - - - - - - - - - - - - - - - -
Sem icassis (E chinophoria) sp -- - - - - - -- - - -- - - - -- - - -- -
S em icassis (T ylocassis) sp -- - - - - - - - - -- - - -- - -- - - -- - - --
Morum (Cancellomorum) of. M. chipolanum Maury I -- - - - - - - -- - - -- - --
Ficus carbasea micronematica (Brown and Pilsbry) -------------- -------------R P. P.- F R --- --Murex (Siratus) of. M. polynematicus Brown and Pilsbry I6 -- - -- - - - - - - -- - --
Mitrellar sp--------------------------------------------- ---- -P.----- ---- ---------- ---- ------ ---- ------P.
Strom bina of. S. quirosana H odson -- - - - - - -- - - - - - - -- - - - --
Cym atophos? of. C. veatchi (O lsson)-- - - - - - - -- - - -- - - -- - - -- - - --
Antillophos?' (A ntillophos?) sp. (sm all) - - - - - - - - - - - - - - - - -
Antillophos? (Antillophos?) of. A. candei gatunensis (Toula) -- - -- - -- -- - -- - --
N orthia? of. N northiae (G ray) - - - - - - - - - - - - - - - - -
Nassarius (Uzita?) praeambiguus (Brown and Pilsbry)-------- ---- ------ C------------------------- ---- C C
Melongena sp ---------------------------------------------P P. RP.-R -- -------------F a scio la ria sp - - - - - - - - - - - - - - - - - - - - - - - -
Fusinus? sp ------------------------------------------ ---R -- --- -- -- ------------------Oliva (Oliva) sp ---------------------------------------- ---- ------ ?F------ ------ -------- ------ ---- C-----Olivella sp------------------------------------------------- ------ ?F ---------- ---- ---- ------ ---- C-----A g a ro n ia sp - - - - - - - - - - - - - - - - - - - - - - - -
X ancus of. X validus (Sow erby) I - - - - - - - - - - - -- - - - -
Lyria? sp ----------------------------------------------------------------P. -- ----- -- -----Marginella (Eratoidea) sp ------------------------------ -- ---R --- --- ----- -----------------Prunum (Microspira) aff. P. apalachee (Gardner) --------------------?P. --------------- -----Prunum (Microspira) sp.......................................................................................----Persicula (Rabicea) venezuelana amydra W oodring, n. subsp-- -- -- - -- -- - -- -- -- -- -- --
C a n cella ria ? sp - - - - - - - - - - - - - - - - - - - - - - -
Conus of. C. planiceps H eilprin - - - - - - - - - - - - - - - - -
Conus aff. C. chipolanus D ali - - - - - - - - - - - - - - - -
Gemmula of. G. vaningeni (Brown and Pilsbry) -------------- ---- ------ ------ F --- --------------Pleuroliria (Polystira?) sp ----------------------------------- R---------- ---- ------ ---------------Pleurofusia? sp ------------------------------------------------ ---- R---- -------------------------C rassispira (Crassispirella) sp -- - - - - - - - - - - -- - - -- - - - -
See footnotes at end of table.









GASTROPODS: EULIMIDAR, MARGINELLIDAR TO HELMINTHOGLYPTIDAE 315


Cut area (Acmaeidae to Ellobiidae)

C, common; A, abundant]


Localities- Cont inued


100b 101 101a 101b 11 101~d 101c 101g 10111 1011 114 116 li5a 115b 116 116a 119 119a 120 126 130








---------- ------ ------ ------ ---- ---- ---- ------ ------ ---- ---- ------ ---- -----U --- -- --- ---R-------- ------ ------ ------ ---- ---- ---- ------ ------ ---- ---- ------ ---- ------ ---U-- --- ------------- ------ ------ ------ ---- ---- ---- A-------- ---- ----------- ----U U ----- --- ------------- ------ ------ ------ ---- ---- ----U--------- ---- ---- ------ ---- ------F--------- -----U ------ -------- ------ ------ ------ ---- ---- ---- C - - -- - -- - -- - -- - --
-- -------- ------ ------ ------ ---- ---- ---- ------ ------UR -- ----- --C --- ----- ----- -------- ------ ------ ------ ---- ---- ---- ------ ------ ---- ---- ------UR -- ------ ----- ------------------- ---- ---- ---- ------ ------ ---- -----?F------ -----U --- -- --- ----- ------------------------------- -------F ---F C ---F --- ----- ------------- --------------- ---- ---- ---- ------ ------ ---- ---- ------ ---- ------F ---------------- -- ------------------ ---- ---- ---- -U - - -- -- - -- - -- - -- - --
-- --------------- ------ ------ ---- ---- ----UR - -- - -- - - -- - -- - --
---------- ------- ---------- ---- ---------- --------- -U -- - -- - -- - -- - --

-- -------- ------ ------ ------ ---- ---- ----A---- -------------------- ------ U------ ----- ----- --------- ------ ------ ------ ---- ---- ----UA - -- - -- R R - -- - --

-- -------- ------ ------ ------ --- ---- ---- ------ ------ ---- ------- U-- F-- A------ ----- ------------- ------ ------ ------ ---- ------------------ ---- ---- ------ ---- ---- --F---- ------ ---- ------ ------ -------- -----------------------------------U
-- -------- ------------ ------ ---- ---- ---- ------ ------U F-- F-- U-------- C U--- U----- ------------- ------ ------ ------ ---- ---- ----U---- ------ -- -----s R A -----------------sp---sp.------ ------ ------U -- p--------- ----- ----- ------ ---- ---- ------ ---- ---- F U----- ---R----------- ------ ------ ------ ---- ------------------ ---- ---- ------ ---- ------U----- ----- ------------- ------ ------ ------ ---- ---- --- -C-- -------- ----U U U ---R ------------ -F-- U----- ------ --- ------ R --F- F ---- U F------ ---------- s-- --- Up- U---R--------------- R---s--------- --------------------U------ ------ ---- -----U R -- --- ----- -------- ------ ------ ------ ---- ---- U U--- - - -- -- - -- - R - -- - --
---------- ------ ------ ---------------- ------ ---C----------U-- R --- -- --- ---------------- ----------- ---- ---- -----U F R-- ---- ---R R --- -- - - -- - - -- -- - -- - ------- ---- ---- R---------- ------ ---- ---- ------ ------ ------ ------ ---- ---- ---- ----C- -- -- - - - - -- - -- - --
-- -------- ------ ------ ------ -------- U U1R - -- - -- - - -- - -- - --
---------- ------ ------ ------ -- ---- --------- -- -U U------ ---- --U---- ------ ----U-- ----- -------- ------ ------ ------ ---- --------- --------- -U- ----- ---- ------ ----- ------------- ------ ------ ------ ---- ------------------------------------ ---------- ---------------- ---- ----- -------- ------ ------ ------ ---- --------- ------------ U-- U----- ----- --- ----- ----- -------- ------ ------ ------ ---- --- --- R ---- -- U -- - -- - -- - -- - --
-- -------- ------ ------ ------ ---- --------- U U --R -- -- --- --R -- -------- ------ ----- ---- ----- ------ R-------------- ---------------- -------------- ----- -- ------ ------ ?- --- ------ ---- ---- ---- ----C- - R - - -- - - -- - -- - --
---------- ------ ------ ------ ----------?---- ------------- -----R---- -----U-- ----- --- -----?F- -------------------- ---- ------ C R-------------- ----------------F U--- ---------- -----? -------------------- ---- ------ R R---------- ---- ----------------F U--- ---------------------------------- ---- ---- ------ ------ ---- ---- R------------------------------ ---------------------- ------ ------ ---- ---- ---- ------ ------ ---- ---- ------ --------- -U-- ----- ------------------------------------- ----------------------- ---- --------------- -U----- --------- - -- -- - -- -- - -- -- -- ? -- -- --------- ---- ---- R------ ---- ------ -------------- ------ ----- ------ ------ ------ ---- ---- ------ ---- ---- ------ -------U--- ------ ----U A-- - ?F - -- -- - -- -- -- -- - -- - FR------ ---- ---- F---------- ------ ---- ----- ----?F ------- ------ ------ ---- ---- ---- -C--- --- F R ----------- U --- -- ----- --------------------- ---- ---- ---- R---------- ---- ---------- ---- F------ ------------------- --- - -- -- - -- -- -- -- -- -- - ------ ------ ---- ---- R---------- ------ --------

-- -- - - -- - - -- -- - -- - --- ---- ------ ---- ---- R------ ---------- ---- ---- - -- -- - -- -- - -- -- -- R - -- ------------- ---------- ------ ---------- ----------------------- ------------ ------------------ -----------------------------U --- ----- ----







316 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE

Gastro pods from La Boca formation of Gaillard


Clathrodrillia sp-------------------------------------Terebra (Oreoterebra) dicheres Woodring, n. sp -----------Strioterebrumi cf. S. clethra (Maury) ----------------------Acteocina sp----------------------------------------Scaphander (Scaphander) cryus Woodring, n. sp.? ----------Atysp --------------------------------------------Retusa? (Cytichnina?) sp.............................--Floribella aidrichi (Dali) -----------------------------Eltobium. aff. E. pellucens (Menke) ----------------------


Localities


98


I Described on page 101 as TUrritella (Torcule?) ameras. I Recorded on page 103 as Turritella (Torcula) altulra, subsp.
3 Described on page 105 as Turritella cf. '1'. subqrundifera. Recorded on page 81 as Trochita cf. T'. trachiformis (Born). Described on page 203 as Meorum ("Oniscidia") cf. AM. chipe anuin. 1)Described oni page 215 as Mlurex (Marex?) cf. 11. plnecmaticus. 7Described on page 286 as Xancus cf. X. rex Pilsbry and Johnson.


99a 99b


R


99c


99d


99e


99f


99g


99h 100 100a


-F


Ga stro pods from La Boca formation of Gaillard Cut area and
occurrence elsewhere and in other Canal Zone formations of
same or related species.- Con tinu ed from p). ;113


Species from La Boca formation


Murex (Siratus) of. M.
polynematicus Brown and
Pilsbry.5
Strombina of. S3. quirosana
Hodson.
Cymatophos? of. C. veatchi
(Olsson).


Anfillophos? (Antillophos?) of.
A. candei gatunensis
(Toula).
Northia? of. N. northiae
(Gray).
Nassarius (Uzita?) prae ambiguus (Brown and Pilsbry). Xancus of. X. validus
(Sowerby) .'

Prunum (Micros pira) aff. P.
apalachee (Gardner).
Persicula (Rabicea) venezuetana
amydra, Woodring, n. subsp.

Conus of. C. planiceps I-eilprirn

Conus aff. C. chipolanus Dall__Gemmula of. G. vaningeni
(Brown and Pilsbry).
Terebra (Oreoterebra) dicheres
Woodring, n. sp.
,Strioterebrum cf. S. clethra
(Maury).
Scaphander (Sea phander) cryus
Woodring?, n. sp.
Floribella aidrichi (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. north iae, H-olocene.

Also Gatun formation.

X. validus, early Miocene,
Dominican Republic. Also
Culebra formation.
P. apalachee, early Mio cone,
Florida.
P. venezuetana 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.
7'. 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, H-olocene.


I Described on p. 101 as Turritella (TerculaI) amaras.
2 Recorded on p. 104 as Turrilella (Terco is) allilira Conrad, subsp.
3 Described on p. 105 as Turritetia cf. T'. subqrundifera.
4 Described on p. 165 as Architctonica (Architectoi tea) cf. A. noil412 Roding. 5Described on p. 203 as Merum (" Oniscidia'') cf. )II. chipolaosnn. 6Described on p. 215 as Murex (Murex?') cf. Al. peiyuemnaticus.
7 Described on p. 286 as Xancus cf. X. rex.


Gas tro pods, Em perador limestone nlemb er.-The Einperador limestone mnem-ber yielded the gastropods tabulated below. Aside from species identified only at the generic level, Cam panile cf. C. herculeanus is the only one Rot in the preceding table. The Emnperatdor species and C. Zierede anus, from the early Miocene Anguilla form-ation of Anguilla, are the last American species of the genus, younger than any in Europe.

Gastro pods from Emperador limestone member of La Boca formation


Turritella of. T.
collazica Maury--Turritetla (Bactrospira?) amaras
Woodring ----Turritella (Bactrospira) altilira
Conrad, subsp-Cam panile of. C.
herculeanus (Cooke)Strombus sp-----Orthaulax sp ----Cypraca of. C. chilona
D all-- -
Ficus of. F. carbasea
micron ematica
(Brown and
Pilsbry) - -
Fusinus? sp---------Xancus sp ------


Localities


Gaillard cut area


117a 1117c


R


F


118




R


R


121


It


123


R


129a


Rt


Madden basin 71 73


R

R


R
R
R


R


Age.-The La Boca fossils have marked early Miocene affinities. They suggest correlation with the Chipola formation of Florida. Nevertheless, the Lat Boca is considered to be older than tile Chipola, chiefly on the grounds that Lepidocyclina, and Neinoeardiuin sur-


316






GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO I{ELMTNTI{OGLYPTIDAE31


Cut area (Acmaeidas to Ellobiidae) -Continued

Localities-Continued
100b 101 l0la 101b 10ic 101d l01c 101g 10111 l01 114 115 115a 115b 116 l16a 119 119a 120 125 130

-- - - -- - - -- - -- ------ R----- ---- ------ -------------- ------ ---- ------ ---- ---------- ------ ------ ---------- ---- ---- ------ ------ ---- ---- R ----------------------------- ------ ------ ------ ---- ---- ---- ------------ ---- ---- ------R -- sp.F --- ----- -------------- ------------------C------ ---- ------ ------ ---- ---------- ---- ---- ------------ ---- ------ ---- -------- ------------ --------------- ---- ---- ------ ------ ---- ----- R------ ---- R ----------------------- -------- ---- --------- ---- ---- R---- ---- ------ ------ ---- ------ ---- ---- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -


viv~ed in La Boca time. To be sure Lepidocyclina is unknown in southeastern United States later than Oligocene. Aside from the sur6ival of ANeiocarditin in the Tamipa limestone (Mansfield, 1937, p. 258; recorded as Protoca)rdia) the L Boca fossils show little resemnlblance to those of the Tampa, although they are thought to be of the same age. Ortiaidax. Globulaiia, Pacliyc2ronunimm?, and Floribella are extinct.
The smaller benthonic, foran-inifera from the La Boca indicate correlation with the early Miocene Sancesian stage of California (Blacut and Kicinpeli, 1969, p. 5). In addition to Lepidocyclina girandi frorn locality 101h, a collection from the east side of Las Cascadas Reach (locality 1011), 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 ni'ratiorensis. Those two species are the last representatives of the genus in the Canal Zone.
The large, high-domed echinoid, Echinolarnpas 8erzo rbis is conspicuous in the Emperaclor 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 amid Alhaj nela 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 H-ighway at locality 74, 1.5 kilometers south of Rio Chilibrillo bridge.


The formation is divided into a lower member and an upper member. Tihe lower member consists of the former calcareous sandstone member of the Caimito formation and includes the lens of sandy limestone formerly designated the Chulibrillo limestone member of thie Caimito (p. 32). The upper member is the former Aihaluela sandstone member of the Caimito. Thej~ 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:

Gastro pods from A ihajuela formation

Localities
Lower member Upper member

77 79 80 82 82a 85 85a 88 89 92 Turriteiia (Bactrespira)
oltilira aitilira Conrad I ---of. Ri --- f. R --------- ---- ------ ? C ?
Tiirritella gatuiieiisi8
Conrad?--------------- ------- ---- ------- R........................---Xernietid? ----------------------- ---- ----- --- --R------R-----Crucibulunmsp ----------- ------- ---- ------- ---- ---- --- ------i
Moalea sp ------------------- -----R ----------- ?R -?R ------Ficus carbasea carbasca
(Guppy)---------------- sp. R ---------- ---- ----- ---- ------ R----IDescribed on p. 102 as Turritella (Torctcla) altilira altitira.

Age.-The upper member contains the typical form of T'urritella altilira and the typical form of Ficu8 ear75asea-the earliest known occurrence of those forms, which are found in the Gatun formation. That the age of the entire Alhiajuela 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


'A


317








318 GEOLOGY AND PALEONTOLOGY OF CANAL ZON~E


Gastro pods from Gatun formation (Eulimidae, Mar ginellidae to

[R, rare; F, few;

Localities

Middle part
Lower part

Eastern area








Turritella abruple Spieker--------------------------------------------------------------- -- ---- -- -- -------I R CR --R
1'etaloconclius sculpturatus .11. C. Lea-----------------------------------------------------H -- -- F---------------------------Ileuisinus eineras Woodring, ni. op------------------------------------------------------ ----------- --- --IEulitia nobitis 6.*upy ------------------------------------------------------------- -- ----- -- I F R C R -- --- HEulina acuta Sowerby---------------------------------------------------------- ------ --- --- -- --- _--- -- ----- -- --- -- --- -- ---- ---- -- ----- --- ----Eulima sarisiforntis (Vilsbry and Johnson)-------------------------------------------- F--- --------F--- -
Balcis (Balcis) jacululum (Mauury)--------------------------------------------- ?------------- --_I H --- --------------------R
Balcis (J3alcis) liparea Woodritig, n. sp-------------------------------------------------- ------F--- F -------- R--------------Balcis (Balcis) alara Woodring, ni. sp -----------------------------------------------------------------------------------
Balcis (13dmi) celia Woodring, ii. op------------------------------------------------- --- _-- --- -- F it RH R R
Nise (Niso) nliesata Woodring, ni. sp-----------------------------------------------t 1-------- -F_ _I _- --- --- _--- ------- ---- -- R ---Golubraria obscure (Heavc) ------------------------------------------------------------------- _-------------------------------------_ ----- --------Plterynetus (Subplerynotus) textilis (Gabb)-----------------------------------------H---------------- --------_ _Typais ('1atityphis) eucleanus Woodring, 11. SI)-----------------------------------------_-- ------- -- --- --- -Volvarina leanaer (Brown and i'ilsliry)--------------------------------------------- -- ----- --- -- ---- --- -- -----C ?It ------ -A A- C
Prunure (Micrespira) rjelunense (Brown and Pilsbry)--------------------------------------3 ------- ----------------------- -----------Plersicule (R1abicea) c)utriana sleriyrra Woodriiig, n. subsp-------------------------------- ? -- ----------------------Gancellerie (Cancellaria) eneontoia Woodrinig, n. op --------------------------------- -- ----- -- ----- -- v------ ----
Gaeellaria (Gancellarie) all. C. oacneili Alansfiold ------------------------------ -- ----- H- -- - -- -- -- ------------(Jancellaria (Casecellaria) tepeiina Woodring, a1. sp -------------------------------------- ----- ----- -- -- - -- ---- -- -- --------- ----(Jencellaria (Cancellarie) epistomilera dariena 'Iou --------------------------------F U it - F It H A A F GC------ -- C __ C -- _---- ---?F__--?
(Janceltaria (Cancellaria) epiot omifera lipara Woodr ing, ni.srsbsp ----------------------- -- _--- -- ----- ----- --- --- ------- ----- -- ----_-----__-------Cancelluria (Gancellaria) epimela Woodrig, n1. op------------------------------------------------------ ---------- --- ----------__--__--___ __ -------_--Gancellarie (Pyruclie) cibarcole cibarrole Anderson ---------------------------------- --H HI F Ht F Ht A C F (3-_--- ---- -- -- -- -Cencellerie (Pyruclia) diadele Woodrin~g, 11isp ----------------------------------------- ----- --- ----------- -- ---- ----- --- -------------------Gancellarie (IEuclia) codazzii Anderson------------------------------------------- A C It H -_ R It A A C GC------ -- F -_ Ht R
Gancellaria (Luclia) dinole Woodring, ii. sp -------------------------------------- F C H- H HtR_ -A C-- C_--------------------------Gancellaria acalypte Woodrig, ii. sp----------------------------------H -------------------------------It------ ---------- --- -------------Cancellaria nattcellaria Woodririg, ni. p-------------------------------------------------- ------ --- ----- ------- --- --------- ----- ----- ---- ---- ----HR C
(Jancellerie (Narene) baryslome Woodring, n. op------------------------------------ -- -- H H Ht R te---- ------Cancelleria (N arona) decaplyx Brow n and I'ilsbry -- - - - - - - - - --- -- --- -- --- - - -- -- --- -- -- -- -- --
Cancelara (Chercollerie) terryi Olsson ------------------------------------------------ -- --- -- H--- --- CR -- -- ----- -- --- -- -- H
Aphera islacolonis (Maury) ------------------------------------------------------ -- ----- -- -- _It --H ---- ---_- -_- ---- -Trigonosloma cf. 2'. scalalelle (Guppy) ------------------------------------------------ --_-- --- _--- --- -- -I -- -- ------- ----'1'rirjo c.n'on u a e 1is y a d J hn o )- -- - -- - - -- - -- -- - -- -l- -- -- - - ---a- -f. _1' -nu-r --l r arid- Jo-nson
Genus receynilus Guppy -------------------------------------------------------------- -_--- - -- - ---- -- -- ------- -C ones m u s teflsio G lsson -- - - - - - - - - - - - - - - - -- -- -- - -- -- -- -- - -- -- - _ -- -- - -
Genus spurius Ginelin....................................................................................................................- --_ -- ----- ---Coitus brerel Spieker ---------------------------------------------------------- F C H- R(_-----c C R H - - -_- -Conas acolus Woodring, n. sp----------------------------------------------- ItF----------- _F------ --- ---- -- -- ------- ----Genus malis Brown and Pilsbry-------------------------------------------- H HCH----------- _R----R-_---H--- H-_----R--- ?H_ R
Coinus aemalator aeinulalor Brown and Pilsbry ----------------------------------- --H Ht H_ H _-i F RH---------H- -- ---- H-Conus cf. C.'catenalus Sowerby ---------------------------------------------------- --- --- -- --------- ----- -- ----- --- _--_ --- -Geitus consebrinus consobrinus Sowerby ------------------------------------------- -- ----- -- -------_--It---- _---------- ---H...........................-- -Genus symmetriCUS Sowerby --------------------------------------------------------- ---- -------- ---_--_--- -_- -- ---- -Coiunuslaphrus ri g, pW- -o----i- -g----n.-- --p-- --- -- --- --- -- ----------- -- ---------------- -- ------------------- -- ----------- ------Genus imitator imuiator Brown and Pilsbry--------------------------------------- --HR F F C Ht_ -- --H ---- -- -_- ----
Genus mulliliralus mulliltralus B bse------------------------------------------------------C H--------- _---_-CR --------C R -------- ---------------Genas burckhardli burckhardli Wiise-------------------------------------------------- -- --- -- ----- ----- --- --- ----- ----- ----- -_-- --H------- ?R _Gentus burcirlardli luarrisi Oisson-------------------------------------------------- ----- -- -- C RH---------- -_ ---- ----- -- -G en u s tortu osostrialu s T ou la -- - - - - - - - - - - - - - - -- -- -- - -- -- -- -- -- -- -- - -- -- -- -- -- -
Gemmiula vaninrieni (Brown and i'ilsbry)------------------------------------------------ ------ --- ---H----- t -- --- -- ----- ----- -----_---- ------ ------R _0Genmula nachapoorensis (Maury) ------------------------------------------------------ -- --- -------I ----__---------_---_---_--------Pleureliria (1'olyslira) Imnages (Gardner)--------------------------------------H------- H H -- ItR_ A C F F ------ -- C It C H H H C
P leu re liria (P elyslir p - - -- - - - - -- - - - - -- - - - --)- - - - - - --op- - _ - -
Plearofusia acre Woodrig, iin. - - - - - -- - - - - - - - - - - - - - - - - -p - - -
Pleuroifusia (Cruzilurricula) fusions fusions (Brown and Pilsbry)_----------------- -- -----HR C F - -A C H F H __-----It ---__-- --- __ F__ C HLeacosyriux xenica W oad(ring, n.s - - - - - -- - - - - - --.- - - - - o--p- - - - - - - - -
Gochlespira (Ancistrosyrinx) cedonulli (Reeve)------------------------------------------- ------ --- -- ----- ------ -- -----_--- --------- ------ -------- -- -.
,Scebinella seorier(i (Cossmanin) -------------------------------------------------------H -_R_------- ---- ---- -- -----Drillie (Needrillia) rioqnrabenis eurysemua Gardnier --------------------------------- _-- --- -- ----- -- R--- --- -- ----- F.................
Garinadrillia (Garinedrillia) zooki (BrownV and P1ilsiiry) ---------- ----_------__-- _----- R_--- ----- ---------------H H -- -- -- ----- R
Garinedrillia (('arinodrillia) cf. CJ. elocala (l'ilsbry andiohuisoi)..- --------_---------- -- -----_-- --- ------_--- -- ------- R
Gtassispoire (Grassispira) henc'keni leptalee W oodrig, ni. outiop-- ----- ----_ -- --- -- -- --------- -
Grassispira (Ilindoiclara) censors censors (Sowerby)----------------------------------- ------ -- (3_ F R H ------ -- F -- R------------- C R
G rassispirap r la)c m to o drlg t p - - -- - - - -- - - - -- - -- - - - - --assp r la cy--ie ---r g -. --Grassispoira (Grassispirella) lyloessa Woodrinig, ii. sp-------------------------------------------------------H
Glalbrodrillia ejalunensis (Toula) --------------------------------------------------------------_--- --C F H -F_ t I F -- R H --I-------- -------Glalhrodrillia seavedrai Woodrinig, n. op---------------------------------------------- ------ --- ------ --- ----- --- --- --- --- ------ -- --- ------- R









GASTROPODS: EULIMIDAE, MARGINELLIDAR TO HELMINTHOGLYPTJDAE Cavolinidae and a few species of other families)

C, commron; A, abundant]


Localities-Continued


Middle part-Continued


Upper part


Eastern area-Continued


Western area


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320 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Gastro pods from Gatun formation (Eulimidae, Mar ginellidae to


Localities

Middle part
Lower part
Eastern area




CS CS CS-C 0' t 0 CIL.


Clot hrodrillia aft. C saarc-drai W oodring, n. sp - - - - - - - - - - -- - - - - - - - - - - - - - - --
Clot hrodrilia aft. C. lelandi (Olsson).-------------------------------------------- -- R----Clathrodrillia cf. C. islalindae (Maury) ----------------------------------------- -- ----- -- ----- -- --- F --R ---- -- -- -- ------
Agladrillia (Agladrillia) characta IWoodrinig, 11. sp---------------------------------- -- ----- -- --?R F ?R --- --------Rt------- ---- -- -- ------- F
Agladrillie (Ag-ladrillia) enneacyma c-anc-acymea (Brown and Pilshry)----------------------- ------ --- ------ --- ----- I R---- ---------------C --A -- F
Agladrillia (Agladrillia) esineacyma (Brown and Pilsbry), subsp -------------------- -- -R -- R --It- -- -- --- -- -- -- ------
Agladrillie (Ayljadrilia) pltc-ngeid-s Woodrinig, n. sp ------------------------------- -- ----- -- ----- -- --- --- --- -- ----- -- --- -- --- -- ---- -- ---- F -- A- F
Agladrillia (Agledrilia) aarna -ct ypha Woodring, ii. suhsp-------------------------- -- --C - F C C F I-- ----- C C Rt R --- -- --- -:-
Agladrillia (Eumc-tadrillia) isthmica (Brown and I'ilshry)-------------------------- ---- F--- -F Rt R ----It------ -- -- ------- ---Agladrillia (Eumctadrillia) acidna W ooding, n1. sp ----- -------------- ---- ----
Microdrillie triva Mansfield----------------------------------------------------- -- ----- -- R---R--- --- -- ----- R F ------ -- ---- ---- --I R-- A-- C
L ee cyic ra heara o a Ghb )pt--- --- -- --- ---a-- --- -- --- --- --- --- -- --- ------------ ---R---- --- --- ----------- --- -- --- --------- -- ------ --- ----Ithycythara defuniac Gardner --------------------------------------------------- ------ ------ R---t --- ------ ----- ------- ---- ---- ------ ----Ithycythara cf. L. elongate (Gabh) --------------------------------------------------------------- ---I t - -- ---- -- -- ------
Cytharella? cf. C. ce-epsacosta (Gardner) ----------------------------------------------- ----- --- It- -- --- ------ -- -Kurtzi-lla (Kartzi-lla) payc-ila Woodrinig, n. sp -------------------------------------- I -- -- F C- IIt --C F------------ ---- CA
Kurtzic-lla (Kurlzic-ila) stenotella Woodring, 11. sp---------------------------------- -- ----- -- ----- -- --- --- --- -- ----- -- --- -- --- ---- ---- ----I R -- IR
Kurtzi-lla (Cr yet rris) ha bra Woodring, u. sp ------------------------------------- -- ----- -- -- R -- ---- -- -- -- ------Knrtzic-lle (Cr getUrriS) Sp ------------------------------------------------------ -- ----- -- ----- -- --- R-------------------------------Nannodi-ila rintriada (Mansfield)-----------------------------------------------------------------------------t--- --- ---------------- ----Nannodiella cf. N. melanitica (IDnll)--------------------------------------------- -- ----- -- ----- -- --- --- --- -- -----It
Itarlathor-lle (Etuclathur-lla) rc-ndrycsiana (1)all) - - - - - - - - - -- -- -- -- -- -- - -- -- -- -- -- -R -- -Euclathur-lla (Miraclathr-lla) cucharis Woodring, n. sp--------------------------- -- -----It F RI- C Rt--- -- ----- C C ------ ---- ---- -- --R C-- It
Dolostoma anorhepes Woodring, n. sp--------------------------------------------- I tI-------------RIt- F--- RRR--------------- --Glyphostome (Glyphostoma) dc-ntifc-rnm Gabbh------------------------------------------------ ? ------R ------------------------t--Glyphostome (Glyphostoma) pyrg/ota W oodrinig, n. sp - - - - - - - - -- -- -- -- -- -- -- -- -- -- -- -- -- --
Glyphostom a (I'i lyphostorna) oissossi W oodring, n. sp-- - - - - - - - -- --- - -- -- -- -- -- -- -- -- -- - -- -- -- -- -- -- --
Glyphostoma (Rhiglyphostoa-) allodapum Wooddli-g, n. sp -------------------------------------------- R--------------------R
IDaphn-lla pagera Woodrig, n. sp...........................................................................................................- -- ----- -- -- -Daphnella? sp------------------------------------------------------------------ -- ----- -Terebra (Oreoterebra) subsesicifera subsulcifera Brown and 1Pilshry------------------------ F R C Rt F -------- -I-It
Teretbre (Oreterebra) subsulcifera cembra Oisson ----------------------------------------C It ?R RIt --- ---- ---- -- ------
Terebra (Oreoterebra) iseerpeiti Maury ----------------------------------------------------------------F
Terebra (Parat-rc bra) aft. T. taurina (Lightfoot) - - - - - - - - - -- - - -- -- - - - - - - - - - - --
Terebra (Panaterebra) csscurrupi-ssis Olnomikado -------------------------------- -- F-- R C C-- A F-- C RI--t- F F -- -- RI --t -- -----
Strioferc-brum spiriferum (1)all) ------------------------------------------------- F C ?R -- C C It A A C A C ?R-R I----- -- -- ------------I
Strioterebrum ga gi(T o la ---o---f- ---n- -i--- -- -- ----a)-- -- -- -------- ----- -- ----- ------- ------- ----- ----- ---------- -- -------------Striotc-rc-brum geusapatam (Brown and Pilsbry)------------------------------------ -- -- C C -- --- --- --- -- ----- -- --- -- --- -- ---- -- -- -- R-----,Striot-rc-brum indocayapum Gisson------------------------------------------------ -- -- A C F A A It C F F--- F -- --- ----- ------ -------C
Strioterebrum orsignum eresic-num Olsson---------------------------------------- -- ----- -- ----- -- --- --- --- -- ----- -- C.-- F C
Strioterebrum oeignum hadrum Woodring, n. suhsp------------------------------- C A --I ---- - ---- ---- ---- -- ------
S triot -rc-b rn m sp - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - -
,Strioercbrunmonidum (Woodrig) ----------------------------------------------------- ---- ---- -- ---- -- -- -- ---------It It,Striotc-rcbrum aft. S. raptum (Gardner) --------------------------------------------------------R ------------------------------------It-Acteon (Acteoss) pounctostriatus (C. B. Adams)------------------------------------------- C------------ ----F--C--- ------------------Aeos (Lissacteon?) sp -------------------------------------------------------- -- ----- -- ----- -- --- --- --- -- ----- -- --- -- --- -- ---- ---- -- ----- F
Rictax~is oryze (Gabh)---------------------------------------------------------- -- ----- -- ----- --I R R -- -----IR F ------ --IR
Ringicala (Ringiculella) semistriata (l'Orhigny------------------------------------------F--F-R ------I--FFC--------C-----------------I-tC
Acteocina bullata (Kiener), small form------------------------------------------- -- ----- -- ----- -- --- --- --- -- ----- R---- -- --- -- ---- ---- -- -----R R-Acteocina clarhista Woodrig, n. ap---------------------------------------------- -- ----- -- F C ------ ---It R -- -------- -- --- -- ---- ---- -- ----- C
Acteocina ruse Gardner-------------------------------------------------------- -- ----- -- ----- --IRt--- --- -- -----IR
Cylichnelle atacate stibare Woodring, n. suhsp------------------------------------- -- --IR -- C A A C F -- ------N A RR It----C--------F
Rexania c/dpolana (Dali) ------------------------------------------------------- -- ----- -- ------- R--- --- -- ----- -- R---- ------ ---- -R--- A
Bulla m umbilicat gsm ll or -- -- -- --din--- -- --small- -- --for--- --R---- ------------ --- ---------- -----------------------------------A tys (A liculastrum ) eurys W oodring, n. sp - - - - - - - - - - - -- -- - - - - - - - - - - - - - --
Atys (Weink-auffia) cadus Woodrinig, ii. sp---------------------------------------- -- ----- -- ----- -- --- --- --- -- ----- F
Retusa (Cylichnina) quercissensis biforis Pilsbry and Johnson ----------------------- -- ----- -- ----- -- --- --- --- -- -----It
Sulcorets-s sulcata lipc-we (Woodig)------------------------------------------------ ------t--- R -- ------ ------ ---F Rt----- -- R----- -- C
Volvitella (Volvulelle) orrytata (Bush) -------------------------------------------- -- --IR -- F C C -- Rt -- R --- C FA
V olvulella (Volvulella) m icratracta B row n and l'ilshry -- - - - - - - - -- -- - -- - -- -- -- -- - --- -- - -- -- -- -- -- -- --

Volvulella fWolvulc-lla) jphoinicoid-s (Gardner)--------------------------------------- ------- --- ---R-- --- -- ----- C -- --- ---- ---- -- ----- ----Spiaella inflatlat (Colln)------------------------------------------------- ---Carolina (C'avolina) triaspis W oodring, n. sp - - - - - - - - - - - - - - - - - - - - - - - - - - -
Caroline (Caroline) cf. C. ventricosa (G uppy) - - - - - - - - - - - - -- - - - - - -- - - - - - -- -








GASTROPODS: EIJLIMIDAE, MARGINELLIDAR TO HELMINTHOGLYPTJDAE Cavolinidae and a few species af other farnilies)-Continued


Localities-Continued


Middle part-Continued


Eastern area-Continued


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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 gcenlus Aphera), 16 conids, .50 turrids (including two species of the new mnangeline genus Dolostoma, and a species each of Euglyphostoma and Rhiglypliostorna, new subgenera of the miangeline genus Glyphostomza), and 13 terebrids (including five large species and subspecies of Terebra, assigned to the subgenera Oreotere bra. Paraterebra, and Panatere bra).
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 deseribed). To that total should be added 5 that remain to be described and 10 undescribed pyram-idellids. For the purpose of talbulation 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.

Gastro pods from Gatun formation


Part


Low er - - - - -
M iddle-- - - -
Upper, eastern ------Upper, western -------


Number of species and subspecies



145 181 82 53


Largest collection


Number of species and subspecies

125
104 35 25


Locality


1 38c 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 Darh~n Province and Chiriqui Province, both on the Pacific side of Panamki, anid also with the m-iddle Miocenie 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, Strombinopho8, Strombiniella, Sub terynotus, Pavainurex, Pilsbrytyphis, Calop)/OS, Rhipophos, Nicemna, Nanariuts, Psilarius, Charcolleria, Scobin ella. and Dolostomna are extinct.
rraxa, 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 sub genera in gastro pod fauna of Gatun formation


Teinostoma (Aepystoma) Solariorbis (Hapalorbis) Heliacus (Astronacus) Rhinoclavis (Ochetoclava) Trochita
Neverita (Glossau lax) Neverita (Hypterita) Malea
Thais (Vascula) Strombina (Sincola) Solenosteira Metula
1 Described on p. 256 as 1-anctia.
2 Described on p. 283 as Tiara.

A few Ipaciphiles, listed
level:


Cymatophos Oliva (Strephonella) Mitra (Cancilla) I Cancellaria (Pyruclia) Canceltaria (Euclia) Cancellaria (Narona) Aphera
Pleurofusia (Cruziturricula) Agladrillia (Agladrillia) Glyphostoma (Euglyphostoma' Terebra (Panaterebra) Rictaxis



below, are at the specific


Paciphile species in gastro pod fauna of Gatun formation Trochita spirata Forbes? I Neverita (Glossaulax) reclusiana xena Woodring Neverita (Hypterita) helicoides Gray Distorsio (Rh ysema) decussata gatunensis Toula Cochiespira (Ancistrosyrinx) cedonulli (Reeve) Volvulella (Volvulella) cylindrica parallela (Pilsbry and Johnson) I Described on p. 81 as Trochita trochiforinis (Born).

Still fewer species survive on both sides of the Panamd. land bridge, as follows: Gastro pod species of Gatun formation that survive on both sides of Panamd land bridge
Architectonica (Architectonica) nobilis Rd6ding 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.

Gastropodts from Cha gres sandstonc at locality 208
(Canccllaridw to Tcrcbridw)

Canodilaria (Cancllaria) aff. C. epistomifera dariena Toula Cancilaria (Cancellaria) sp. Cancellaria (Pyruclia) diadcla Woodring, n. sp.? TJrigonostomia ii. sp.
Conus imitator Brown and Pilsbry? Conus tortuosostriatus Toula Plcuroliria (Potystira) ecuadoriana (Olsson)


322







GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO H-ELMINTHOGLYPTIDAE 323

Occurrence elsewhere of gastro pods from Gatun formation


Teinostoma (P8eudoretclla) pycnum (Woodring) --GycIOtTOescus (POnocyclUS) Pentagonus (Gabb)--Turritelta (Bactrospira) altilira altitira Conrad 5--Turritella (Bactrospira) altilira praecellens Pilsbry
and -Brow n 5- - - - - - - - -
Turritella abrupta Spieker....................---Turritella 7natarucena Hlodson-----------Turrilella gatunensis gatunensis Conrad ------Serpulorbis papulosus (Guppy) ----------IPetalocossehus scuiptur atus 1-1. C. Lea 6-------Alabtisa asperoides asperoides (Gab b) -------Alabina asperoides canaliculata (Gabb)-------Arelitectonica (Architectonica) nobilis nobilis Ro6disg- lieliacus (Astronacus) stonemnanae (Maury)-----Scalina pseudoleroyi (Maury) -------------------Eulimia nobilis Guppy----- ----------Eulimna acute Sowerby --------------Eulima sarassiformis (Pilsbry and Johnson)----Balcis (Balcis) jacululum (Maury)---------C'elyptraea centrali ((Conrad) -----------777ochita spirata Forbes? 7 -------- -----C'rucibulum (Crucibulumi) chlpolanumn Dali ----Crucibulum (Dispotaca) sprinqvaleense Rutscb --Crepidula plana Say------ -- -------Xefnophora delecta (Guppy)---------------------Strombus gatuflensis Toula-- -----------Siphocypraea (Muracypraea) henekeni (Sowerby) 8 Atlanta (Atlantidee) lisea Woodring --------Natica (Natica?) bous Browun and Pilsbry-----Sligmaulax ciuppiana (Toula) -----------Polinices brunneus subclausus (Sowerby)---------Polinices staniolIasmeunseri Maury ---------Neverita (1-yplerita) helicoides (Gray)............--Smnum euryhedra Woodring ------------Semicassis (Tylocassis) recluse (Guppy) ------Sconsia laevigata sublaevigata (Guppy)-------Colubraria obscure (Reeve.)------------Distorsio (Rh goema) decussata gatunensis Toula --Bursa (Colubrellina) caelata aniphdtrites Maury---Melee censure Guppy-- ------ -------Ficus carbasee cai asee (Guppy)----------Mjurex (Marex) recurvirootris recurcirostris Broderip Pterynetus (Subpterynotus) textilis (Gabb)-----Paziella (Panarnurex) gatunensis (Brown and
P ilsb ry) -- - - - - - - - - -
Eupicara thompsoni Woodring-----------Typhis (Talityphis) alatus obesa8 Gabb ------Mitrellehlimonensis (Gabb) ---------------------Anachis (Costoanachis) mnira mire (Dali)------Anachis (Costoanachis) mire fugax Brown and
P ilsb ry - - - - - - - - - -
Strombine (Strombina) lessepsiana Brown and
P ilsb ry - - - - - - - - - -
Strombina (Sincola) chiriquiensis Olsson__ -----Solenosteira dalli dalli Brown and Pilsbry --------Solenooteira dalli mnedioamericena Olsson 9 -----Cymatephos veatchi veatchi (Olsson) --------Cymatophos subsemnicostatus (Birown and Pilsbry) - Antillophos (Antillephos) candei gatunensis (Toula) Antillophos (Antillophos) mnexicanus (Btise) ----Nesariur- (Uzita) celcadensis (Maury)-------Psila? ins leptus (Woodring) 10 -----------Mfelongena snelongena conso? s (Sowerby) ----------Fasciolaria gorgasiana Brown and Pilsbry-----Fasciolaria yorgasiane Brown and 1'ilsbry, subsp -Olive (Strephonella) colpotus Woodring, n. name UAncille (Eburna) pinguis (Guppy) --------Olivelle (Niteoliva) terryi Olsson ----------Olivella (Toroliva) gjoliath Olsson----------Agaronia testacea mancinella (Glsson) -------Mitra (Pleioplygma?) limnonensis Olsson ------Mitra (Cancihla) longa lange Gabb 12......... Mitre (C'ancilla) dariensis Brown and Pilsbry 12 ... Xeocas falconensis Hlodson 13 -----------Veluta eurytera Woodring 14 .............
Prunum (Microspira) gatunense (Brown and Pilsbry) Can cellaria (Can cellaria) epoistomifere dariene Toula Cancellaria (Cencellerie) epistomzifera lipara Woodring, n. subsp -- - - - - - - - -
Cencellaria (Pyruclia) cibarcole cibarcole Anderson --Cencellerie (Enclia) codazzii Anderson-------Cencellerie (Charcolleria) terry Dlsson-------Aphere islacolonis (Maury) ------------See footnotes at end of table.


Gatun formation


'




x


x
x
x
x


x

x
x

x
x
x
x
x
x
x
x

x
x




x

x


x







x


x
x


'7


x
x


x

x


x


x


x


x


Early Miocene


S.
0
C)
0-C) C) '0 04-'
n's C.)


x


x


x


x


x


x


x


x
x


x













x


Middle Miocene


cd
0


0
WC


-->
x






x


-->
x

x
x


x


x

x

x
x

x
x


x


x
x


10 '0


x




x~>U

x x x
-->
x x x






x



x x


x
x


x


x


S. W
04




x x
- - - - -


x
x

x



x
-->


x
-->
x
x

x
x


-7
x





x


-->

x
x
x
x






-->
x
x

x


-->
x

x


x



x


-C3

00


.0


Late Miocene


S.
0g


.0


-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
x


x
x


x


-x
---------


x
x






-->




x
x


x


x


Early Pliocene


Late RooPlio- ceno cene


0
0


S.
0
.-C)
~
on
00


.0
0


x

x

x



x





x


x






x






-->


x


x


x








324 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE Occurrence elsewhere of gastro pods from Gatun formation-Continued


Gatun formation


04


0
0~ '0


Early Miocene


Middle Miocene


-- ______ 1_____ _______-______ ______________


a
0
0


C)0

0


Genus recoginitus Guppy--- - - - - - - - -- - --
Genus mnasaensis Olsson ------- ----------- X
Conu3 spurius Gmelin -------------------------- ------ x --Genus bravoi Spicker-----------------------------X x X --Genus molis Brown and Pilsbry -------------------X X x X
Conus aemulator aemulator Brown and Pilsbry ---X X --Genus consobrinus consobrinus Sowerby ------------- X x x
Genus symmetricus Sowerby-------------------- ----- X----Genus imitator imnitator Brown and Pilsbry ----------X X x X Genus multiliratus multiliratus B~se ----------------X X x x
Genus burckhtardti burckhardti Bo6se---------------- ------ X X
Gonus burckhardti harrisi Olsson------------------- X x
Gonus tortuosostriatus Toula---------------------- ------ X x
Gemimula vaningeni (Brown and P ilsbry)------------X X x X
Gemmula macoapoorensis (Maury) -----------------X ----Pleuroliria (Polystira) tenagos (G~ardner)-------------X X x x
Pleurofusia (Gruziturricula) Jo sinus fusinus (Brown
and Pilsbry) ----------------------------------X X x X
Cochiespira (Ancistrosyrinz) cedonulli (Reeve) ------- ------ X x
Scobinelta morierei (Cossmann)---------------------- ------ X
Dritlia (Neodrillia) riogurabonis eurysoma Gardner X X --Crassispira (Ilindsiclava) censor s conscrs (Sowerby) X X x Clathrodrillia gatunensis (Toula)-------------------X X x
Agladrittia (Agladrillia) acaria ectypha Woodring, n.
subsp -------------------------------------- X X --Microdrittia trina Mansfield -----------------------X X x X
Lepicythara heptagona (Gabb)--------------------- X x --Ithycytliara defuniak Gardner---------------------- X x .--Nannodielta rintriada (Mansfield)------------------ ------ X x
Euclathuretta (Euctathurella) vendryesiana (DalI) ----- X----Euctathoirella (Miractathuretta) eucharis Woodring,
n.sp ----------------------------------------- X X --Dotostoma anorhepes Woodring, n. sp---- ------------ X X --Otyphostoma (Glyphostoma) dentiferum Gabb ---- ----) X X --Terebra (Oreoterebra) subsulcifera subsulcifera Brown
and Pilsbry-----------------------------------X X x X
Terebra (Oreoterebra) subsulcifera cembra Olsson----X --- -Terebra (Oreoterebra) isaacpetiti Maury -------------- ------ X x
Terebra (Panaterebra) cuo currupiensis Oinomikado~ x X --Strioterebrum spiriferusn (lDall)--------------------X X x x
Strioterebrum wotfgangi (Toula)--------------------X X x X
Strioterebrum gausapatumn (Brown and Pilsbry) ---X X --Strioterebrusn indocayapum Olsson ------------------x X x
Strioterebrurn oresignum oresignum Olsson ----------- ------ X --Strioterebrum monidum (Woodring) ----------------- X X
Acteon (Acteon) punrtostriatus (C. B. Adams) --------X X --Rictaxis oryza (Gabb)---------------------------- X X --Ringicula (Ringicutetta) semiotriata d'Orbigny ---- ----X X x Acteocina buttata (Kiener)------------------------- ------ X X
Acteocina ruse Gardner--------------------------- X X --Cylichnetta atacata stibara Woodring, n. subsp --------X X X Roxania chipotana (lDall) -------------------------) X x X
Butta umbiticata Rilding, small form ---------------- X X --Retusa (Cytichnina) quercinensis biforis Pilsbry and
Johnson--------------------------------------- ------ X X
Sulcoretusa sulcata tipara (Woodring) --------------- X X --Ivolruletta (Volvuletta) ozytata (Bush)---------------- X X --Votruletta (Votrutetta) micratracta Brown and
Pilshry--------------------------------------- ------ X --Volvutetta (Volvutella) cytindrica parattela (Pilsbry
and Johnson)--------------------------------- ------ X
Votrutetta (Volvutelta) phoinicoides (Gardner) --------1 X x X
Spirateettainflat a eterata (Collins) ------------------X ----Vaginetta undutata (Gabb) ----------------------- ------ X


x













x





x
x


0

' 09-


x



x


I Early Miocene part of Uscai i shale, southeastern Costa Rica; llaitoa formation, Dominican Republic; Thomonde formation, hlaiti; Aymiamr6i limlestone, Puerto Rico; deposits of early Miocene age in Sinfi area, Colombia; Las Perdices shale, Colombia; outcrop La Rosa foirimation, Venezuela; Subihaja formation, Ecuador; lower part of Zorritos formation, I erf
2 Presumably Cercado or Gui abo formation, Dominican Republic (Gabb's stratigraphically unallocated species); Tub-ir formation, Colombhia; subsurface La Rosa formation, Venezuela; middle Miocene part of lirasso formation, Trinidad; San Jos6 calcareous silt member of Manzanill o formnation, Trinidad; Agueguexquite formation, M6x leo; middle Miocene deposits in Cbhiriqui P royince and I)arliti province, Panan am middle Miocene deposits in valley of Rio Saii Juan, southwestern Colonibia; Angostura, Picaderos, Progreso, and Daule formations, Ecuador; Upper part of Z/orritos formation, and Cardalitos and Montera formations, Perui.
3 Deposits of late Miocene age, Col10mb ia; P'un ta Gavilln form ationi, Venezuela; Savaneta glauconitic sandstone meiiber aiid Melajo clay mnember of Sprigvale formiation, Trinidad; deposits of late Miocene age (Spencer's Coatzacoalcos formation), Tehuantepec area, Mdxico; deposits of late Miocene age, including Olsson's Pinecrest


x


x




x
x


x










x


co





0



x


x




x
x




x


x


x



x


x


x


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x





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x


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0

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x
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x
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x


x


x


0l


'0


x
x


x


x


x

x


Late Miocene


0
x




x




x


x

x




x






x
x


0


x
x




x
x
x


x


Early Pliocene


0


0


x

x
x


Late Piocene


0


x


x


bieds and Mansfield's Cancettaria zone, Florida; 1)uplin marl, North Carolina; Estneraldas formation, Ecuador.
4 Deposits of early Pliocene age(, iloeas Isla id, l'anaintl; Mare and Play a Grande foriIations, Veniezuela; deposits of early I'l iocene. age iii Ciian9.i area, Venezuela; Matura formation, Trinidad; Charco Azul formation, southeastern Costa Rica and southiwesterni Paaniim.
5 Assigned to subgenus '1orcula on p. 102 and 105, respectively. G D escribed on p). 161 as Pet alocoomchas alt. P. floridanus.
7 D escribed on p). 81 as Troch ibm troch iforom is.
8 Described on p). 194 as Cgpraea (Ifuracypraea) henekeni.
9 Assigned to genus Ifaaeia on p). 256 ainl 257, respectively. 10 Described on p). 272 as Leptorias teptits.
11 Described on p). 278 as Oliva (S'trephoivtta) plicata. 12 Assigned to subgenus TJiara o p). 2S3 and 284, respectively.
03 Described on p). 286 as Xacus vatidosfatcoeniisi.
ii Described oil p. 287 as Volitta alfaroi otrylera.


324







GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE


Plcurofusia cf. P. fenimorci (Bartsch) Scobincila ecuadoriana Gisson Carinodrillia (Carinodrittia) sp. Darbya? (Buridrillia?) sp.
Grassispira (Hindsiclava) pyrgoma Woodring, n. sp. Cla throdritlia? sp.
Strioterebrum sp.

The gastro pod 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 endem-ic and four others evidently are. The occurrence elsewhere and in the Gatun formation of the same or related species is tabulated below.

Gastro pods 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.

Epitoniurn ("Dc pressiscala")
eucteanum Woodring.
Sthenorytis toroensis cuthynta
Woodring.
Scalina weigandi (Bi~se)---Bath ygalea (Miogalea) hadra
Woodring and Olsson.
Stigma lax guppiana (Toula) --

Distorsio (Rh ysema) decussata
gatunensis Toula.
Ficus carbasea carbasea
(Guppy).Solenosteira dalli Brown and
Pilsbry, subsp.2

Cymatophos? acolus Woodring-Amarophos bothrus Woodring___.

Latirus (Polygona) anapetes
Woodring.
Voluta eurytera Woodrino 3


Cancellaria (Cancellaria) aff.
C. epistomif era dariena
Toula.
Cancellaria (Pyruclia) diadela
Woodring, ni. sp.?
Conus imitator Brown and
Pilsbry?
Conus tortuosostriatus Toula

Pleuroliria (Polystira)
ecuadoriana (Olsson).
Pleurofusia cf. P. fenimorci
(Bartsch).
,Scobinella ecuadoriana Olsson-Crassispira (Hlinds iclava)
pyrgoma Woodring-, n. sp~.


Occurrence elsewhere and in Gatun formation of same or related species

C. aurora Dali, liolocene.

Middle Miocene, Ecuador;
late Miocene, Venezuela,
Panamnti, M6xico.
E. scipio (Dali). Hiolocene.

S. toroensis toroensis (D~ali),
Toro limestone member. Late Miocene, Tehuantepec
area, M6xico.
B. andersoni (Abbott),' early
Miocene, Colombia.
MIiddle to late M\iocene. Also
Gatun formation.
Middle to late(?) Miocene.
Also Gatun formation.
Early to late Mioeene. Also
Gatun formation.
Subspecies of S. dalli range
from middle to late Miocene.

A. dentalis (Olsson), early
Mioeene, Costa Rica.
L. taurus Olsson, late Miocene, Panami.
V. alfaroi Dall, middle Miocene, Costa Rica. Also
Gatun formation.
C. epistomif era 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, Pananit,
Ecu ad or.
P. fenimorei, Hlolocene.

Late Mioeene, Ecuiador. C. consors (Sowerby), early
Miocene to late Pliocene.


Recorded on 1). 198 as Bathyjelcea dalli. 2Assigned to genus 1-lanelia on 1). 258. 3Described on p. 287 em Veluta alfaroi eyrytcra.


Though the Chiagres 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 ae range of none, except a related species, includes Pliocene. So far as known, however, three species are closely related only to living species. Mliogalea, Amarop/io8. and Scobiiiela are extinct.
Two of the Chagres tnrrids 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 suggrestingy a correlation between the Chagres and Esmeraldas a comparable moderately deep-water facies rather than age is being correlated. Nevertheless both formations overlie middle Miocene deposits. In three widely separated areas-the Tehuantepec region of Mexico, northern Panamil, aiid northern Ecuador-moderately deep-water formations seem to be of late Miocene age.
Only eight gastropods have been recovered from the TrIo limestone member, the shiallow-water basal part of the Chiagres. Six, represented by molds, are aragonite-shieled, one of which is identified as Tiwrritella alidv a. The other two are calcite-shelled epitonids: Sthevoi~ytis pernobilWs (a living species) and S. toroeinsis 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 onl page 50. The following radiocarb~on dates on black organic miuck and wood were measured in the TTSGS laboratory: Radiocarbon dates on muck and wood from Atlantic and Pacific muck


Lab. No. W-959







W-960


Location of sample


Core 1-lole BBR 53, west side of Pacific
entrance to Panama Canal, 9 kilometers west-northwest of Point Farf an; depth 10 meters, 4.2 meters below top of mnuck, which is 9.9 meters thick.
Black organic muick.
Core Hlole BBR 128, east side of Pacific
entrance to Panama Canal, 0.5 kilometer east of terminuls of Thatcher Ferry; depth 22 meters, 9 mneter's below top of muck, which is 10.5 meters
thick. Black organic muck.
Core ilole Mindi 2, Milidi DIairy Farm,
6.4 kilometers south of Col6n; depth 10.6 meters, 1.5 meters below top of mnuck, which is 3 meters thick. Wood.


Age (years before present) 6, 720 300 7, 680 300 7, 240 300


325






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.
Dali (1912) and Brown and Pilsbry (1913a) described a few species of mollusks from marine layers in the muck near Mount H-ope on the Caribbean side of the Canal. Three of them are now known to be living off Payardi Island, 8 kilometers northeast of Col6n: ll inioliva myrrnecoon (Dall), Str b-oferebpurn spei (Brown and Pilsbry), and Ad'rana perprot'racta (Dali). 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 Chiagres, sloping northwvestward 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, Spillway, and Loecs described the bedrock surface as follows: "The rock surface under the Gatun Locks was vecry unievenl 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 MOLLUSKSCONTINUED FROM CHAPTER C
GASTROPODS-CONTINUED FROM CHAPTER C
Family EULIMIDAE
The family EulimidT. was omitted in its proper place.

Genus Eulima Risso
Risso, Histoire naturelle des principales productions de l'Europe
m~ridionale, v. 4, 1). 123, 1826.
Type (logotype, Herrinarnsen, indicis generum malacozoorum,
v. 1, p. 431, 1847) : Turbo subulatits Donovan (=Strombiforrnis glaber da Costa), living, eastern Atlantic Ocean.
Eulima cf. E. jacksonensis Gregorio Plate 48, figure 8
Very small, slender, 9-whorleci. Protoconch acte, tip slightly bulging. Post protoconcli whorls practically flat-si decd. Suture indistinct. Aperture elongate, narrow, ovate.


Height (incomplete) 3.1 mm (estimated restored height 4 mm), diameter 1.1 mm (figured specimen).
rThe marine member of the Bl~oilo(?) formation on Trinidad Island yielded five small specimens of Eulirna. They resemble E. jacksonensis Gregorio (Palm-er ;n H-arris and Palmer, 1946-47, p. 224, pl. 26, fig. 16, 1947; middle and late Eocene, southeastern tV. S.), but are about half as large, even when specimens having the same number of whorls are compared.
Occurrence: Marine mem-ber of Bohio (?) formation (late Eocene or early Oligocene), locality 42.

Eulinia nobilis Guppy
Plate 49, figure 1
Eulima (Liostraca) nobilis Guppy, in Guppy and Dali, U.S.
Natl. Mus. Proc., v. 19, p. 315, p1. 30, fig. 9, 1896 (Miocene, Jamaica).
M1elancila (Eulim a) cecadica Maury, Bull. Am. Paleontology,
v. 5, no. 29, pl. 25, fig. 1, 1917 (Miocene, Dominican Republic).
Strornbiformis prailubrica Pilsbry and Johnson, Acad. Nat. Sdi.
Phila. Proc., v. 69, p. 183, 1917 (Miocene, Dominican Republic). Pilsbry, Idem, v. 73, p. 395, p1. 35, fig. 10),
1922 (Mliocene, Dominican Republic).
Strornbiforntis ischna Gardner, U.S. Geol. Survey Prof. Paper
142, p. 573, p1. 55, fig. 17 (misprinted 16), 1947 (Miocene,
Fla.).
Small, slender, 11- or 12-wvhorled. Protoconch acute, tip sl](i gtl y bulging. Post-protoconch whorls practically flat-sided. Suture indistinct. Aperture elongate, narrow, ovate.
H-eight 5.9 mm, diameter 1.7 mim (figured specimen).
rType: Lectotype, herewith designated, USNM 107071.
Type locality: Blowden, Jamaica, Bowden formation.
E ulhna imobilis, represented by 35 specimens, is fairly wdly distributed in the lower and iddle parts of the Gatun formation, but is nowhere abundant. The Gatun shells are smaller than those from Jamaica, the Dom-inican Republic, and Costa Rica, which reach a height, of 9.5, 10, and 8.5 millimeters, respectively.
rTlie type lot of Guppy's mnisnamed species consists of four specimens. The lectotype is the largest and was illstated. Ma(ury's synionoiny and discussion indicate that she intended ileclanclla cercadwca to be a substitute name for Gabb's LUirna acicida)i, a homonym, although she did not, have access to Gabb's type. Nevertheless, her exp~ressionI "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 p)roperly pr )1oposed substitute name for01 Gabb's species was published. it is not the same as Maury's. (See the second following species.)


326






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE32


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, 147b5 14Th, 153a, 155 (identification doubtful), 159d. Cercado and Gurabo (USGS 8735) formations (middle Miocene), Dominican Republic. Bowden formation (middle Miocene), Jamaica. Middle Miocene deposits, Costa Rica (USGS 5882f and other localities). Shoal River formation (middle Miocene), Florida.
Eulimia acuta Sowerby
Plate 51, figure 13
Eutima acuta Sowerby, Zool. Soc. London Proc., p. 8, 1834
(living, eastern Pacific Ocean).
Leiostraca acuta (Sowerby), Sowerby, Thesaurus conchylioruin,
p. 803, p1. 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, p1. 47, fig. 2 (enlargement of Sowerby's
illustration), 1917 (Sowerby's record).
Eulima scotti Maury, Bull. Am. Paleontology, v. 4, no. 21, p. 30,
p1. 7, fig. 21, 1910 (Miocene, Florida).
Stromiforrnis scotti (Maury), Gardner, U.S. Geol. Survey Prof.
Paper 142, p. 573, p1. 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, p1. 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. ata has been used, for example by Dall 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 Republic. Middle Miocene deposits, Costa Rica (USGS 5882 in). Deposits of late Miocene age, Florida. Caloosahatchee formation (Pliocene), Florida. Living, North Carolina to West Indies; Pacific coast of Panaink.

Eulimia sarissiformis (Pilsbry and Johnson) Plate 49, figure 2
Eutima acicularis Gabb, Am. Philos. Soc. Trans., n. ser., v. 15,
p. 227, 1873 (Miocene, Dominican Republic). Not Eu~ima
acicularis A. Adams, 1861.
Stronmhiformis sarissiformis Pilsbry and Johnson, Acad. Nat.
Sdi. Phila. Proc., v. 69, p. 183, 1917 (Miocene, Dominican Republic) ;new name for Eulima acicutaris Gabb. Pilsbry, Idem, v. 73, p. 394, p1. 35, fig. 9, 1922 (Miocene,
Dominican Republic).
Relatively large, slender. Protoconch and early postprotocoiicl whorls m-issing. Remaining post-protoconch whorls practically flat-sided. Suture distinct or moderately distinct. Aperture elongate, narrow ovate.
Height (incomplete) 11.5 mmn (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 f rom the lower part of the Gatun formation. The protoconch and early post-protoconch whorls are issuing 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, pl. 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 Re.public.
Genus Balcis, Leach
Leach, Annals and Mag. Nat. History, v. 20, p. 271, 1847. Type (monotype) : Balois rnontagui 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.
Subgenus Balcis s.s.
Balcis (Balcis) jacululum (Maury) Plate 51, figures 1, 2
Melancila (Eulima) jacululurn Maury, Bull. Am. Paleontology,
v. 5, no. 29, p. 143, p1. 25, fig. 3, 1917 (Miocene, Dominican Republic).


368-278 0 70 3


327






328 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Eulima rotusta Gabb, Am. Philos. Soc. Trans., n. ser., v. 15,
p). 227, 1873 (Mliocene, Dominican Republic). Not Eulima
rob usta A. Adams, 1861.
Hlclanclla asfuta Pilsbry and Johnson, Acad. Nat. Sdi. Philadelphia Proc., v. 69, p). 182, 1917 (Miocene, Dominican Republic) ;new name for Eulirna robusta Gabb. Pilsbry, Idem, v. 73, p). 394, p1. 35, fig. 7, 1922 (Miocene, Dominican Republic).
Small, slender, about 8-whorled. Protoconch missing. 1Post-protoconcu whlorls lpracticallIy flat-sided. Suture fairly distinct. Body whorl rounded. Penult whorl practically flat or faintly angulated just above bodywhorl suture. Aperture short, ovate.
H-eight (not quite complete) 3.3 mmni, diameter 1.1 mmi (figured specimen).
Type: Cornell University.
Type locality: Bluff 3, Cercado de Mao, Rio Mao, Dom-inican Republic, Ce ucado formation.
Six small, slender fossils in five lower and middle Gatun collections are identified as Balcis Jacuuun. TIheyT are considerably smaller than topotypes of that species, which reach a height of .5.5 mim. Maury emphasized the "overbianging whlorls," that is, the faint angulation of the peniult whorl just above the bodywhorl suture. TFI a t feature, however, is iiot 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. maoi'ca (Maury, 1917, p. 142, pl. 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), 1471), 159d. Cercado formation (middle Miocene), Dom-inican 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- p rotoconcl whorls slightly bulgoing, remainder practically flat-sided. Suture (distinict. Aperture short, ovate.
H-eight (not quite complete) 8.3 mmii, diameter 2.5 mmi (type).
Type: TTSNM (46076.
Type locality 138 (UJSGS 16909, North and south sidles of Transisthian H-ighway, 1.6 kilometers northeast of Canal Zone boundary, Panamd), lower part of Gatun formation.
Baicis li]para is represented by 13 specimens from the lower aud idle parts of the Gatun formation.


It. is characterized by its faintly concave spire and slightly bulging last two whorls. Those features distinguish it from B3. jamaiceensis (C. B1. Adams) (Clench and Turner, 1050, p. 296, pl. 36, fig. 5; living, Jamaica) and a similar species in USNM collections from localities betwNNeen Massachusetts and F~loridla. rThis Atlantic coast species seemns to be the same as that named B. 1bai't.scb (Gardner and Aldrich) (1919, p. 37, pl. 2, fig. 1) for late Miocene fossils and R. callcim~i (Qisson and Harbison) (1953, p. 332, pl. 59, fig. 5) for Pliocene fossils.
Occurrence: Lower and middle parts of Gatun formiation middlee MTilocene). 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 lpost-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 mm11 (type).
Type: UTSNM 646077.
Type locality: 138 (see preceding species), lower part of Gatuni formation.
Th'lough time outer lip) of the type the only specimnen-is defective, Balcis aidaca is a distinctive species. In suiture and level ing adjoining the suture it is similar to time next species, but the shell is smaller and more slender, and the whiorl height is greater.
Occurrence: Lower part of Gatun formation (middle Miocene), locality 138.

Balcis (Balcis) cetia Woodring, n. sp.
Plate 49, figure S

Very large, sli mder, about 18 -whorled, whorl height 10wN. 1Protoconch missing. Post -protoconch whorls slightly bulging. Sutuire deeply impressed. Base of all except first few spire whorls beveled adjoining suture. Ap~erture short, ovate. Columiellar lip thick.
H-eight (not quite complete) 28.3 mm, diameter 8.7 mm11 (type).
T~ype: TTSNM 646078.
Type localitY: 1'Sf' (UTSGS 23663, South side of Transisthiian 1Inay1~, hillIside excavation at Colchoneri a Yero, about 450 meters southwest of.Cativa, Panami) lower part of Gatun formation.


328






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO IIELMINTHOGLYPTIDAE


Fourteen specimens of this large Niso-like species were found in the lower and juiddle parts of the Gatun formnation. 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. rTle earliest and sm-allest (height 20 mmn) of these species occurs in the Thom-oncle formation of Haiti (USGS 9946), the last and largest in the Savaneta, glauconit ic sandstone miem-ber and Melaj o clay miemiber of the Springvale formation of Trinidad, bo0th of late Miocene age: Balcis egiregia (Guppy) (Jung, 1969, p. 465, pl. 47, figs. 1, 2; height 50 mmn). The fourth species, B. niaki.ta (Gardner) (1926-47, p. 574, pl. 72, figs. 6-8, 1947), occurs in the middle Miocene of Florida. Like B. cetia, it is of intermediate size (estimiatedi height 93 1111. It is owvr moreslne than the Gatun species and the, base of its spire whorls is not bevelled.
B. dalli (B~artsch) (191'7, p. 302, pl. 35, fig. 5; Gulf of California) is a large species (height 20 mim), but its outline, suiture, and whorl height are like those of smaller species.
Occurrence: Lower and middle parts of Gatun foriation (iddle 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 1'Europe
meridionale, v. 4, 1). 218, 1826.
Type (monotype) :Niso cburnca Risso, Pliocene, France and
Italy.
Subgenuis Niso s.s.
Niso (Niso) unibilicata (Lea)?~
Plate 48, figure 7
Of miediumii size, moderately inflated, about 10whorled. Protoconch mssing. Spire whorls low, practically fiat-sided to slightly bulging. Body whorl roumndled to faintly angulateh. U~mbilicus partly exlposed, apparently narrow. Peristomec missing.
H-eight (incomplete 11.2 1111n (estimated restored height 15 mmn), diameter 5.5 mmn.
rile marine member of the Bohio (?) formation ait the submerged Vamnos Vamios locality and Palenquilla Point yielded eight specimens doubtfully identified as Niso urn bliwata. 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 (Palm-er, 1937, p. 66, p1. 6, figs. 22-25; Palmer in Harris and Palmner, 1.946-47, p. 225, pl. 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, la ter~ whorls practically flat-sided, last few slightly bulging. Intermediate whorls weakly
anguatedat base adjoiingr suture, angulation fainter or absent on last few whorls, including corresponding p~art of body whorl. Occasional microscopic axial threads on some specimens. Umbilicus moderately wide, bordering ridge subdued. Basal lip missing.
H-eight (incomplete) 14 mmn (estimated restored height 18 inun) diameter (incomplete) 6.4 mm (type). Height (not quite complete) 21.1 mm, diameter (not quite complete) 8 mm (paratype).
Type: USNM 646079; pairatype USNM 646080.
rType locality: 138c (USGS 21956, About 100 meters north of r1aisisthl~a1 H-ighiway and about 75 meters west of road to refinery site on Payardi Island, Panama.d; 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, ltud902'Nlongitude 790 52' WI., plus 1,010 feet [310 meters], middle part of Gatun formation.
Thug 61 specimens of Niso am esata are in collections from the three parts of the Gatun formation, 80 percent are mature, including all from-1 six localities. The early and intermediate whorls of the paratypethe largest specimen-are worn. The type shows the
anuation of inemdaewhorls.
Several similar species, differing chiefly in apical angle,0 are found in the Miocene of the Caribbean region. N. qr~andis Gabb (Pilsbry, 1922, p. 394, pl. 34, figf. 17 ; 1)ominican Republic) and its probable synionym, N. striatula Bbse (it Bose and rrolla, 1910, p. 227, pl. 12, fig. 7 ; Tehuantepec area) have the widest alpical angle, and a species from the Cercado farmation of the Domninican Republic, (USGS 8534) the narrowest. (rile type, of A. grandis is a fragment of four whorls. A lO-whorled spechinen, lacking a few early whorls (height 23.8 inin) has been found in the Gurabo formation at UTSGS locality 8519.) The apical angle of N. vesata is of intermediate width; wider, howeethan that, of a smaller species from the Bowden formation of JTamaica and the Lim6n formation of the Bocas (del. Toro area.
A'. mesata is more closely related to the Panamic N. excolpa Bartsch (1917, p. 3048, pl. 48, fig. 4) than to


329







GEOLOGY AND PALEONTOLOGY OF 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 MYARGINELLIDAE
Unlike most of the families of gastropods under consideration, the family Marginellidve has a richer representation in the Carribbean Sea than in eastern Pacific waters.
In addition to the species described or mentioned, unidentified m arginellids, or probable marginellids, are in collections from the Caimito and Culebra formnations.
Genus Marginella Lamarck
Larnarck,!Soc. Histoire Nat. Paris M(Sm., p. 70, 1799. Type (monotype) :Voluta glaticlla Linn6, living, west Africa.
Subgenus Eratoidea Weinkauff
Weinkauff, Systematischcs Conchylien-Cabinet von Martini uand
Chemnitz, v. 5, pt. 4, p. 140, 1879.
Type (logotype, Cossmann, Essais pal~oconchologie compare&,
pt. 3, p. 87, 1899) :Margincila 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 seTata Gaskoin, living, Maurit ius) 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 Eburnospira (Olsson and Harbison, 1953, p. 201; type (orthotype) : Marginella eburneola Conrad, Miocene, Virginia, and North Carolina) is available.
Marginella (Eratoidea) aff. X. mollitor Dall
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 mmn (figured specimen). Estimated restored height 11 mm, diameter 6 mmn (largest specimen).
Three damaged specimens are referred to this species: one from the upper part of the Bohio forma-


tion 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 Dali (1915, p. 52, pl. 12, fig. 1; Tampa formation) and ff[. eitancycla Gardner (1926-47, p. 388, pl. 46, fig. 20, 1938; Chipola formation). M. hernatita Kiener (1834-41, p. 11, pl. 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 Panamd 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) :Marginclia nitida Hinds, living, locality unknown.
Hinds' statement that Mfarginella 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). Mar ginela nitidla was based on a specimen (or specimens) of unknown habitat in the Cuming collection. Sowerby's (1847, p. 389, pl. 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, V~olvarina 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 columnellar 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 (Tomalin, 1917, p. 288; Woodring, 1928, p. 237), Schumacher'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


330







GASTROPODS: ETJLIMIDAE, MARGIINELLIDAE TO HELMINTHOGLYPTIDAE 3


smaller than his shell, but is much better depicted than by an illustration in a publication by Schr6ter.
Volvarina leander (Brown and Pilsbry) Plate 51, figures 4-6
Margincila lean der Brown and Pilsbry, Acad. Nat Sci. Phila.
Proc., v. 63, 1p. 347, p1. 24, fig. 13, 1911 (Miocene, Canal Zone). Gisson, Bull. Am. Paleontology, v. 9, no. 39, p.
98, p1. 6, fig. 22, 1922 (Miocene, Canal Zone).
Marginella aff. nit ida Hinds, Toula, K. k. Geol. Reichsanstalt
Jahrb., v. 61, p. 504, p1. 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 mmn (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 (pl. 51, fig. 5). The largest (pl. 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 H. nitida is a typical example of Volvarina leander.
V. collina (Olsson) (1922, p. 97, pl. 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 V7. avena (Valenciennes) (in Kiener, 1834-41, p. 17, pl. 6, fig. 24, 1834) is a little larger than V. collina, has a higher spire, and lacks the sharply bordered outer lip of V.* collina and V. leander. V. avena has been recognized in deposits of late Miocene age in Costa Rica (Olsson, 1922, p. 97, pl. 7, figs. 21, 28). V. tceniolata Mdrch ('Coan and Roth, 1966, p. 287, p1. 50, figs. 48-55), which ranges from southern California to Costa Rica,


has the outline of V.* 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, 177b, 177c, 177d, 178 (identification doubtful) ; western area, localities 183, 185 (identification doubtful).

Genus Prunum Herrniannsen
Herrmannsen, Indicis generum malacozoorum, v. 2, supplement,
p. 113, 1852.
Type (monotype and tautotype) : TVoluta prunum Gmelin, living, West Indies.
At the present time it is appropriate to use Pi-unum at the generic level for a large group of maginellids, the largest group of fossil species in the Caribbean region. The species assigned to Prunmm 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 Pr-unum 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 31icrospira Conrad
Conrad, Am. Jour. Conchology, v. 4, p. 66, 1868. Type (monotype) :Volutella (Microspira) oviformis Conrad,
Miocene, Virginia.
The type specimen of the type species was illustrated by Gardner (1926-47, p. 397, pl. 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 Joussea-une = Mar givella ainygdala 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.


331







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 narrowV, its inner edge smooth. Parietal callus thin.
H-eight 10.2 m-m, diameter 5.9 nun (figured specimnen).
The La Boca form-ation at locality 116,a near Paraiso yielded a small specimen of Prun urn. Though it is not well preserv~ed, it evidently represents a small species allied to P. apalachee (Gardner) (1926-47, p. 395, pl. 47, fig. 5, 1938), of the early Miocene Chipola formation of Florida. The spire of the La Boca species i* a little lower and its outer lip is narrower.
P. aff. P. apalach-ee is doubtfully recognized at locality 99b.
Occurrence: La Boca formation (early Miocene), localities 991) (identification doubtful), 116a.
Prununi (Microspira) gatunense (Brown and Pilsbry)
Plate 51, figures 10, 11
jlfarginclia qatuncnsis Brown and Pilsbry, Acad. Nat. Sci.
Pbila. Proc., v. 63, 1). 347, pl. 24, fig. 10, 1911 (Miocene,
Canal Zone).
HarginclIa coniformis Sowerby, Brown and Pilsbry, Idem, p.
348, 1)1. 24, fig. 12, 1911 (Miocene, Canal Zone).
Alargincla (JBulata) mindiensis Cossrnann, Jour. Conchyliologie, v. 61, p). 61, pl. 5, figs. 13-15, 1913 (Miocene, Canal
Zone).
11(1rfincllct mindicnsis Cossmann, Olsson, Bull. Am. Paleontology, v. 9, no. 39, p). 96, pl. 6, figs. 16, 23, 1922 (Miocene,
Canal Zone).
Mla r!in ella latissima pilsbryi Olsson, Idem, 1). 96, pl. 10, fig. 2,
not fig. 1, 1922 (Miocene, Canal Zone, Costa Rica).
Of miediumi size, moderately to strongly inflated, spire extendling well above posterior end of outer lip. Aperture very narrow. Outer lip strongly thickened, its inner edlge, weakly denticulate. Callus extending from outer lip over much, or exceptionally all, of ventral face of spire, thickly covering parietal wall and thinmung somewhat on columiellar lip. Outer edge of callus generally fading out on parietal wall and columellar lip, exceptionally sharply defined.
H-eight 17.8 mmii, diameter 12 mmin (larger figured specimen).
T~ype: Acadl. Nat. Sci. Phila. 1706.
Type locality: Gatun Locks excavation, Canal Zone, middle part of Gatun -formation.
Unfortunately the type of Jfarginella gatunensis is imm-rature (height 11.4 mun), whereas the type of ill. mnindiegsis is mature.
Thou0~gh Prunum. gatunevse is of modest size, it is the largest mnarginellid in tihe 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. nmindiense and Qisson's illustrated specimens referred to that species are moderately inflated, like the immature type of P. qa/une)?se and the immiiature shell shown on plate 51, figure 10. Brown and Pilsbry's P. conifonnis is mature andl inflated. Olsson's illustrated P. latissima pilsbryi from. the m-iddle part. of the Cxatiun in the western area is strongly inflated, lbut not as strongly inflated as the illustrated specimen from- Rio Banana, Costa Rica, which 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. nindiense has exceptionally thick and sharply limited calluLs. A moderately inflated specimen (height 20.3 mim, diameter 12 mi-) from- locality 1,57 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.
Prunuin cons for'ne (Sowerby) (Pilug, 1961, p. 55, pl. 15, figs. 12-6, 8, 9: Miocene, Dominican Republic, Jamaica) is larger, less inflated, less heavily callused, and has a lower spire. P. pr ecqtrsor (Pall) (1890-1903, p. 47, pl. 5, fig. 4, 18.90; Pliocene, Florida) also has a lowver spire. P. latisimum (Pall) (in Guppy and IDall, 1896, p). 08S, pl. 29, fig. 11; Miocene, Dominican Republic; P~liocene, Costa Rica) is smaller, more inflated, and its aperture is dilated near the base of the outer lip. P. storeiriqn (Couthiouy) (1837, p. 440, pl. 9, figs. 1, 2), now living along the Caribbean coast of Pailnm, is less inflated and its outer lip is smooth. Coan and Roth's (1966, p. 280, pl. 48, figs. 7, 8) illustrations of an unnamed shell dredged off Costa Rica indicate that a species similar to P. qatunense is living in eastern Pacific waters. As they noted, the inner edge of the outer lip of Brown and Pilsbry's Marginella coniformnis (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, 138(1. 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 Rica. Middle Miocene, Chiriqui area, Paiiam~i (USGS 7955)).

Genus rersicula Schumacher
Schumiacher, Eissal d'un nouveau syst~ne des habitations des
vers testaces, p. 235, 1817.
Typ1e (monotype andl tautotype) : Persicula variabilis. Schumacher (= Voluta persicula Linn6), living, west Africa.


332






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE33


Subgeniis Gibberula Swainson
SNwainson, A treatise onl malacology, p. 323, 1840. Typ~e (moniotype) :Gibbcrila zoniata Swainson (= TVolvaria oryza Lamarck) living, west Africa.
Gibbe)uda is low-spired, whereas Persicula s.s. has no spire; in fact, the, type has at 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 eolge of outer lip finely denticulate. Siphional canal notched. Columiellar lip bearing five or six folds.
Height 6_.5 mmn, dia meter 4 mim (figured inflated sipecimeiii). Restored height about 6.7i mmi, diameter 3.6 inn moderatelyy slender specimen).
Two silicified small miarginei lids from the Gatuncillo formation are similar to Peisicula semen, a middle and late Eocene species of southeastern United States (Palmer, 1937, p. 4122, pl. 67, figs. 13), 14, 16-19). They hiave a slightly higher spire and narrower aperture than P. semeui. Onie of thle fossils from the Canal Zone is inflated andl the other is fairly slender. The preservation of bo0th is poor, as the granular silica, is too coarse for satisfactory 1)reserN7ationi 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 (monlotype) :Pcrsicala intcrrupta (Lamarck) (Marginella
in trrupta Lamarck=TVolitta intcrruptoiincata Megerle),
living, west Africa and Caribbean Sea.
The heavy, more or less bifid second fromn anterior end) fold on the colinmiellar lip, callus spread over the entire lparietal wall, anid absence of an apical depression distinguish flabiea from Peirswula 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 l ip. Aperture very narrow. Outer lip wide, weakly bordered by low ledge. Inner edge of outer lip finely clenticulate. Siphional. canal notched. Five folds visible on columieilar 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 b)ands that have lost their color, or the spaces between color bands.
Height 7.6 mm,11 diameter 4.9 mmii (type).


Type: USNM 645860.
Trype locality: 116a (USGS 20956, east bank of Panama Canal at Canal station 1870 near Paraiso, Canal Zone; 700 feet (213 me-iters) southeast of locality 116), La B~oca formation.
This mnarginellid 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 observ-ed on shells of the Eocene species P. semen (Lea) (Palmner, 1937, p. 4-23) and on shells of P. 'reflez?(laria qvezuelana (F. Hodson, in Hodson, Hodsoil, and H-arris, 1927, P. 77).
P. vegeelafa amydra agrees with P. venezuelana VeneZUelana (F. Hodson) (in Hodson, H-odson, and Harr-is, 1927, p. 77, pl. 40, figs. 13, 14) in essential features, but is less inflated and a. little smaller. The noimite sulbspecies occurs in strata of early Miocene age in Venezuela. A largTer middle and late Miocene subspecies in Venezuela has been named P. venezuelava lacelana (F. H-odson) (in H-odson, H-odson, and Harris, 1927, p. 78, pl. 40, figs 3, 10, 11-, Rutsch, 1934, p. 91, p1. 6, figs. 9-11, 12).
P. vevezitelana is more inflated and has a less distinctly bordered outer lip than P. interruptolineata (Megerle) (Kiener, 1834-41, p. 25, pl. 5, fig. 21, 1834; as Martyiela interrupta Lamarck), the type of Rlabicea. Thug those two species are similar, P. cercadeinsis (Mlanrv) (1917, p. 73, pl. 11, fig. 7), a small, slender middle Miocene species from the Dominican Rlepub~lic, perhaps more closely resembles the type species. Despite its small size (height 7 mm), P. cercadei'sis has a wide, sharply bordered outer lip, like the type species. P. hnterrauptolineata is living in both west African and Caribbean waters. A comparable, if not, identical, species, P. inbricata (H-inds) (Coan and Roth, 1966, p. 283, pl. 48, fig 18, pl. 49, figs. 19-33), is found in the Panamnic, region.
Occurrence: Culebra formation (early Miocene), locality 106. La, Boca formation (early Miocene), localities 114, 1 15b, 116a.

Persicula (Rabicea) couviana stenygra Woodring, n. subsp.
Plate 51, figure 7
Of miediumii size, strongly inflated, apex about at level of posterior~ end of outer 1l1). Aperture moderately narrow. Outer lip) narrow, weakly bordered by a low ledge. Denticles on inner edge of outer lip fairly strong. Siphonal canal notched. Columiellar lip bearing seven


333






334 GEOLOGY AND PALEONTOLOGY OF 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, Panamd; 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.
Persioula couviana stenygra is less inflated and a little smaller than P. couviana couviana (Maury) (1925, p. 202, pl. 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. pro peobesa (Mansfield) (1925, p. 41, pl. 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 f auna 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, pl. 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 WXilson (1948). His upper Miocene collections from peninsular Florida include a loosely coiled Trigonostoina allied to T. mullei-, recently described as T. lwrrlei (Olsson, 1967, p. 24, pl. 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 p)resen~t Caribbean region, but survive in the Panamic region. Unidentified cancellarids, or doubtful cancellanids, are represented by poorly preserved material in the marine member of the Bohio (?) formation, the upper part of the IBohio formation of Barro Colorado Island, the Caimito formation of the Gatun Lake area, and the La Boca formation.

Gen-as Cancellaria Lamarck
Lamnarck, Soc. Histoire Nat. Paris Mrm., p. 71, 1799. Type (monotype): Voluta reticulata Linn&, living, Cape Hatteras to Caribbean Sea.

Subgenus Cancellaria s. s.
Cancellaria (Cancellaria) anomola Woodring, n. sp.
Plate 52, figures 1, 2
Small, moderately inflated, spire distinctly turreted. Protoconch relatively large, naticoid, 31/4-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 iiarrowv 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 lies 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 CancellaTia aiuoinoia was found in the lower part of the Gatun formation at locality 138c. rThe same locality yielded a minute shell, consisting of the protoconch and 21/4 sculptured whorls, and three imperfect shells, one of which is a little larger than the


334






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTIIOGLYPTIDAE35


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. rowel/i 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, pl. 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 Xanslleld Plate 52, figures 3, 4
Small, moderately inflated, spire distinctly turreted. Protoconch worn, naticoid, 2l/2-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 Cancel/aria 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. maeneili (Mansfield, 1937a, p. 609, pl. 85, figs. 1, 4; middle Miocene, Florida), if not that species. The early sculpture of C. Icevescens Guppy (Woodring, 1928, p. 220, pl. 12, figs. 7, 8; middle Miocene, Jamaica), is reticulate and stronger than that of C. macneili, whereas C. rowe//i Dall (in Guppy and Dall, 1896, p. 307, pl. 29, fig. 1; middle Miocene, Dominican Republic) 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 mmn (type).
Type: USNM 645714.
Type locality: 182a (USGS 8488, Caribbean coast east of San Miguel [Rio Miguel], station 25 plus 4:00 feet (120 meters), Panama.), upper part of Gatun formation.
Cancellaria tapeina 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. anominoa, C. aff C. mac'neili and C. tapeina are more similar to C. reticul at a 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) epistoniifera dariena Toula
Plate 52, figures 11-18
Gancellaria dariena Toula, K. k. Geol. Reiclisanstalt Jahrb., v.
58, p. 703, pl. 28, fig. 2, 1909 (Miocene, Canal Zone).
Brown and Pilsbry, Acad. Nat. Sci. Philadelphia Proc., v. 63, p. 345, pl. 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, p1. 6, fig. 8, 1922 (Miocene, Canal Zone, PanamA, Costa Rica). Weisbord, Idem, v. 14, no. 54, p' 50, p1. 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, p1. 75, fig. 4, 1965 (Miocene, Canal
Zone).


335







336 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


CaneceIlaria dariena n. sp., var., Toula, K. k. Geol. Reichsanstalt
Jahrb., v. 58, p. 704, p1. 28, fig. 1, 1909 (Miocene, Canal
Zone).
Canceilaria dariena track yostraca Brown and Pilsbry, Acad.
Nat. Sd. Philadelphia Proc., v. 63, p. 345, pl. 24, figs. 1, 2,
1911 (Miocene, Canal Zone).
Cancellaria daricncnsis Toula, Cossmann, Jour. Conchyliologie,
v. 61, p. 51, pl. 4, figs. 9, 10, 1913 (Miocene, Canal Zone). Canccilaria epistorniftra Guppy, Cossmann, Idem, p. 53, pl. 4,
figs. .5, 6, 1913 (-Miocene, Canal Zone).
Uxia rniocanica Cossrnann, Idem, 1p. 54, p1. 4, figs. 11, 12, 1913
(Miocene, Canal Zone).
Cancellaria (Can cellaria) daricna Toula, Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst., p). 118, 1964 (Miocene, Ecuador).
?Canccllaria (Canccllaria) daricna Toula, Oinomikado, Geol.
Soc. Japan Jour., v. 46, p. 623, p1. 29, fig. 19, 1939 (Miocene, Colombia).
Not Canceclaria? cf. C. dariena n. sp., Toula, K. k. Geol. Reichsanstalt Jahrb., v. 58, pl. 25, fig. 13, 1909 (Miocene, Canal Zone) ;=Antillophos candci gatienensis (Toula).
Of medium size, inflated to moderately slender, spire slightly to strongly turreted. Protoconch naticoid, 3whorled, 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 flaringr and bearing a, shallow notch. Siphioial f asciole strongly inflated. Parietal callus moderately thick.
Height 34 mm, diameter includingg strong term-inal varix) 21.5 mm (figured inflated specim-eni). H-eight 31 mim, diameter (including. weak term-inal varix) 16.5 mim (figured slender specimen).
rType material :Lectotype (herewith dlesiginated), rrechi I-ochischule, Vienna.
Type locality: Presumably Gatun Locks site, C!anal Zone, middle part of Gatun formation.
Cancellairia epinstoiif era dariena is the most widely distribultedl 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 iii the upper part of the Gatun in the western area, two incomplete worn shells, listed as C. aff. C. epistIomifera darieiia. were found in the Chagres sandstone.
A p)ronounced range of variation, in outline and sculp~ture, is shown by the fossils referred to C. epistomifera da?-iena. Trhe greatest range in outline is afforded by the samples from the lower part of the Gatun. An inflated specimen (p1. 52, figs. 17, 18), a slender specimen (p]. 52, figs. 15, 16), and a distinctly tulrretedl specimen (p]. 52, figs. 13, 14), were selected from the large collection just mentioned. That collection includes 13 distinctly turreted shells. A strongly turreted specimen (p1. 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. epistoinifera darieva 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 (p]. 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 fiat siphonal fasciole 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 rnge is shown on plate 52 for specimens from the lower part of the Gatun. rrol'ls and Brown and Pilslbry's illustrations serve the sam-e purpose for the mid(Ie part, and Cossmnann's for thme upper part in the eastern area. Fiv-e of somie 250 shells in the collections at hiand lack secondary spiral threads. At the other extremle, one (locality 138c) has exceptionally strong secondary threads.
C. epis tornif era epistfomifera Guppy occurs in the (1ercado and Gurabo formations of the Dominican Repulic. A lectotype, presumably from the Gurabo forilation, was selected andl ilhmstra-tecl by Pflug (1961, p. 52, pl. 14, figs. 1, 2, 9). Its lprotoconcll is not preServeN(l. MlaUry (1917, p. 63) pointed out that two types of 1)rotoconecl and first p)ost- protocoinch whorl are represented in her collections: (1) protoconch small, axial ribs appearing at beginning of first post-protocleiwhorl- (2) protoconch large, axial rb per


336






GASTROPODS: EUIJMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAB 3


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. C. 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, pl. T5, figs. 3, 4, respectively).
The chief difference between C. epistomif era dariena and the nominate subspecies lies in the greater range of variation, in outline and sculpture, of the Canal Zone subspecies.
The subgenus Ca'neellaria s.s. is an appropriate assignment for C. epistoinif era, although that species formerly (Woodring, 1928, p. 221) was referred to the subgenus later named JBivetiella by Wenz (1943, p. 1356; type (orthotype: Cancellaris simd;is 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 161,161la, 161c, 170 (identification doubtful). Upper part, eastern area, localities 172, 175, 176, 176a, 177 (identification doubtful), 177b, 178. Middle MioCenle, Costa Rica, Colombia, Darliin area (USGS 8477), Chiriqui area (USGS 7955). Lim16n formation (Late
Mi'ocene), Bocas del Toro area, Panaimt. Miocene, Ecuador. Chagres sandstone (late Miocene), locality 208 (C. aff. C. epistom-ifera dariena).
Cancellaria (Cancellaria) epistomifera lipara Woodring, n. subsp.
Plate 52, figures 7, 8
Smaller than Canela)ia 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 mim (largest specimen).
Type: USNM 645719.
Type locality: 183 (USGS 8487, Caribbean coast east of Rio San Miguiel [Rio Miguel], station 4 plus 40 feet (12 meters), Panamd), upper part of Gatun formation.


Can cellaria epistornifera 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. Ti marked contrast to the diversity shown by C. epistoinifera 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 C. epistomifera darieiia and C. epistomiifera epi stoinif era. Despite their small size, the largest specimens of C. epistomifera lipa ra. like the small form of C. epistomifera epistomifeira in the Gurabo formation of the Dominican iRepublic, have a swollen siphonal fasciole and heavy columellar folds, indicating that they are mature.
The Lim6n formation, of late Miocene age, on Ca-yo de Toro, in the lBocas del Toro area of northwestern Panamd. (USGS 8326), yielded one specimen of C. epistoimifera lipara (height 31 mm), associated with four of C. epistomifera dariena.
The axial ribs of an immature shell (height 16.3 mim) in the large collection from the lower part of the Gatun formation at locality 183c are even more closely spaced than those of C. epistornifera lipara, and so are the spiral cords. This fossil is doubtfully identified as ain aberrant form of C. epistoinif era dariena.
Occurrence: Upper part of Gatun formation, western areat (middle Miocene), localities 182a, 183. Lim6n formation (late Miocene), lBocas 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 p~reserved sculpture consisting of axial ribs and eight low, flat spiral cords. Axial ribs strong and widely spaced onl 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 mim, diameter 13 mmin (type).
Type: USNM 645720.
Type locality: 182at (USGS 8488, Caribbean coast east of San Miguel [Rio Miguel] station 25 plus 400* feet (120 meters), Panamai, upper part of Gatun formation.
Trlis 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.


337







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 Cancella'wia 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.
Subgenus Pyruclia Olsson
Olsson, Bull. Am. Paleontology, v. 19, no. 68, p. 160, 1932. Type (orthotype): Canceclaria solida Sowerby, living, Gulf of
California to Perul.
The earliest species of Pyruolia 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 Panami, Colombia, Venezuela, and Trinidad-and also in the iDarie'n area, Ecuador and Perid. The latest species in the present Caribbean region occur in deposits of late Miocene age.

Cancellaria (Pyruclia) cibarcola cibaxcola Anderson
Plate 52, figures 9, 10; plate 53, figures 8, 10-12
Gancellaria cibarcota Anderson, Calif. Acad. Sdi. Proc., 4th ser.,
v. 18, no. 4, p. 116, p1. 14, figs. 1-3, 1929 (Miocene, Colombia). Barrios, Colombia Servico Geol. Nac. Bol. Geol., v. 6, nos 1-3 (Informe 1082), p. 290, p1. 11, fig. 7, 1960
(Miocene, Colombia).
Not Cancellaria eibarcola Anderson, Rutsch, Naturforsch. Gesell. Basel Verh., v. 54, p. 163, p1. 9, figs. 6ah, 6b, 1942
(Miocene, Trinidad) ;-C. an riculaperta Yokes.
Of medium size to large, slightly to moderately shouldered. Spire of varying height, slightly to distinctly turreted, strongly inflated. Protoconch naticoid, 2 /-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 At1l.ntico, Colombia, Tubard formation.
CancellaTia 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 (pl. 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 fasciole is a little wider than on Gatun specimens. These Colombian fossils have also a slightly more abruptly inflated siphonal f asciole and a slightly channeled suture. Specimens from the Darien area closely resemble those from the Gatun formation.
C. pyeta Olsson (1964, p. 122, pl. 21, figs. 3, 3a) is considered to be a middle Miocene Ecuadoran subspecies of C. cibareola, distinguished by the persistence of sculpture to a slightly later stage.
A late Miocene species from Trinidad, C. auriculapert a Yokes (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. iRutsch, however, thought that C. auriculaperta is C. cibarcola. In general features C. cibarcola is similar to C. 8olida (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, Dari6n area (USGS 8433, 8476, 8493), Paniamk.

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-


338







GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE33


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 [IRio Miguel], station 25 plus 600 feet 150 meters, Panamd), 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. 8oheibei Anderson (1929, p. 115, pl. 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 f rom the upper part of the Gatun formation in the western area (C. tapeina) C. diadela is based on only one specimen. A fragmentary shell f rom 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 pal6oconchologie compar&e, pt. 3, p. 10, 1899): Gancellaria cassidliformis Sowerby, living, Gulf of California to Perd.
EFuclia, 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 ais that of Pyritclia: Panam Colombia, Venezuela, Trinidad, Dari~n area, Ecuador, and Peril. 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
Gancellaria codazzii Anderson, Calif. Acad. Sci. Proc., 4th ser.,
v. 18, no. 4, p. 116, pl. 14, figs. 4-7, March 29, 1929 (Miocene, Colombia). Barrios, Colombia Serviclo Geol. Nac., Bol. Geol., v. 6, nos. 1-3 (Informe 1082),y p. 291, pl. 11,
fig. 5, 1960 (Miocene, Colombia).
Cancellaria icarsteni Anderson, Calif. Acad. Sci. Proc., 4th ser.,
-v. 18, no. 4, p. 114, p1. 10, figs. 7-9, 1929 (Miocene,
Colombia).
Cancellaria hcttneri Anderson, Idem, p. 114, p1. 10, figs. 5, 6,
1929 (Miocene, Colombia).


Cancellaria (Euecia) maldonadoi Olsson, Neogene mollusks
from northwestern Ecuador, Paleontological Research Inst., p. 1.22, p1. 21, figs. 5, 5a, 1964 (Miocene, Ecuador).Of medium size, moderately slender to moderately inflated. Body whorl angulated to subrounded at shoulder. Protoconch naticoid, 2 /-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; 7 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 fasciole. 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 mmn, diameter 24 mm. (figured moderately angulated, moderately spinose specimen).
Type: Calif. Acad. Sci. 4645.
Type locality: Between Chorrera and Cibarco, Dept. of Atlkntico, Colombia, Tubara' formation.
This variable species is widespread and locally abundant in the lower part of the Gatun f ormation 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 (pl. 54, figs. 3, 4, 7, 8) are most abundant; those that are sharply angulated. and strongly spinose (pl. 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


339







340 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


occur together (Can cellai a karsteni and C. heitneri). C. codazzii is given preference, as the other two names were used for extreme forms. In the Dari~n area the range of variation is not as great as in Colombia and in the Gatun formation.
C. mnaldonadoi represents the subrounded, nonspinose form of C. codazzii. The spire whorls of C. 'werenf c/si .Jung (1965, p. 552, pl. 75, figs. 9-11; middle Miocene, Venezuela) are more distinctly turreted.
A middle Miocene Venezuelan species, C. veneouelana H. K. H-odson (Hodson and Hodson, 1931, p. 45, pl. 23, figs. 1, 4), is more inflated than C. codazzii and has coarser spiral sculpture. C. rnontserratensis Maury (Rutsch, 1942, p. 163, pl. 9, figs. 7a, 7b), a late Miocene Trriinidad species, also has coarser spiral sculpture and includes distinctly tabulated shells. In general features C. codazzii is similar to the type of Fuel ja, 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 Panam~i Bay: C. ba/bow. Pulsbry (1931, p. 439, pl. 41, figs. 7, 8). The type of C. ba/bowe is lost or misplaced and the paratype, is immature. Twenty-two specimens were collected recently from dredge dumpings at Fort Amador, Canal Zone, by IR. H-. Stewart, Joanne L. Allen, and Douglas Allen. According to this sample, C. ba/bowe is similar to the slightly or moderately angulated form of C. codaozii, but has fewer axial ribs on spire whorls. The type of C. ba/bowe (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, 139c, 139f, 155. Middle Miocene, Colombia, TDari~n area (UTSGS 8429, 8430, 8453, 8476, 8477), Panaimi. Angostura formation (middle Miocene), Ecuador.

Cancellaria (Euclia) dinata Woodring, n. sp.
Plate 54, figures 1, 2; plate 56, figures 5, 6
Of medium size, moderately inflated. Body whorl rounded at shoulder. Protoconch naticoid, 212-whorled. End of protoconch marked by appearance of sculpture. First sculptured whorl or two angulated or subanguilated 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) oin 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~ 0o penult; those on main part of body whorl almost as strong as pi-inaries. One or two (generally two) widened ribs on lhody whorl and preceding two whorls.
H-eight (practically complete) 29.9 mm, diameter 18 mmi (type). H-eight 34.9 mim, diameter 21 mmn (exceptionally large figured specimen).
Type: USNM 645728.
Typ~1e locality: 138c, (IJSGS 21956, about 100 meters north of Transisthinian H-ighway and about 75 meters west of road to refinery site on Payardi Island, Panamd; immediately east of Cativa and 100 meters north of locality 138), lower p~art of Gatun formation.
In marked contrast to the diversity shown by Canellaria codazzii. C. dinota is fairly uniform. The spacing of axial ribs on tile last third of the body whorl, however, is not uniform. Also on a few specimens the spiral thread ait the shoulder and those between the shoulder and tile 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, pl. 3, figs. 1, 2), a Pliocene species from Ecuador. That species is larger (height 43 min), 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. Immnature shells may be distinguished from immature shells of C. codazz:ii by the finer sculpture of C. dinota.
Occurrence: Lower part of Gatun formation (middle Mioceine), localities 136, 136a, 137a, 138, 138a, 138c, 138d, 138f.
Subgenus?
The following species is distinguished by its small size, slightly inflated spire whorls, narrowly reticulate sculpture, strong coluinellar folds, widely swinging si plional fasciole and acconi)anying relatively wide umbilicus. The general facies of form- and sculpture and the strong, columellar folds suggest Cancel/aria s.s., whereas the widely swinging siphional fasciole suggests Bi'vetopsia Jomisseaume (1887, P. 193; type (logotype, Cossinalnn, 1899, P). 9) : Cancellaia chrysostorna Sowerlby, living, Panannt to Peril) and Rii'etiella Wenz (1943, p. 1,356; type (orthiotype) : Cancel/aria similis Sowerby, living, west Africa). Contrary to the view


340






GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO IHELMINTHOGLYPTJDAE34


expressed in 1928 (Woodring, 1928, p. 221), when B'ivetopseis was inadvertently used for Bivetopia, Bivetopsia and Bivetiella 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 2 /-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, liram on interior of outer Ip, 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 meterss, long 790 49' 11., Panan-ii; same as ITSGS 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 aire 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 anid the lip of theo other specimens is dainaged.
The axial ribs of Cancellaria laqua Mansfield (1935, p. 26, pl. 2, fig. 5; middle Miocene, Florida), C. lavelava H. K. Hodson (in Hodson and Hodson, 1931, p. 44, ph. 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.

Subgenus?
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 pen-Llt, 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 fromn base) folds present or absent. Interior of outer lip bearing moderately strong lirx. Siphional 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 mmi, diameter 5 mm (largest specimen).
Type: USNM1 645731.
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.
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 Caveellaia vavcellar-ia. 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 f airly 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. qiancellaria is a minute species. The columiellar folds, the short siphional canal, and the position of the siphonal fasciole suggest Canoe lliia s.s.,


341






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., ann~e 9,
p. 222, 1887): Oanccllaria clavatula Sowerby, living,
Mexico to Perd.
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 bulibrooki Mansfield (1925, p. 31, pl. 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: Cancellariqa atraktoide8 Gardner (1926-47, p. 376, pl. 45, figs. 13, 14, 1938; middle Miocene), C. blountiana Mansfield (1935, p. 27, pl. 2, fig. 4; middle Miocene), C. agria uaquala Mansfield (1935, p. 27, pl. 2, figs. 7, 8; middle Miocene) C. ag7ia agria Mansfield (1930, p. 48, pl. 3, fig. 1; late Miocene), and C. coensis Mansfield (1930, p. 49, pl. 3, figs. 3, 4, late Miocene).

Cancellaria (Narona) barystoina Woodring, n. sp.
Plate 53, figures 5, 6
Small, slender, distinctly turreted. First two sculptured whorls weakly angnlated at shoulder, later whorls rounded. Protoconch small, naticoid, 2lA4whorled, 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. Lira,- 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 fasciole 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, Panamd; immediately east of Cativa and 100 meters north of locality 138), lower part of Gatun formation.
In essential features Cancella'ria 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 liroe 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 C. barystoma is represented by 13 specimens (12 from the lower part of the Gatun formation and one f rom a horizon near the base of the middle part), the type is the only mature shell. The next ]argest has a height of 10 mm. As is to be expected, its apertural folds and lira-, 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.

Canceliaria (Narona) decaptyx Brown and Pilsbry
Ciancllaria dzecaptyx Brown and Pilsbry, Acad. Nat. Sci. Phila.
Proc., v. 63, p. 346, pl. 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 C. barystovna 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 C. 75arystoma (11 on penult). The columellar folds and lira-, on the interior of the outer lip are of about the same strength as those on a specimen of C. barystorna 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.


342






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE34


Subgenus Charcolleria Olsson
Gisson, Bull. Am. Paleontology, v. 27, no. 106, p. 61, 1942. Type (orthotype): Cancellaria (Uharcoileria) Verdiciana Gisson,
Miocene, Colombia.
The large, fusiform, reticulately sculptured shells of Charcoller-ia 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 111assyla (H. and A. Adams, 1854, p. 278; type (monotype): Cancellaria corrugata Hinds, living, Ecuador to Perd) and Mass yla is generally treated as a subgenus of Cancellaria. The type of Mass yla is a rare species and no specimens are available. According to Hinds' (1844-45, p. 42, pl. 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 Mass yla for European Miocene species (Sacco, 1894, p. 67, pl. 3, figs. 80a, 80b; Cossmann, 1899, p. 39-41, pl. 2, figs. 13, 14) is unjustified.
Charcolleria has been found so far in the lower Miocene of Colombia, the middle Miocene of Colombia and the Caribbean part of Panamd, 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 Panamd. 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, pl. 28, figs. 8, Sa, 8b), which was assigned to Mass yla by Olsson and Harbison. Cancellaria pro pevenusta Mansfield (1930, p. 47, pl. 17, fig. 2; upper Miocene, Florida) may be a predecessor of Cancellaria venusta and perhaps Cancellaria pro pevenusta arose from Cancellaria run chcena Gardner (1926-47, p. 375, pl. 45, figs. 8, 9, 1938; middle Miocene, Florida). The resemblance of Cliarcolleria to Cancellaria venusta is attributed to convergence.

Cancellaria (Charcoileria) terryi Olsson
Plate 54, figures 5, 6, 9, 10'
Canoe ilaria (Charcolleria) terryi Gisson, Bull. Am. Paleontology, v. 27, no. 106, p. 62, p1. 8, fig. 1, 1942 (Pliocene, Costa Rica). Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst., p. 124, p1.
22, fig. 2, 1964 (Miocene, Ecuador). Jung, Bull. Am.
Paleontology, v. 49, no. 223, p. 556, p1. 75, figs. 17-19,
1965 (Miocene, Venezuela).


Cancellaria (Charcolleria) sp., Olsson, Neogene mollusks from
northwestern Ecuador, Paleontological Research Inst.,
p. 124, pl. 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 fasciole slightly inflated, swinging some distance from edge of columellar lip.
Height (incomplete) 53.5 mm, diameter 24 mmn (larger figured specimen). Height 42.3 mm, diameter 19 mm (smaller figured specimen).
Type: Paleontological Research Inst. 4045.
Type locality: Quebrada Pefiites, Puntareiias 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: Cane ilar-ia perdiciana (Olsson, 1942, p. 61, pl. 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 Cancelaria venusta (height 22 to 26 mm)-the Pliocene species already mentioned-and has more inflated spire whorls and stronger axial sculpture. Canelaria pro pevenusta
-also already mentioned-is low spired and inflated in outline; otherwise it is similar to Cancelaria venusta, aside from the difference in size. The type of Cancellaria pro pevenusta 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. Ciantaure formation (middle Miocene), Venezuela. Punta Gavi1kin formation (late Miocene), Venezuela. Esmeraldas formation (late Mio-


368-278 0 70 4


343






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 Recent Mollusca, v. 1, p. 277,
1854.
Type (monotype): Cancecliaria tessellata Sowerby, living, Gulf
of California to Per~l.
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 Paiiam6, southeastern Costa Rica, M6xico, Florida, and Perd; and tho upper Miocene of Per6. In other Nvords, Aphera (as well as Pyruclia, L'uclia, 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, pl. 3, figs. 76-79; Cossmann, 1899, p. 18, pl. 1, fig. 11) lack the distinctive apertural features of that genus.

Aphera islacolonis (Maury)
Plate 56, figures 1, 2
Cancellaria tcsscllata Sowerby, Gabb, Am. Philos. Soc. Trans.,
n. ser., v. 15, p. 236, 1873 (Miocene, Dominican Republic). Cancellaria (Aphcra) isiacolonis Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 65, pl. 10, figs. 12, 12a, 12b, 1917
Miocene, Dominican Republic).
Can cellaria islacolonis Maury, Olsson, Idem, v. 9, no. 39, p. 86,
pl. 6, fig. 12, 1922 (Miocene, Costa Rica) ;assigned to
subgenus Aphera.
Cance liaria ellipsis Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 73,
p. 333, pl. 22, figs. 8, 9, 1922 (Miocene, Dominican
Republic).
Small, elliptical, low-spired. Protoconch naticoid, 2 /-whorled, end marked by gradual appearance of sculpture. Sculpture reticulate. Spiral cords strongly overriding axial ribs, wider than the ribs on spire wh~orls, 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; lir.T on its interior strongly thickened just within aperture. Columella bearing two strong folds. Apertural shield thick; traces of underlying sculpture reflected on lparietal part; low, irregrular, elongate denticles on part opposite end of colume-


lar folds. Upper part of parietal wall bearing an elongrate denticle.
1-eigrht: 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 kmn above ford at Cercado de Mao), Dominican Republic, Cercado formation.
Apliera islaeolofli8 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 138,a) 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 lirTe on the interior of the outer lip are thicker just within the aperture. Perhaps these coarser features indicate that the Panamd fossils lived in shallower water than those in the Dominican Republic and they may eventually be given subspecitic rank. Olsson's figured specimen and the type of Pilsbry's Can cellaria ellipsis are not entirely mature. In M~xico (USGS 999.5, near Santa Rosa, Vera Cruz) A. islacolofli8 reaches a slightly larger size than elsewhere (height 18.5 mm).
A. islacolorvis is more similar to A. tessellata than to the other described species of Aph era. That living species is larger (height 26.5 mm), more slender, and has a higher spire. It is doubtful whether A. wig ginsi 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 foriation (middle Miocene). Lower part, localities 138a, 138c, 138f. Middle part, western area, locality 161c. Usciari shale (early Miocene part), Costa Rica. Middle Miocene (deposits, Costa Rica, MC16xico. Cercado formiation (middle Miocene), Dominican Republic.

Genus Trigonostoma Blainville
]Ilainville, Manuel (ie mnalacologie et de conchyliologie, p. 652,
1827.
Type (monotype an(l tautotype) :Dclphinula trigonostorna Lamarck (= T'rigona peilucida Perry), living, tropical western Pacific Ocean.


344






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTIIOGLYPTIDAE34


Petit (1967, p. 217) has pointed out that Lamarck's well known name for the type species is a junior synonym of Tr-igo'na pellucida.

Trigonostoma of. T. scalatella. (Guppy)
Small, slender, scalariform, sutural area sloping toward shoulder. Protoconch and spire whorls utp 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 suttural area. Near base a faint secondary thread appearing in two of spaces between spiral threads. Aperture and umbilicus narrow. Lirve 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 Trigonostorna 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, pl. 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. fu'niculatum (Hinds) (1844-45, p. 43, pl. 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)
Cancliaria (Trigonostoma) aff. Can cellaria Lullata Sowerby,
Toula, K. k. Geol. Reichisanstalt Jahrb, v. 61, p. 504,
p1. 30, fig. 10, 1911 (Mliocene, 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 Trigonzostonia i'nsuare (Pilsbry and Johnson, 1917, p). 163; Pilsbry, 1922, p. 334, 1)1. 22, fig. 11) and Olsson thought it probably is his T. toroense (Olsson, 1922, p. 84, pl. 6, fig. 4). The early finely reticulate sculpture of the Gatun species suggests that of 7'.


toroeflse, but the ribbing on the body whorl is more like that of T'. hsulare.
T. iiisulare is found in the Dominican Republic, presumably in the Cercado or Gurabo formation, and T'. toroe'nse in the late Miocene Limn6n formation of the iBocas del Toro area of northwestern Panamai.
Occurrence: Middle part of Gatun formation (middle Miocene), Gatun Locks excavation.

Trigonostonia 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.
H-eight (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 Trigonostorna woodr-ingi Jung (1965, p. 557, pl. 76, figs. 1, 2; middle Miocene, Venezuela) are wider.
Occurrence: Chagres sandstone (late Miocene), locality 208.
Family CONIDAE
Genus Conus Linn6
Linn6, Systema nature, 10th ed., p. 712, 1758. Type (logotype, Children, Quart. Jour. Sd. Lit. Art, v. 16, p.
69, 1823): Conus marmorens Linn6, living, western Pacific Ocean.
Though many attempts have been made to recognize subgenera, or genera, in the traditional genus 6'onus, 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 Chiagres 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. Thle exceptions are an


345







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

Closely allied or identical living species
Species in Gatun formation ____________Panamnic region Caribbean region

C. recognitus Guppy-- C. patricius Hinds --------C. spurius Gmelin ------------------------- C. spurius
Gmclin.
C. molis Brown and C. fergusoni ------Pilsbry. Sowerby.
C. consobrinus Sowerby- C. emersoni Hanna--------C. imitator Brown and C. arcuatus Brodcrip ------Pilsbry. and Sowerby.


Despite the large number of species in the Gatun formation, Conu8 pla'niliratus 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 ILim6n 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. 8timpofi is similar to C. planiliratu8 (Woodring, 1928, p. 211). The recently described C. austini (Rehider and Abbott, 1951, p. 22, fig. 7) is even more similar. In fact, it is doubtful whether C. awstini can be distinguished consistently from C. planiliurats by any feature other than the slightly larger size of the Recent form.
The status of C. tortuosopu'nctatus, 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, p1. 16, fig. 1, 1866 (Miocene, Jamaica). Not Conus solidus Gmelin, 1791, or Conus solidus Sowerby,
1834 (?).
Conus rccognitus 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, p1. 7, fig. 9, 1917 (Miocene, Dominican Republic). Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 73, p. 327, pl. 19, fig. 2, 1922 (Miocene, Dominican Republic). Olsson, Bull. Am.
Paleontology, v. 9, no. 39, p. 46, pl. 2, fig. 9, 1922 (Miocene, Costa Rica). Maury, Brazil Servi~o, Geol. y Mineral. Mon. 4, p1. 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) rccognitus Guppy, Pflug, Acta Humboldtiana, Geol. Pala~ontologica Ser., no. 1, p. 59, p1. 18, figs.
12-15, 1961 (Miocene, Dominican Republic).
Conus pyriformis Reeve, Gabb, Am. Philos. Soc. Trans., n. ser.,
v. 115, 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 f aint 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 Norte, 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 IL. 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


346






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE34


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. 8purius 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. recog'nitus; 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. 8olidus. 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. recog'nitus 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. williamgabbi (Maury, 1917, p. 36, pl. 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, pl. 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, pl. 8, pl. 10, figs. 4, 5), appropriately designated C. pyriformis by Reeve. Pilsbry pointed out that the tuberculate shoulder of C. recognitus disappears at an earlier stage than that of C. patriciws. 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, pl. 1, figs. 1, 2, 1844). The largest recorded C. recognitus is half as large as the exceptionally large C. patirwats recently illustrated by Hanna (1963, pl. 8; height 140 mm). C. patricius is the type of the recently proposed su-bgenus Pymiconus (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. Tubar~i formation (middle Miocene), Columbia.

Conus mnusaensis Olsson
Plate 57, figure 2
Gonus muswensis Olsson, Bull. Am. Paleontology, v. 9, no. 39,
p. 47, pl. 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, ll/2-whorled. Early postprotoconch whorls sharply carinate at shoulder. Carina gradually suppressed with further growth. Anal f asciole 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 mmn (figured specimen).
Type material: Lectotype (herewith designated), specimen represented by Olsson's fig. 24), Paleontological Research Inst. 20887.
Type locality: Rio Banana, Lim6n 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.
Tlhe small size and general appearance suggest C. jaspideus Gmelin, of the present Caribbean fauna, especdaily the round-shouldered form, for which Reeve's name C. pygmnwus is -used, either as a subspecies of C. Yasp? .deus (Clench, 1942, p. 12, pl. 7, figs. 1, 2), or as an infrasubspecific form of that species (Abbott, 1958, p. 89, 1)1. 3, fig. j). That the resemblance is superficial is indicated by the noncylindrical, more inflated protoconich and somewhat concave anal fasciole of C. jaspideus.


347






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 Rica.

Conus spurius Gmelin
Plate 55, figure 7
(Jonus spurius Gmelin, Systema naturm, 13th ed., p. 3396, 1791
(living, locality unknown). Clench, Johnsonia, v. 1, no. 6, p. 19, p1. 10, figs. 4, 5, 1942 (living, Bahamas, West Indies). (Jonits proteus Hwass, variety B, in Brugui~re, Encyclop~die
m6tlhodique, Histoire naturelle des vers, v. 1, p. 682, pl.
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, 1). 42, p1. 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, pl. 2, figs. 3, 4,
1922 (Miocene, Costa Rica, Panam6.).
Conus (Lithocorius) proteus Hwass, Woodring, Carnegie Inst.
Washington Pub. 385, p. 204, p1. 9, fig. 4, 1928 (Miocene,
Jamaica).
(Jonus iconinus 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 at 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 57)ur't8, formerly designated C. [pro teus, 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. rihe color pattern of the Gatun fossil is like that of C. spuriu.s atlaicius Clench (1942, 1) 20, pl. 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 ijrbbethat the color pattern can be used to distinguish geographic races in C. 8purius of Miocene age.


Occurrence: Middle part of Gatun formation, eastern area (middle M~iocene), locality 155. Middle Miocene deposits, southeastern Costa Rica, Colombia. Bowden formation (middle Miocene), Jamaica. Cercado and Gurabo form-ations (middle Miocene), Dominican Republic. Deposits of late Miocene and Pliocene age, Bocas del Toro area, Pa nami, Lim~n Peninsula, Costa Rica. Living, Florida, Bahiamas, and *West Indies.

Conus species
A small poorly preserved cone (restored height about 24 mmli, diameter 13.5 mm) from the Caimito formation of the Gatun Lake area has a moderately low, eel tapering spire and a wide shoulder. The shouldris abruptly roumndedl andl 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 Co'nus cf. C. marginatus Sowerby of H-ubbard (1920, p. 162, pl. 24, fig. 16), which occurs in limestone of early Miocene age in Puerto Rico (Aymam-in limnestone of present terminology). Conus marginatus Sowerby (1850, p. 44; not Conus mnarginatus J. de C. Sowerby, 1840), as identified by Guppy (1876, p. 528, pl. 29, fig. 5), has relatively strong spiral sculpture on the body whorl.
Occurrence: Caimito formation, Gatun Lake area (late Ol1igocene), locality 55b.

Conus bravoi Spieker
Plate 56, figures 10, 11
(Jonus molis var. bravo Spieker, Johns Hopkins Univ. Studies
in Geology, no. 3, p. 41, pl. 1, fig. 6, 1922 (Miocene, Peril). (Jonus (Dcndroconus) 1ravoi Spieker, Olsson, Bull. Am. Paleontology, v. 19, no. 68, 1p. 151, pl. 16, figs. 1, 3, 4, 1932 (Miocene, Perfu). Marks, Idem, v. 33, no. 139, p. 137,
1951 (M.Niocene, Ecuador).
Jons molis Brown and Pilsbry, Barrios, Colombia Servicio
Geol. Nac., Bol. Geol., v. 6, nos. 1-3, (Informe 1082), p.
294, 1)1. 12, fig. 8, 1960 (Miocene, Colombia).
(Jonas rios an tiagcws s01sson, Neogene mollusks from northwestern Ecuador, P~aleontological Research Inst., p. 85,
1)1. 13, figs. 1, la, 1b, 1964 (Miocene, Ecuador).
Moderately large, broad-shouldered. Edge of shouldeCr marked by angulation, generally pronounced, but of varying strength. Spire low, profile smoothly conical1, except part formed by strongly emerging early wiorl s. Protoconch mi ssing. Early post-protoconch wh-lorls worn, but showing suggestion of tuberculate si oulder. Succeeding whorls overlapping to edge of shoulder, or not quite to edge. Anal fasciole flat or somiewh-at 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.


348






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE34


Height (incomplete) 64 mm, diameter 44 mm (larger figured specimen). Height 59.4 min, diameter 36.6 mm (smaller figured specimen).
Type: Peabody Museum, Yale University.
Type locality: Zorritos district, Perii, Zorritos f ormation.
This Peruvian and Ecuadorean species is fairly abundant in the lo-wer 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 fasciole, which lacks spiral sculpture or is faintly to weakly sculptured. Three immature shells from locality 138c have a more distinctly concave anal fasciole 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 Tubard 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. bra coi. 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 Gatnn forination (middle Miocene). Lower part, localities 136, 136a, 137a, 138c, 138e, 138f. Middle part, eastern area, localities 155, 159d. Tubarat formation (middle Miocene), Colombia,. Progreso and Angostura, formations (middle Miocene), Ecuador. Zorritos, Cardalitos, and Montera, formations (middle Miocene), Perfi.

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 Ipost-protoconch whorls worn, shoulder angulation exposed on later whorls. Anal fasciole concave, marked only by growth lines. Lower part of body whorl sculptured with widely spaced spiral grooves.
Height 35 mmi, 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 790 49' W., Paniamk; same as USGS 16912), lower part of Gatun formation.
Conu8 acolu8 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. b'ravoi, with which it is associated, by its more evenly conical spire, more concave anal f asciole, and stronger spiral sculpture on the lower part of the body whorl.
C. cercadens8 Maury (1917, p. 43, pl. 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, althougrh in reduced form. In outline C. acolus suggests a small version of C. berinudenmis Clench (1942, p. 34, pl. 13, fig. 4), a living western Atlantic species from Bermuda and the east coast of Florida.
Occurrence: Lower part of Gatun formation (middle M~iocene), localities 136, 136a, 138c. Middle Miocene deposits, Chiriqui area, (USGS 7955, identification doubtful), Panamai.

Conus cf. C. sauridens Conrad
The silicified fossils of the Rio Casaya area include a small incom-plete, elongate, low-spired Conus that probably is allied to C. saul ide'n8, a species of southeastern United States ranging in age from middle Eocene, to early Oligocene (Palmer, 1937, p. 458, pl. 71). The shoulder is sharply carinate and the anal fasciole 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 fasciole, but the granular silica is too coarse to reproduce low threads like those of C. sauridens. The restored height is 30 mm,1 the diameter 15 mm.
A similar late Eocene and late Eocene or early Oligocene cone from Per~t has been inmed C. chirwnsis (Olsson, 1930, p. 39, pl. 5, figs. 1, 2, 4, 10). C. chircensis, treated as a subspecies of C. sam-iden8, has been identified in deposits of late lEocene age ini Columbia (Clark, in, Clark and Durham, 1946, p.* 46, pl. 24, figs. 11, 16, 18, 19).
Occurrence: Gatuncillo formation, Rio Casaya, area, (middle Eocene), locality 38.

Conus cf. C. suiculus Dail
Trhe upper part of the JBohio formation on Barro Colorado Island and the Caimito formation at a nearby locality yielded poorly preserved remains of a small


349






GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


species of Conus. 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 f asciole 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, pl. 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 jolaniceps Heilprin (1887, p. 110, pl. 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 fasciole 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. pla'nicep8 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, pl. 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. Dall's 1890 illustrations (Dall, 1890-1903, p. 25, pl. 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 M6xico, recently identified as C. plarneeps (Perrilliat Montoya, 1963, p. 27, pl. 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 101h, 115a, 115b, 116a.


Conus molis Brown and Pilsbry
Plate 55, figures 8-10
Conu flmrolis Brown and Pilsbry, Acad. Nat. Sci. Phila. Proc.,
v. 63, p. 343, pl. 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, pl. 2, figs. 1, 2(?), 1922 (Miocene, Costa Rica, Panami, Canal Zone). Weisbord, Idem, v. 14, no.
54, p. 56, p1. 6, fig. 1, 1929 (Miocene, Colombia). Anderson, Calif. Acad. Sd. Proc., 4th ser., v. 18, no. 4, p. 109,
1929 (Miocene, Colombia).
fWonus 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, p1. 23, figs. 5, 6, 1911 (Miocene, Canal Zone). Olsson, Bull. Am. Paleontology, v. 9, no. 39, p. 43, 1922. (Miocene, Canal Zone, Panamd). Anderson, Calif.
Acad. Sd. Proc., 4th ser., v. 18, no. 4, p. 110, 1929 (Miocene, Colombia).
Conus (Lithoconus) concavitecturn Brown and Pilsbry, Cossmann, Jour. Conchyliologie, v. 61, p. 43, pl. 4, figs. 3, 4,
1913 (Miocene, Canal Zone).
?Conus domintiensis Sowerby?, Brown and Pilsbry, Idem, p.
341, 1911 (Miocene, Canal Zone).
Not Gonus moli Brown and Pilsbry, Barrios, Colombia Servicio
Geol. Nac., Bol. Geol., v. 6, nos. 1-3 (Informe 1082), p. 294, p1. 12, fig. 8, 1960 (Miocene, Colombia) ;=C. 'bravoi
Spieker.
Exceptionally large, elongate, moderately wide at shoulder. Edge of shoulder generally abruptly angillated. Spire of moderate height, its profile concave. Protoconch consisting of 2 / 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 fasciole generally concave, bearing spiral threads. Growth lines of fasciole 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 Co'nu8 8purfi8s.
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.
conu8 MORiS, based on a mature shell, is given precedence over C. concavitec turn, the type of which is immature.
This large species-the largest American fossil species-is found throughout the Gatun formation. A total


350






GASTROPODS: ETJLIMIDAE, MARGINELLIDAE TO IHELMINTHOGLYPTIDAE35


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 tijan that of the others and their anal fasciole is flat or almost flat. Though the shoulder and fasciole suggest C. ammulator (the next species described), these two fossils are larger than that species and have the elongate outline of C. molis.
The distribution of C. m08i is clustered around the southeastern border of the Miocene Caribbean Sea: in Costa Rica, Panama., and Columbia. It has a close Miocene ally in Ecuador and Perfi: C. ca#cuminatus Spieker (1922, p. 40, pl. 1, fig. 5), which has more strongly tuberculate early whorls, a more deeply concave anal fasciole, and is not known to be of comparable size. C. haytensis Sowerby (Pflug, 1961, p. 60, pl. 16, figs. 1-5; Cercado and Gurabo, formations, Dominican Republic)
-the most similar fossil species in the West Indiesis wider at the shoulder, and has a less concave anal fasciole and more strongly tuberculate early whorls. It is likely that Maury's C. molis3 is an immature C. haytensi8. On the contrary, Brown and Pilsbry's C. haytensis and C. doiningensis?, both from the Gatun formation, are likely to be immature C. mo/is. The stubby outline and flat anal fasciole of the small middle Miocene Venezuelan cone identified as C. aff. C. mo/is (Jung, 1965, p. 575, pl. 78, figs. 1, 2) indicate that it is more similar to C. ammulator than to C. molis.
No similar species is living in the Caribbean Sea, but C. fergusoni Sowerby (Hanna, 1963, p. 42, pl. 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 fasciole of C. mo/is 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, 176a, 177, 177a, 177b, 177c; western area, locality 185. Middle Miocene, deposits, Costa Rica. Tubardi formation (middle Mio-


cene), Colombia.. Lim6n f ormation (late Miocene), Bocas del Toro area, Panamak.

Conus aemulator aemulator Brown and Pilsbry
Plate 55, figures 5, 6; plate 56, figures 4, 8
Conus wmulator Brown and Pilsbry, Acad. Nat. Sci. Phila.
Proc., v. 63, p. 342, p1. 23, fig. 9, 1911 (Miocene, Canal Zone). Pilsbry and Brown, Idem, v. 69, p. 32 (list), 1917
(Miocene, Colombia).
Not Conus proteus wvmulator Brown and Pilsbry, Rutsch,
Schweizer. Pala~ont. Gesell. Abh., v. 55, no. 1, p. 104, pl. 9, 'figs. 7-11, 1934 (Miocene, Venezuela) ;-C. avmulator
nwnzanillmnsis Mansfield.
Conus haytensis 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,
p1. 2, figs. 5, 8, 1922 (Miocene, Panamal, Canal Zone).
Woodring and Mansfield, in Geology of Republic of Haiti, p. 175, pl. 15, figs. 1, 2, 1924 (Miocene, Haiti).
Anderson, Calif. Acad. Sdi. Proc., 4th ser., v. 18, no. 4, p. 108, 1929 (Miocene, Colombia). Trechmann, Geol. Mag., v. 72, no. 858, p. 545, pl. 21, fig. 10, 1935 (Miocene, Carriacou). Gisson, Neogene mollusks of northwestern Ecuador, Paleontological Research Inst., p. 82, 1964 (Miocene,
Ecuador).
?Conus sp., Toula, K. k. Geol. Reiclisanstalt Jahrb., v. 58, p.
710, pl. 25, fig. 18, 1909 (Miocene, Canal Zone). Engerrand and Urbina, Soc. Geol. Mexicana Bol., v. 6, p1. 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 f asciole practically flat or slightly concave, bearing spiral threads. Lower part of body whorl bearing weak spiral sculpture; the threads f aintly pustulose on very young shells.
Height 45 mm, diameter 28 mmn (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 Co'nus emulator. The type (pl. 55, figs. 5, 6), photographed by A. A. Olsson, is immature (height 22.7 mm) and therefore does not show the fiat-topped late spire whorls. The largest specimen examined (height 59 mm) was found on Rio Chico, in Dari~in (USGS 8433). The less elongate outline, fliattopped late spire whorls, and flatter anal fasciole distinguish mature shells of C. wemulator from immature shells of C. mo/is. The two species are associated at many localities and in many features are similar.


351






GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


C. ceinulator is fairly widespread inl the Gatun formlation, 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 Ipart of the Gatun in the western area-the only s1)ecilens 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 Contis sp. is missing in Ilis collection. Ills illustration suggests anl immature C. cemulator.
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. manzanillmnsis Mansfield (1925, p. 12, p1. 2, figs. 5, 10; San Jos6 calcareous silt member of Manzanilla, formation, Trinidad), is assigned to subspecific rank under C. cemlator. It is smaller than the nominate subspecies and has faintly pustulose spiral sculpture onl the lower part of the body whorl, like immature specimens of the nominate subspecies. The anal fasciole of the type is very weakly sculptured, but the sculpture is stronger on other specimens. The largest in a collection of three (height 36 mlm) from the San Jos6 calcareous silt member on Guaracarita River (USGS 21098) is high-spired, like Rutsch's Venezuelan specimens.
Occurrence: Lower, middle, and upper parts of Gatun formation (middle Miocene). Lower part, localities 136a, 138, 138a, 138c, 138d, 138e) 138f. Middle part, eastern area, localities 139c, 139e, 139f, 14711, 155, 155a, 155b, 155c, 159, 159d, 160 (identification doubtful). Upper part, western area, locality 182. lBaitoa formation (early Miocene), Domlinican Republic. Thomonde formation (early Miocene), Haiti. Deposits of early Miocene age, Carriacou, Grenadine Islands. Deposits of early (?) Miocene age, 1]cuador. Coeado and Gurabo formations (middle Miocene), Domlinican Republic. Tubar. formation (middle Miocene), Colombia. Middle Miocene deposits, IDari~n. area (USGS 8429, 8433, 8477, 8479, 8492), Panamla. Liml6n formation (late Miocene) Blocas del Toro area, Panama.

Conus cf. C. catenatus Sowerby
Sinall, moderately slender, a ugul ated at shoulder. Spire high, its profile concave. Angfulated and tuberculate shoulder ap~pearingr onl first lpost-protoconch whorl. Trubercles gradually disappearing on third whorl. Last few whorls overlappin g almost to shoulder of preceding whorl. Anal fasciole fiat or slightly conicave, bearing weak spiral threads or none. Spiral scullpture of body whorl weak, faintly to distinctly pustulose or nonpustulose.
Height 25.7 mil, diameter 12 mml (largest specimen).


Four small cones in two collections from the Gatun Third Locks site are identified as Conu8 cf. C. catenat.s. Without much doubt they are immature. In some features they suggest C. catenatus (Pilsbry, 1922, p. 328, pl. 22, figs. 3, 4; Woodring, 1928, p. 213, pl. 11, figs. 4, 5)), a middle Miocene species from the Dominicall Republic and Jamlaica. At the growth stage represented by the last few whorls of the Gatun fossils the shoulder of C. catenates is not as strongly angrulated.
Occurrence: Middle part of Gatun formation, eastern area (middle MIiocene), localities 155, 155a.

Conus consobrinus consobrinuls Sowerby
Plate 56, figures 3, 7, 9
Conus3 consobrinus Sowverby, Geol. Soc. London Quart. Jour.,
v. 0, p. 45, 1850 (Miocene, Dominican Republic). Maury, Bull. Am. Paleontology, v. 5, no. 29), p). 39. 1p1. 6, figs. 5, 6, 1917 (Mliocene, Dominican Republic). Pilsbry, Acad. Nat.
Sdi. Phila. Proc., v. 73, p. 330, pl. 20, figs. 7, 7a, 7b, 1922
(Mliocene, Dominican Republic).
Con us (Lcp to con us) conso brin us Sowerby, Woodring, Carnegie
Inst. Washington Pub. 385, p. 214, pl. 11, figs. 6, 7, 1928 (M.Niocene, Jamaica). Perrilliat M.Nontoya, M6xico Univ.
Nac., Inst. (leologia Paleontologia Mexicana, no. 8, p. 26, 1)1. 4, figs. 3, 4, 1960 (Miocene, M4'xico). Pflug, Acta i-Inmboldtiana, Geol. Palmeontologica Ser., no. 1, p. 62,
1)1. 17, figs. 1-10, 1961 (MAiocene, Dominican Republic). ?Conuls consobrinus Sowerby?, Anderson, Calif. Acad. Sdi. Proc.,
4th ser., v. 18, no. 4, p. 111, 1929 (Miocene, Colombia). Not Conu.s con sobrinuts Sowerby, Gabb, Acad. Nat. Sci Phila.
Jour., 2d1 ser., v. 8, p. 359, 1881 (Pliocene, Costa Rica)
= C. (Oooobri ii. ultin 118 Pilsbry and Johnson.
Not Con111s consobrinus Sowerby, Brownm and Pilsbry, Acad. Nat.
Sdi. P~hila. Proc., v. 63, p. 341, 1911 (Miocene, Canal
Zone) ; =C. tortlloostriat u Ton.
Not Con us (Chl (qon us) con solrinus Sowerby, Cossmann. Jour.
(oncblYliologie, v. 61, p). 46, pl. 3, figs. 17, 18, 1913 (Miocene, -Martinique).
Conus (Conos pira ) lurilici Cossmann. Jour. Conchyliologie, v.
61, p). 38, 1)1. .4, figs,. 1, 2, 1913 ('Miocene, Canal Zone).
Con 15!K 1 (IZana U to i(', -Maury, B~ull. Am. Paleontology, v. 5,
no0. 29, p. 39, 1)1. 6, fig. 7, 1917 (Miocene, Dominican
Republic).
Of mediumii size, elongate, moderately wide at shouldecr. Shoulder sharply angrulated. Anal fasciole slightly sloping, p)rodlucing an almost square shoulder. Spire highl and its P~rofile almost smoothly conlical, or moderately high and its profile slightly conlcave. Proto(011(41 and~ earliest p)ost-protoconlch wvhorls missing. Shoulder of all except last, or last two, spire whorls strongly tuberculate ; tublercles gradually disappearing. Anal fasciole c oncave, bearing faint or distinct spiral threads. Lower part of body whvlorl. weakly sculptured; the threa"Is faintly lpustulose onl immature shells.
H-eight 61.7 mdiamleter 30.3 mil (larger figured specim-en).


352






GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO IHELMINTHOGLYPTJDAE35


Type material: Lectotype, British Museum (Natural History) G83962.
Type locality: Valley of Rio del Norte, Domiinican Republic, Miocene.
Conies consob2-inus eonsobrinus 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 Mindi, 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 (pl. 56, fig. 9) is the only one that is fully mature. Its spire is somewhat worn. Gatun specimens are alm-ost 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. scaliav (Bd6se, 1906, p. 51, pl. 5, figs. 41, 42) suggests an immature C. eansobrsnus. His type (height 15.2 mmi), however, lacks spiral sculpture on the anal fasciole. That C. consob', nus co'nsobrinus 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 U. S. National Mnseum (USGS 10346).
The high, or moderately high, strongly tuberculate spire is a diagnostic feature of C. consobr-inus. 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 specimeins that have a height of 50 mm. This formn is C. covsobriflus udtimus Pilsbry and Johnson (Pilsbry, 1922, p. 330, pl. 20, fig. 8). It first appears in the late Miocene TLim6n formation of southeastern Costa Rica and continues in the Pliocene Momn formation of that area. C. torornsis Glsson (1922, p. 48, pl. 2, fig. 7) may be treated as a weakly sculptured late Miocene subspecies. C. em ersofli Hanna (1963, p. 25, pl. 1, fig. 2), dredged off Cape Sami Lucas, Baja California, is probably a descendant of C. cons ob rin'us. It is roundshouldered and lacks tubercles on the shoulder of the body whorl, and therefore is similar to C. consobrimus cofsobriflus.
Occurrence: Lower, middle, and upper parts of Gatun formation (middle Miocene). Lower part, locality 138c. Middle part, eastern area, locality 139d; west-


ern area, locality 161b. Upper part, eastern area, Cossmann's record; western area,, locality 185. Middle Miocenie deposits, Dari~in area smalll race, USGS 8430, 8477), Panama. Agueguexquite formation (middle Miocene), Tfehulantepec area, M6xico. Bowden form-ation (middle Miocene) Jamaica. Gurabo formation middlee Miocene), Dominican Republic.

Conus symmetricus Sowerby
Plate 57, figures 13, 14
Con us symmetricus Sowerby, Geol. Soc. London Quart. Jour.,
v. 6, 1. 44, pl. 9, fig. 1, 1850 (Miocene, Dominican Republic). Maury, Bull. Amn. Paleontology, v. 5, no. 29, p. 36, p1. 7, figs. 7, 7a, 1917 (Miocene, Dominican Republic).
Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 73, p. 328, pl. 20, figs. 2, 2a, 2b, 1922 (Miocene, Dominican Republic). Not
Conies syrmmetricus Sandberger, 1862.
Conus (Lcptoconus) symenmtricus Sowerby, Pflug, Acta Humboldtiana, Geol. Paleontologica Ser., no. 1, p. 63, p1. 18,
figs. 1-11, 1961 (Miocene, Dominican Republic).
Conuts symmcntricus scmiobsolctus Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 37, p1. 7, fig. 8, 1917 (Miocene, Dominican Republic).
Con us synim ctricus doniingen sis Sowerby, Sowerby, Maury,
Idem, p. 37, p1. 4, fig. 10 (Miocene, Dominican Republic). 2 Conus (Litliocon us) 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 postlprotoconich 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, arrangyed in axial series, or not so arranged.
Height 20.4 mm, diameter 11.8 mm (figured specimnen).
Type material: Lectotype, British Museum (Natural History) G83696.
Type locality: Valley of Rio Yaque del Norte, Domninican Republic, Miocene.
Two specimens, both immature and both collected at the Gatun Third Locks site, are referred to this small, broaci-shoul dered, delicately sculptured, pustulose species. Trhe larger of the two is illustrated. On the early p~art 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, Jama1"ica, miay be a corroded Conus syinmetricus, this is the first unequivocal record for that species beyond the Dominican Republic, where it is abundant in the Gurabo formation. Sowerby's illustrations is unmistakable.


353






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. 8ymmetricus senciol38oletus. The spire is low and may be so low that the shell is almost flat-topped : Maury's C. sylnmetricus 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.

Conus 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 t~o edge of shoulder of preceding whorl, bearing a narrow channel adjoining suture. Anal fasciole 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 mmn (type).Type: IJSNM 645748.
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.
Conus aff. C. chipolanus Dali
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 B~oca formation. The Bobio 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 mmn and a diameter of 15.2 mm. Its shell material, like that of the others, is replaced by calcite.
Thug these fossils are unsatisfactory, they are allied to C. chtipolanus, which is discussed under the next heading, and show that in the Canal Zone the lineage of C. 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 C. aff. C. 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 101h, 115a, 116a.

Conus imitator imitator Brown and Pilsbry
Plate 55, figures 1, 2
(Jonus imitator Brown and Pilsbry, Acad. Nat. Sci. Phila. Proc.,
v. 63, p. 342, p1. 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, 1. 45, p1. 2, fig. 6, 1922 (Miocene, Canal Zone, Panami., Costa Rica). Pilsbry, Acad. Nat. Sci Phila. Proc., v. 73, 1). 327, 1922 (Miocene, Dominican Republic). Anderson, Calif. Acad. Sdi. Proc., 4th ser., v. 18, no. 4, p. 108, 1929 (Miocene, Colombia). Perrilliat Montoya, M6xico Univ.
Nac., Inst. Geologfa, Paleontologia 'Mexicana, no. 14, p.
27, p1. 6, figs. 6, 7, 1963 Miocene, Mkxico). Barrios, Colombia Serviclo Geol. Nac., Bol. Geol., v. 6, nos. 1-3 (Informe
1082), p. 295, pl. 12, fig. 6, 1900 (Miocene, Colombia).
Not Conus imitator Brown and Pilsbry, Li, Geol. Soc. China
Bull., v. 9, p. 275, p1. 8, figs. 72 (mislabelled 70), 72a, 1930 (Miocene, Panama Bay;- C. mahogani 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, p1. 10, figs. 5, 6, 1928 (Miocene, Jamaica).
?Conus cf. imitator lius Woodring, Olsson, Neogene mollusks
from northwestern E~cuador, Paleontological Research
Inist., p. 84, 1964 (Miocene or Pliocene, Ecuador).
(Jonus dalli Toula, K. k. Geol. Reichsanstalt Jahrb., v. 61, p.
509, p1. 31, fig. 23a-d, 1911 (Miocene, Canal Zone). Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 48, p1. 7, fig. 15, 1917 (Miocene, Dominican Republic). Alancister-Ibarra, Asoc. Mexicana Ge~logos Petroleros Bol., v. 2,1). 570, fig.
17, 1950 (Miocene, M~xico). Not Conu8 dalli Stearns,
1873.
Conus (Lithoconus) dalli Toula, Cossmann, Jour. Conchyliologie, v. 61, 1). 41, p1. 3, figs. 30, 31, p1. 4, figs. 7, 8, 1913
(Miocene, Canal Zone).
Of medium size, moderately wide at shoulder, shoulder strongly carinate. Spire high, its profile almost


354






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE35


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 caninate 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 fasciole 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., diameter 22 mm (larger figured specimen).
Type: Acad. Nat. Sci. Phila. 1688.
Type locality: Gatun Locks excavation, Canal Zone, middle part of Gatun formation.
C onu8 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 fasciole, 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. chipotanus Dall (Gardner, 1926-47, p. 360, pl. 43, fig. 6, 1938), or from the late Oligocene and early Miocene form identified as C. aff. C. chip olanus. 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 Dari6n area, Jamaica, and the Dominican Republic. C. sophus Olsson (1932, p. 154, pl. 16, figs. 6, 8, 9; middle Miocene, Peril 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, pl. 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 fasciole 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, pl. 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, pl. 3, figs. 4-6), now living at moderate depths (15 to 50 fathoms) in the eastern Pacific Ocean, from the Gulf of California to Panamd. 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.


355






356 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


C. imitator imitator is more similar to C. arcuatu8 than to the western Atlantic species C. floridanus Gabb (Clench, 1942, p. 27, pl. 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 fasciole of late whorls is flat or practically flat; the protoconch is blunt and 112-to 2-whorled. A small middle Miocene Costa Rican form has been named C. floridanus costaricernsis (Olsson, 1922, p. 45, pl. 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 chip olanus -imitator lineage adapted to shallower water than C. arcitatus.a few fathoms to 25 fathoms, exceptionally up to 40 fathoms. Since Miocene time C. floridarus, like Antilloplios 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; wvestern area, locality 161a. Upper part, eastern area, localities 171, 172, 173, 175, 177, 177al 177b, 177d, 178; western area, locality 182. Chagres sandstone (late Miocene), locality 208 (Conus imitator?). Middle Miocene deposits, southeastern Costa Rica, Tehuantepec area, Mexico, Colombia, Darien area (USGS 8477), Panam. B~owden formation (middle Miocene), Jamaica. Cercado and Gurabo formations (middle Miocene), Domninican Republic). Lin16i formation (late Miocene), Boca,s del Toro area, Panaimi, TLim6n Peninsula, Costa Rica.
Conus multiliratus multiliratus Bose Plate 57, figures 3, 4
onuss agas~sizi rnultiliratus lBise, Inst. Geol. ixico Col. 22,
p. 49, pl. 5, figs. 34-38 (mnultistriatus in explanation of
plate), .1906 (Miocene, Mexico).
('onus8 (Leptoconus8) muitiliratus Bise, Marks, Bull. Am. Paleontology, v. 33, no. 139, p. 139, 1951 (Miocene, Ecuador). (onus n tistriatits B~se, Olsson, Idem. v. 9, no. 39, p. 54, p1. 1,
figs. 21, 23,1922 (Mioceiie, Painmd).
Conus gaza Johnson and Pilsbry, in Brown and Pilsbry, Acad.
Nat. Sd. Phila. Proc., v. 03, p). 342, p1. 23, figs. 2, 3, 1911 (Mioeene, Dominican Republie, Canal Zone). Pilsbry and Brown, Idem, v. 09, p. 32 (list), 1917 (Miocene, Colombia) Maury, Bull. Am. Paleontology, v. 5, no. 29, p. 46,
pl. 7, fg. 12, 1917 (MNioeene, Dominican Republic).
Not Conits mulitiliratu8 gaza JIohnson and Pilsbry, Spieker,
Johns Hopkins lUivN., Studies in Geology, no. 3, p). 37,
1922 (AMioceiie, 1'erfi).
('onuts (Lcptoconits) inultiliratits gaza Johnson and Pilsbry,
Woodring, Carnegie Inst. Washington Pub. 385, p. 212, pl. 11, fig. 3, 1928 (Miocene, Jamaica) Olnomikado, Geol.
Soc. Japan Jour., v. 40, p. 620, 1)1. 29, fig. 30, 1939 (Miocene, Colombia).


('onus (Leptoconits) planiliratus Sowerby, Cossmann, Jour.
Conehyliologie, v. 01, p. 48, p1. 3, figs. 24-26 (not fig. 27; = C. plan iliratuts Sowerby), 1913 (Miocene, Canal Zone,
Jamaica).
('onus niargin atits Sowerby, Maury, Bull. Am. Paleontology, v.
5, no. 29, p. 40, p1. 7, fig. 11, 1917 (MINiocene, Dominican
Republic).
Small, biconica I, wide at shoulder, shoulder strongly carinate. Spire high, its profile alinost smoothly conical. Protoconch slender, cylindrical, 212- 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 wvhorl overlapping almost to carina. Anal fasciole slightly concave, bearing growth threads exaggerated at fairly regular intervals. Entire body whorl sculptured with strong-, narrow, stralplike 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 mmn, diameter 14.4 mmn (figured specimen).
Type: Apparently lost, but may eventually be found at time Instituto de Geologia, Universidad Nacional de. M16xico.
Typec locality : rruxtelpec, Oaxaca, MN6xico, Miocene.
Though this strongly sculptured cone occurs in the three divisions of the Gatun formnation-but not in the upper~ part in time 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 lby its Ibroad-sh ouldered, biconical outline, and exaggerated growth threads on the anal fasciole and in the channels between the straplike spiral cords. The protoconch and spire whorls are like those of Conus imitato) imitator, except that spiral threads are not discernible on the anal fasciole. One inimnature shell (height 17.8 mumi, locality 138c) shows low tubercles on the spiral cords.
Contrary to the view that was adopted in 1928, C. qaza is treated as a synonymn of 0. inultiliatus mulltilira/us.- Th'ough no specimens of the nomiinate subspecies are avilb, I hse's il1 ustrat ions sh ow the, essential fea,tures of' C. gaza. C.~ imdfltiuats noltiratus is recognizedi in middle Miocene formations in the Tehuantepec area, I unani a, imort-l easteimi and southwestern C'olumbia, ,Jamiaica, the iDominican Republic, and pos*i1)1y in 1cudor aund in delposits of late Miocene age ill hart HINNwesteiim1 P11mauu. A k comparable. form of tme Sp~ecie~s, perhaps the same as that in Ecuador, occurs iii time mi(lhle i\I jocemme of' time. Iari6n area. it. lacks strong growth threads between the spiral cords, but the ab-


356






357


GASTROPODS: EIJLIMIDAEI MARGINELLIDAE TO HELMINTHOGLYPTIDAE


sence of that feature may be a matter of preservation and perhaps should not be overemphasized.
C. mnultilirats walli Mansfield (1925, p. 13, pl. 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 Perfi, identified as C. inultilirat.s gaza, 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 inultiliratus. C. multdsiratus spiekeri Olsson (1932, p. 153, pl. 16, figs. 5, 7), also from the Zorritos formation, is sculptured with heavy tuberculate spiral cords and may be given specific rank. C. tutrbinopsis Gardner (1926-47, p. 361, pl. 43, figy. 12, 1938; Shoal River 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, pl. 4, figs. 5, 6).
C. elarki IRehder 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 fasciole and between the spiral cords. Its outline, however, is turbinate and the shoulder of all the postprotoconch whorls is tuberculate. C. frisbeyae Clench and Pulley (Clench, 1953, p. 369, 1)1. 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, M6xico, northeastern and southwestern Columbia. Bowden formation (middle Miocene), Jamaica. Cercado and Gurabo formations (middle Miocene). Dominican Republic. Limi6n formation (late Miocene), IBocas del Toro area, Panama.

Conus burckhardti burckhardti Bbise
Plate 57, figures 19, 20
Con's biirckhardti Bise, Inst. Geol. MWxico Bol. 22, p. 50, p1. 5,
figs. 39, 40, 1906 (Mliocene, Mwxico). Olsson, Bull. Am.
Paleontology, v. 9, no. 39, p. 224, pl. 3, figs. 4, 5, 1922 (Miocene, I.anami, Costa Rica). Maury, Idem, v. 10, no.
42, 1). 18.7,1)1. :34, fig. 5, 1925 (.Nioceiie, Trinidad). Rutschi, Schweizer. Palmont. Gesell. Abhi., Band 55, no. 1, p. 102,
p1. 4, figs. 3, 4, 1934 (Miocene, Venezuela).
9Uonus barckhardti Bise?, Anderson, California Acad. Sdi.
Proc., 4th scr., v. 18, no. 4, p. 110, 1929 (Miocene, Colombia).


?Conus bitrclclardti. B~se, Alenchaster-Ibarra, in Masson and
Alencaster-Ibarra, Asoc. Mexicana, Geilogos Petroleros Bol., v. 3, p. 210, fig. 25, 1951 (Miocene, Mexico). Alencaster-Ibarra, in Rios -Macbeth, Idem, vol. 4, p. 347 (list),
1)1. 21, figs. 14, 15, 1952 (Miocene, MeN1xico).
?Conus (Leptoconias) bitrckhardti Bise, Perrilliant Montoya,
Mexico Univ. Nac., Inst. Geologia Paleontologla Mexicana, no. 8, p. 26, 1)1. 3, figs. 20, 21, 1960 (Miocene,
Mexico).
Con us tortuoso pun eta tits Toula, Gisson, Bull. Am. Paleontology,
v. 9, no. 39, 1). 54, pl. 3, fig-s. 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, 234- 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 fasciole slightly concave, bearing exaggerated growth threads at fairly regular intervals and faint spiral lineation, or more distinct, though faint, spiral strike. 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.
H-eight (practically complete) 42.3 mm, diameter 17.7 mm (larger figured specimen). Height (practically complete) 33.7 mm, diameter 14.8 mim (small figured specimen).
Type: Instituto de Geologia, Universidad Nacional de M6xico.
Type locality: Tuxtepee, Oaxaca, Me'xico, Miocene.
Thirty cones from 10 localities in the Gatun formation are identified as Conus burckhardti hurckhardti: 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 shlarpness of the shoulder carina, the absence or presence of spiral strip on the anal fasciole, and the development of tubercles on the spiral cords. One shell from the upper part of the Gatun in the eastern area and another






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 C. imitator imitator and C. multiliratus multiliratus.
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 Perrilhiat 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 burckhardti is not a suitable name for the Gatun fossils. Mature shells from the type locality are needed.
Beyond the type region C. burckhardti burekhardti, as identified in the present report, has been recognized in middle Miocene formations in southeastern Costa Rica, Panami, possibly Colombia, and Trinidad, and in deposits of late Miocene age in the Bocas del Toro area, Panami, and Venezuela. Cossmann's illustration of C. marginatus boussaci (Cossmanni, 1913, p. 46, pl. 3, fig. 16) indicates the occurrence of a similar cone in the Miocene of Martinique.
C. harveyensis Mansfield (1930, p. 33, pl. 1, fig. 12; tipper Miocene, Florida) lacks tubercles on the carina of early post-protoconch whorls and on the body-whorl spiral cords. The type-evidently the only specimenis small (height 20.9 mm). The type of C. corrugatus Gardner (1926-47, p. 360, pl. 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 t~o the type of C. 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 burckhardti is more similar to the eastern Pacific species C. tornatus Broderip (Hanna, 1963, p. 39, pl. 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 C. burckhardti burekhardti is greater and its anal fasciolp. is flatter.
C. tortuosopunctatus Toula (1911a, p. 507, p1. 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 formnation (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, M6xico, southeastern Costa Rica, Trinidad. Lim6n formation (late Miocene), Bocas del Toro area, Panam6. Punta Gavilkn formation (late Miocene), Venezuela.
Conus burckhardti harrisi Olsson Plate 57, figure 17
Conus harrisi Olsson, Bull. Am. Paleontology, v. 9, no. 39, p. 53,
p1. 3, fig. 1, 1922 (Miocene, Costa Rica). Rutsch, Schweizer, Palmont. Gesell. Abh. v. 55, no. 1, p. 104, pl. 9, figs.
4-6, 1934 (Miocene, Venezuela). Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research
Inst., p. 88, pl. 11, fig. 6, 1964 (Miocene, Ecuador).
Resembling ('onus burekhardti bu'rckkardti, 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, Lim6n Prov., Costa Rica, middle Miocene.
This slender subspecies of ('onus burckhardti is represented by 22 specimens from the lower and middle parts of the Gatun formation. An exceptionally slender shell f rom 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 C. burekltardti 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. ala quaensis Mansfield (1935, p. 18, pl. 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,


358






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO H3ELMINTHOGLYPTIDAE39


localities 136, 136a, 138a, 138c, 138d. Middle part, eastern area, localities 139c, 139e, 155. Upper part, eastern area, locality 177d (identification doubtful). Deposits of early(?) Miocene age, Ecuador. Middle Miocene deposits, southeastern Costa Rica. Punta Gavildn 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 Conu8 pertvaflus Olsson (1930, p. 40, pl. 5, figs. 13-15), a late Eocene Peruvian species.
Occurrence: Caimito formation, Gatun Lake area (late Oligocene), locality 54h.

Conus tortuosastriatus Toula
Plate 57, figures 5, 6
Conuts (Chelyconuts) tortitosos triatits Toula, K. k. Geol. Reichsanstalt Jahrb., v. 61, 1). 508, p1. 31, fig. 22a, b, 1911 (Miocene, Canal Zone). Gisson, Bull. Am. Paleontology, v. 9,
no. 39, p. 49, pl1. 1, fg. 15, 1922 (Mliocene, Panami).
Govius (11cmicon us) tortitosostria tis Toula, Cossmann, Jour.
Conchyliologie, v. 61. 1). 40,)1 pl. 3,figs. 28, 29, 1913 (Miocene,
Canal Zone).
Not Conius tortutosostriatits Toula, M.Naury, Bull. Am. Paleontology, v. 5, no. 29, p. 41, p1. 6, fig. 9, 1917 (Miocene, Dominican Republic) ;=C. ga?.bi Pilsbry and Johnson.
Conuts tortuoso pun ea tits Toula, Cossmiann, Jour. Conchyliologie,
v. 61, p. 47 (discussion), pl1. 3, figs. 22, 23, 1913 (Miocene,
Canal Zone).
Conuts granozonatus Guppy, Brown and Pilsbry, Acad. Nat. Sdi.
Phila. Proc., jr. 63, p. 341, 1911 (Miocene, Canal Zone). (Janus consolrinus Sowerby, Brown and Pilsbry, Idem (Mliocene, Canal Zone).
Of medium size, slender to inflated, shoulder anoulated 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, tuibercles flattened parallel to shoulder and whorls overlapping to edge of shoulder. Tubercles continuing, in reduced form, on shoulder of body
-whorl. Anal fasciole 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.
H-eight (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 Piios qnetdoides (p. 267), is mnislabelled "Ponton, Dominican Republic." 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 mc.ncthyi 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. qnazei Deshayes has generally been used for that species (Clench, 1942, p. 17, 1)1. 9, figs. 1, 2). The slender, high-spired species C. larvatus Pilsbry and Johnson (Pilsbry, 1922, p. 332, pl. 21, fig. 10; Miocene, Dominican Republic; 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. "7arvats" 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. gracilissim77us Guppy (Woodring, 1928, p. 216, p1. 12, figs. 3, 4; Miocene, Jamaica), C. gabbi Pilsbry and Johnson (Pilsbry, 1922, p. 332, ph. 21, figs. 8, 9; Miocene, Dominican Republic), and C. 8tibarfl8 Woodring (1928, p. 21, p11. 12, fig. 5, Miocene, Jamaica). In addition, C. stibartis has fewer and coarser shoulder tubercles.
Occurrence: Middle and upper parts of Gatum formiation (middle Miocene). Middle part, eastern area, Gatin Locks site (Toula's and Brown and Pilsbry's records). Upper part, eastern area, localities 175, 1T6, 177, 1I7b. Chagres sandstone (late Miocene), locality 208. Lim6n formation (late Miocene), Bocas del rToro area,, Panama.


368-278 0 70 5


359







GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Family TURRIDAE
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 b *ringing 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 subgreneric rank: Polystira, Fusiturricula, Cryoturris, Sacckarotrrs, MiraclathurelIla, 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 mnangilenes and daphunellines, occur in the Gatun formation.
Species and subspecies of turrid gastro pods


Subfamily


Turrinac -------------Turriculinac ----------Borsoninac ------------Clavinac -------------Mangeijinae ----------Daphnellinae ---------Total ----------


EoCelle or
011go
cene


Oligocene


C' 0q


1


1


c
2
3


2



7


Miocene


C C)
C C) .5
0

1


C C)
a
C C)
C C)

1
1


2


1 4


4


4
1
20 1s

50


t 10
4

7-


Three of the species listed-one from the Culebra formation and two from the La l3oca formation-are merely mentioned. In addition to those listed, all the formations, except the Gatun and Chiagres, contain unidentifiable turrid material.
Subfamily TURRINAE
Genus Gemmula Weinkauff

v. 2, p. 287, 1875.


Type (logotype, Cossinann, Essais paleloconchologie compare lpt. 2, p). 62, 1896): Plcurotorna gcnimata Hinds 1P.
1/cniinuta Reeve, 1843, not P. gcnata Conrad, 1835,Genniua hindsianu Berry], living, Baja California and
Gulf of California.
The type species has been illustrated by Harris (1937, p. 10, pl. 1, figs. 33, 33a) and Keen (1958a, p. 250, p1. 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 ait the present time, the type species survives in the eastern Pacific Ocean and G. pe7i seelida (Dall) (1889, p. 74, 1)1. 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 G. cf. 0. va'ningeni.

Gemmula cf. G. amica Casey
Small, slender. Protoconch missing. Peripheral band coarsely geminate. 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 Cemmida is represented in the shallow- water facies of the Caimito formation of the Gatun Lake area by four specimens. All aire incomplete and their shell material is replaced by calcite. This species is larger and more strongly sculptured than C. ainica (Casey, 1903, p. 270; Harris, 1937, p. 13, pl. 1, figs. 26-28), of the early Oligocene Red Bluff clay of Mississippi.
Harris and Palmer (1946-47, p. 420, 1947) assigned G. amica to Coronia (Gregorio, 1890, p. 23; type (logotype, Cossmtanii, 1896, p. 78) : Pleurotoma acutirostra Conrad, Eocene, Alabama), which they treated as a sil)fenuis of Cevo nida. Pleurotoina acuttirostra was idlentified by H-arris (1937, p. 12, pl. 1, fig. 22) as a forin of Piettrotomna children i Lea. Though C. ainica, lik e other Oligocene and Eocene species of Cemmula, is smaller than the late Trertiary and living species and has a shorter aperture, its strong and strongly geminate p~eriphleral band favors assignment to Cemnuda.
Occurrence: Caimito formation, Gatun Lake area (late Oligocene), locality 56.


360






GASTROPODS: EIJLIMIDAE, MAR GINELLIDAE TO I3ELMINTHOGLYPTIDAE36


Gemmula vaningeni (Brown and Pilsbry)
Plate 57, figures 11, 18, 22; plate 65, figure 1
Piro torna (Gcm mula) van ingeni Brown and Pilsbry, Acad.
Nat. Sd. Phila. Proc., v. 64, p. 505, pl. 22, fig. 4, 1913 (Miocene, Canal Zone). Cossmann, Jour. Conchyliologie, v. 61, p). 19, p1. 2, figs. 21, 22, 1913 (Miocene, Canal Zone).
Mansfield, Florida Dept. Conservation, Geol. Bull. 12,
1). 10, 11 (lists), 1935 (Miocene, Florida).
Turrjs (Gcrnrula) vaningcn i (Brown and Pilsbry) Olnomikado, Geol. Soc. Japan Jour., v. 46, p. 624, p1. 29, fig. 14, 1939 (Miocene, Colombia). Marks, Bull. Am. Paleontology, v. 33, no. 139, p). 126, 1)1. 8, fig. 11, 1951 (Miocene,
Ecuador).
Gcmnzula vaningeni (Brown and Pilsbry), Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst., 1p.93, 1)1. 20, fig. 12, 1964 (Miocene, Ecuador). Ilcriplcurotorna cileta Gardner, U.S. Geol. Survey Prof. Paper
142, p. 290, p1. 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 1129 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 geminate 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 mmn largestt figured specimen).
Type: Acad. Nat. Sci. Phila. 3855.
Type locality: Gatun Locks excavation, middle part of Gatun formation.
Though Gemmula is rare in tile Miocene deposits of tile Caribbean region, 0 'cavivqeni is represented iii 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. TJwenty-six of these fossils show the protoconch, but on many of them at least some of the protoconch whorls are somewhat worn.
Two imi-mature shells are labeled, in Dall's writing, types of Pie urotoma povtonensis Dall and are alleged to have been collected by "Bland ?" at Ponton, in the Dominican Republic. This record, like that for the type lot of Plios metidoides (p. 267), is presumed to be ain erroneous record for the present Mount Hope. If the rejected record is ignored, Geinmula is unknown in the Dominican Republic. Pleurotoma pontoneiisis, mentioned by Mansfield (192-5, p. 15) and Gardner (192647, 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 17t5 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 siphional 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 (1)1. 57, figys. 11, 18). The anal sinus of the smallest of these shells is partly covered by the attached valve of an anomid (?). The attachiment, 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 malformatioii is shown by other turrids. Those that have come to my attention are as follows: species in the genera Cenmm/lda (Glibert, 1954, p. 7, p1. 2, figs. 3a, b; MacNeil, 1960, p. 104; Powell, 1964, p. 253) ; Pi gidgemmnula (MacNeil, 1960, p. 103, 104; Powell, 1964, p. 277, 279, pl. 215, figs. 5, 6) ; Ptychosyrinx (Powell, 1964, p. 2-89, 290, 291, pl. 223, fig. 1, pl. 224; Okutani, 1964, p. 425, pl. 4, fig. 9) ; Aforia (Dall, 1918, p. 319) ; and a species assigned to D7 11ia (Melvill, 1917, p. 150). Though various explanations for the malformation have been proposed, its significance, as poiiited 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 Dali, who later (Dall, 1921, pl. 11, fig. 6) illustrated a specimen of Afovia showing the anteriorr sulcus."
liipleurotoma eileta, of the Shoal iRiver formatioii of Florida, is considered to be a synonym of C. vaningeni. The largest Florida fossils are a little smaller than Gatun shells aiid some, including' the type, have a slightly narrower and slightly less coarse-


361






362 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


ly greminate peripheral band. Incomplete shells indicate a mnaximium 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 C. '?aningeni. 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 IBrasso formation of Trinidad (Glob orotalia fobsi 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 lBrasso formation (USGS 9219) were identified by Mansfield (1925, p. 15) as Turri8 vain )gemi var. qnachapooren8s (Maury). Their relatively low peripheral band and subsutural collar suggest Maury's (1925,1p. 191, 1)1. 32, figs. 1, 14) [unnumbered on plate] Drillia vaflingefli var. 8avotiandrew. The status of Mansfield's and Maury's specimens is uncertain.
Illustrations of G. ldcsiava, the type of the genus, suggest that it is smaller and more slender thanl G. vaningeni, and has a narrower and lower sutural collar. Nevert]heless G. ,vnigeqi is much more closely related to G. llindsiagia than to 0. 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. Dall's record, "near Monosquillo," was based on a cataloger's error in transcription.
Occurrence: Lower, middle, and upper parts of Gatiin formation (middle Miocene). Lower part, locality 138c. Middle part, eastern area, localities 147b, 147g, 147h, 1,51, 155, 155a, 155b, 159d. Upper part, eastern area, localities 173a, 175, 176, 177, 17Th, 178. Middle Miocene deposits, Dari6n 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 lBrasso formation, Trinidad. Shoal River formation (middle Miocene), Florida.

Gemmu-tla mnachapoarensis (Maury)
Driflia vaningcni var. macha poorcns 'Maury, Bull. Ain. Paleontology, v. 10, no. 42, p. 191, pl. 32, figs. 5, 9, 1925
(Mliocene, Trinidad).
Turris brasswnsis 1Aiansfleldl, U.S. Natl. Mlus. Proc., v. 66, art.
22, 1). 14, 1)1. 2, figs. 7, 8, 1925 (Mliocene, Trinidad).
Iicmiphurotonma bitropis Gardner, U.S. Geol. Survey Prof.
Paper 142, 1). 290, p1. 38, fig. 29, 1938 (MNiocene, Fin.).
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. B~othi edges of peripheral band of post- protoconch whorls raised to form distinct spiral threads. Subsutural primary spiral forming a strong, wide-bsd narrow -crested collar. Exaggerated, uniformily 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 form-ation].
The only specimen, from the lower part of the Gatun formation, is immature. Though the lprotoconch whorls are partly corroded, the features mentioned in the description are discernible. Despite the unpromisin g 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 subsmtural collar. Also as shown by growth lines on shells from Trinidad and Florida, the anal sinus is typically that of Gemmula.
G. mnachapoore)?sis evidently is a small species. Maury estimated 19 mim 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 ITSNM\ 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 Glob orotalia fo~isi zone of the Brasso formation yielded all this Triniidad m-aterial (USGS localities 8302, 9212, 19856, 1987-0, 21234). According to H-. G. Kugler, who supplied the zonal data for these localities, and also for 21230 cited under G. vaningeni, the areal ge ology


362






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO IHELMINTI{OGLYPTIDAE36


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. vani2gefli and C. 9nachapoorensis are associated at such widely separated localities as Trinidad, Panaind, and Florida.
Occurrence: Lower part of Gatun formation (middle Mioene), locality 138c. Early middle Miocene part of Brasso formation, Trinidad. Shoal River formation (middle Miocene), Florida.
Genus F
"Gemmula"l species
Plate 48, figures 19, 20
Small, moderately slender, biconic. Protoconch missing. Peripheral band wide, coarsely geminate. Suture bordered by a geminate collar, less coarsely geminate than peripheral band. Other spiral sculpture consisting of narrow primary and secondary threads, faintly noded by conspicuous, exaggerated growth threads. Aperture long, narrow. Siphional canal virtually absent; aperture ending in a, shallow siphional notch. Siphional fasciole slightly inflated. Anal sinus moderately deep, apex UJ-shiaped, located on peripheral band.
Height (almost complete) 20.7 mm, diameter 18.2 mm figuredd specimen).
This unnamed species is represented by a defective specimen and a minute tip from the moderately deepwater facies of the Caimnito formation on Barro Colorado ]Island. It is to be assigned to an apparently unnamed turrine genus that has a Genmqda-like anal sinus and geminate peripheral band, but, unlike Gemnmula. has a long, n arrow aperture, siphional notch, and siphional fasciole. Though the anterior end of the aperture is defective, growth lines show the shallow siphonal notch. The g-eminate sutural collar is a noteworthy feature. The outline suggoests Scobhvella or a small slender Rath ytoma.
Occurrence: Caimito formation, Gatun Lake area (late Oligocene), localities 54g, 54j.

Genus Pleuroliria Gregorio
(iregorlo, Aiales, Gcol. lPaN-mitologie, p~t. 7,1p. 38, 1890. Type (orthotype) :P~curotovia supramirifica Gregorlo, locality
not speified (presumned to be Plemrotoma cochlcaris Conrad, Oligocene, mississippi).
Should Gregorio's type designation be considered equivocal, Cossmann unequivocally cited Pieurotogna suprarniri/ica as the type four years later (Cossmiann, 1893 (1894), p). 43).
Harris (19.37, p. 8, pl. 1, figs. 5, 5a, b) reproduced Gregorio's illustrations of Pleurotoma supramnirifica.


There is no reasonable doubt that it is the Oligocene species Pleurotogna cocleari8. The known Eocene species are inuch smaller.

Subgenus Polystira Woodring
Woodrig, Carnegie Iiist. Washing-ton Pub. 385, p. 145, 1928. Type (ortliotype): Plcntrotonia albida Perry, living, southern
Florida, Gulf of _Mexico, and WVest Indies.
Bartsch (1934, p. 8) claimed that Pleurotorna albida is a western Pacific species and that Miurex 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 Zealaind and Lord Howe's Island" (Perry, 1811, explanation of pl. 32, fig. 4), his illustration is considered to be a reasonably satisfactory representation of the western Atlantic species later named Mlurex v2i.rgo. 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 Pleuroliq a. It is interpreted to be the direct descendant of Pleuro7ha 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 numher 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 Pol?/stira, 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 namned-a procedure that has been justly criticized (MacNeil, 1960, p. 100).
Both Pie uroliria s.s. and Polys tira embrace American species. Plenroliria s.s. is of Eocene and Oligocene age. The agre range of Polystira is early Mliocene to the present time and the living species occur in western Atlantic and eastern Pacific waters. They aire 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 Pleuroliq a.
Pleuroliria albidoides (Gardner) (1926-47, p. 287, pl. 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 Dali (1915, p. 38, p1. 5, fig. 13) as Turn is al1bida without any indication that the specimen is a, Chipola fossil, not a Tampa fossil. The specimen hie illustrated


363






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 protoconchPli uoliria tenaqgas' the next species described was designated as the type of Jose phina (Gardner, 1945, p. 246), proposed as a section of Pie uroliria. That proposal followed my suggestion (Woodring, 1928, p. 145), which overemphasized the distinction between blunt and acute protoconchis. The protoconch illustrated by Gardner (1926-47, p. 288, pl. 38, fig. 23, 1938) as the protoconch of Polystira virgo is that of a small shell (height 11.6 mim) dredged off Grenada ait at depth of 73 fathoms. It presumably is one of the relatively small living West Indian species, for which six names have been proposed.
Lophiotoma (Casey, 1904, p. 130; type (logotype, Woodring, 1928, p. 146) : Plettrotoma tigina Lamarck
(=P. actta 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, pl. 173), Poliystira 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 Pleurotoina ailbida var. tellea (Dali, 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 sinushas been illustrated recently (Abbott, 1954, p. 268, pl. 13, fig. in), 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, fig-ures 12, 23, 24; plate 65, figure 2
Pleurotoma (ilida Perry, Brown and Pilsbry, Acad. Nat. Sci.
Phila. Proc., v. 63, p. 343, 1911 (-Miocene, Canal Zone). Turi8s albida. (Perry), Olsson, Bull. Am. P~aleonitology, v. 9,
no. 39, P. 58, pl1. 4, figs. 1, 2, 1922 (Mliocene, Panaind).
Plcurotowa haitonsi8 Sowverby, Cossmnann, Jour. Conehyliologie,
v. 61, pl. 16, p1. 2, figs. 1-4, 1913 (M.Niocene, Canal zone). Polilstira (Plcuroliria) t('nagos0 Gardner, IJ.S. Geol. SuIrvey
P~rof. Paper 142, p. 288, 1)1. 38, figs. 25, 26, 1938 (Miocene,

?Plcuroliria tcnagos (Gardner), Gardner, Geol. Soc. America
Mlei. 11, p). 246, 194.5 (Miocene, MeN1xico).
Of medium size to moderately large, moderately slender to moderately inflated. Profile of late whorls slightly to dec idedl y iodli fled by peripheral carina. Protoconch acute, 3- to 4-whorled (generally 3 to 3 /). Last whlorl, 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.
H-eight 58.8 mm, diameter 16.6 mm (largest figured specimenn. Estimated restored height 75 mm, diameter 20.8 mnm (largest specimen).
Type: USNMT 351134.
Type locality: USGS 3742, Shell Bluff, Shoal River, Walton County, Fla., Shoal River formation.
Pleuroliria tevaqos 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 1.38c) and 24 in a miiddle Gatun collection (locality 161a). The largest specimens occur in the upper part of the formation in b)0th eastern and western areas. Though they are incomplete, they indicate a heighlt 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 th)e strength of the subsutural primary spiral thread, are vari able. 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 (p1. 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 Giatlin specinieil of plate 5-7', figure 23. Trhe fossils of the three parts, however, do not uniformly show a ,graded profile series. Ten of the 14 smallest shells (mfaximumiii heighit 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 spediincus froin the u1 )1)eL part of the G'atun in the western ai lea hias a jpreser\'ed lprotoconcli. A minute middle Gatlin shell eightht '5.2 11uu1) the only shell from locality 1(1 (-has a 1)luut lprotoconch of two whorls, the last~ half whiorl of which hears axial riblets. Thils fossil


364






GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE36


At present it is uncertain what species other than Pleurotoma ion girostropsi8 are to be referred to the subgenus P/euro/usia s.s. The protoconch of the type species is unknownv, and some post-protoconch features appear to be unique. Eocene species of Pleuro/usia hame been described by H-arris (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; Pleuro toma collavis Casey (1903, p. 270; H-arris and Palmer, 194647, p. 4301, pl. 59, figs. 13-15, 1947), for example, seems to be a clavine species. The lprotoconch of not all of the long-pillared Eocene species have been described. Those that are known are blunt, smooth, and about 2-whorled.
Pleuro fusia servata (Conrad) (1847 (1848), p. 284; H-arris, 1937, p. 50, p1. 10, fig. 4, not fig. 5; Harris and Palmer, 1946-47, pl. 59, fig. 21, 1947), from the Oligocene Byram mnarl, has an acute, 312- to 4-whorled protoconch. rTle later part of the last whorl bears four to six axial riblets. Pleurofusia olivia (Casey) (1903, p. 269), from the Oligocene Red Bluff clay, has a similar protoconch. This type of protoconch was assumed to be typical for Pleuro fusia when Fyusiturricula (Woodring, 1928, p. 165; type (orthotype) :Turns8 (Surcula) fusing el/a 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 Fusitrricula, which has a blunt or moderately acute, 1 /- to 234whorled, smooth protoconch (last half whorl rounded or slightly bulging), is uncertain, Onl accou-nt of the uncertainty concerning Pleurofitsia 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 Fusiurricu/a. 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 (Blartsch) (1934, P. 13, p1. 3, figs. 1, 2, 10; height 25 mmi), a similar, thinshelled, deep-water species.
P. enae is represented by eight specimens, taken in hiauls at five stations on the south border of the Puerto Rico Trench, off Puerto Rico, and the Virgin Islands, at depths of 240 to 360 fathoms. Three of these thin shells, including the type, as shown in Bartschi's figure 2, have a perfectly preserved anal. sinus, showing the wide forward sweep of the lower limb. Until this species and P. fe2'norei (Bartsch) (cited under P. cf. P. fenirnorei) were described, Pleuro fusia was unknown in the western Atlantic Ocean.


Subgenus?
Pleurofusia species
Plate 48, figure 27
Of mnediumn size, apex and pillar broken. Sculpture
conistng f wdelow axial swellings (four on pen-Llt whorl), overridden by two modern tely 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 oil depressed subsutural. area.
H-eight (incomplete) 18.7 mim (estimated restored height 30 m-m), diameter 9 mm (figured specimen).
The description of this unnamed species is based on all 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 Gatuin species P/euro fusia fusin us. but lacks secondary spiral threads. Its protoconch and outer lip are, of course, unknown.
A poorly preserved fragment of two whorls from the La B~oca formation is listed as P/etro fusia? sp.
Occurrence: Caimito formation, Gatun Lake area (late Oligocene), locality 56.
Pleurofusia acra Woodring, ni. sp.
Plate 57, figure 8; plate 65, figure 3
Of medium size, Fusivus-like in outline and gross sculpture, pillar long and slender. Protoconch acute, 131/9-whorled, last two whorls carinate below middle of whorl. Post- protoconch whorls bearing closely slpaced, wide axial swellings below anal fasciole (8 oil body whorl), disappearing oil body whorl below shoulder. Axial swyellings overridden by spiral threads (5 on lpenult whorl). Similar threads oil body whorl below lower end of axial swellings undulated by low, elongYate nodes, except near end of pillar, where tihe threads are smooth. Anal fasciole 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
1.5 mm11 (type).
Trype: TTSNM 645764.
Type locality: 155c (USGS 16915, Gatun Third Locks excavation, east side of excavation, 1 m1-ile (1.6 kin) north of Gatun Lake, Canal Zone), middle part of Gatun formation.
Though the type of this species, which may be immiature, is the only specimen, it combines three un-


367






366 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Subfamily TURRICULINAE
Genus Zemacies Finlay?
Finlay, New Zealand Ti)st. Trans., v. 56, p). 252, 1926. Type (orthiotype): Zcmacies clatior Finlay, Miocene, New
Zealand.
The following' two early Tertiary turriculine species are noteworthy for their slender outline, long( narrow aperture, and fiat, closely spaced, narrow spiral bands. They represent Zcmacics or a related genuis. Through both evidently are new species, the material is too imperfect to name. In New Zealand Zeinacics has an age range of Paleocene to Pliocene (Powell, 1966, p. 36).
Zemacies? species a
Plate 48, figure 13
Of medium size, slender, sulbaliulated at shoulders, Protoconcli corroded, apparently consisting of abot three rapidly enl arging -whorls. Shoulder bearing low, ~xally elongate swellings, proo-ressively more subdued Oil late whorls. Spiral sculpture consistiug- of fiat, closely spaced, narrow bands. Subsuitural band niarrow, an adijoinig- lbaiid wider. Anal sinus apparently onl narrow, concave space between shoulder and suiture. Aperture lou g,, iiarrow.
H-eight (incomplete) 19.3 mmi (estimated restored height 27 m-m), diameter incompletet) 8.5 mmn (estimated restored diameter 9 mm).
The inarne member of the IBohio (?) formation at Vamos Vamios yielded two imperfect specimens of this species.
Occurrence: Marine inemlber of Bohiio (?) formation (late Eocene or early Oligocene), locality 40d.

Zemacies? species b
Plate 48, figure 21
Of mediumn size, slender, late whorls subang' ul ated at shoulder. Protoconchi and early post-protoconch whiorls missing. L-ate wI ion s beariug(- wi del y sp acedl, obliquely elong-ate nodes onl shoulder. Spiral sculpturle, consisting- of flat, narrow, closely spaced bands. Suibsutural band relatively wide. Anal sumis located onl narrow, concave space between shoulder and suture. Aperture long, narrow.
FHeighlt (incomplete) 32 mmi estimatedd restored height i18 mmi) diameter (incomplete) 11.6 mim (estimiated restored diameter 13 mnm).
This strongly iioced species is represented by an incollete specimen from ti e moderately (leel)-water facies of the Calimito formation onl Barro Colorado Island.


Occurirence: Calinito formation, Gatun Lake area (late Olio-ocene). locality 54n.

Genus PleurofuLsia Gregoria
Gregorio, Aniiales Gaol. PaI~ouitologile, pt. 7, p. 33, 1890. TypIe (ortliot ype) :1(107to)il (Pleurofusia) 7ongirostrops?.3
Gregorio, locality :11)(l age not specified.
Tlioiigh (regorio (didl not specify aI locality or age for1 P/cuovt()mna /0 /i(//i osbop.4., hie presuimably thioughit it is a Claiborne species, that is, according to p)LC5Clt terminology, a species trom the middle Eocenie (4osport sand(i t Clailborne, Alabama. Dr. K. V. W. 1Pahmuer, D~irector of the Paleontological Research Institution, informs me that the Institution has received from the University of Palermo (the repository of Cregorio's collections) three specimens labelled P/citma /0') gb os/tiop.s, s ut none resembles the Mistraton 1llllisiedl by G'regorio as plate 2, figure 26. Hiarr~is, whio reproduced Greglorio's figure 26 (lHarris, 19.37, p). '5l,Ipl. 10, fig. 6) was unable to identify tis species. N_\o species closely resembling Gregorio's Mistratin is known to occur at Claiborne or elsewh-lere in the Eocene or Olig(ocenie deposits of the Southeastern states. (+regori o, of course, may have had a rare species or his drwn ly be faulty. A minute specimen In the IJSNM collection from Claiborne, both ends of wich are, broken (hieighlt 4.9 mmi) has the anial sinu in a strongly constricted sutural area marked with ,onisp~icuous growth threads, as shown in
Grgros iue but, lacks the two strong eipea
spiral threads onl spire whiorls and the subangulateci early wi ion s of that illustration. Casey (1904, p. 152) tlhoiigh-t that "a specimen in the cabinet of iMr. Aid.. richi, from the o-wei' Cl aiborne" greatly resembles Greg-orio's illustration. A turrid in the Aldrich collection label led "P/c uroftisia loon girostropsis Greg., per Casey, Lee Co., Texas, IL. C., Eocene," doubtless is the specimen (Casey biad in mind. It is not the species represented by (Aregroriols ilutrton regorio compared Ills species within P/ci'tloma sc)viwta Conrad, ,an Olioocei(e, spec(ies froin the Bvram miarl, cited in the third I l lowiug- panagrai)li ideed, despite the lack of sumi1 a cit v, lie thiough-t It iay lbe a v-ariety of the Olioyouele Species.
It is iinlortiuate, that, nothing more than Gregorio's drawing i av-ailaible for the type species of P/ewrofu.\i. Neerthleless, it is prop~osedl to treat the namle
asthiamie for a, genus eunhracnvr- small to moderatel ,v larg-e tuir cuulinc turirids that have a FitsinusIlike, oiutlinle and sciil ptiire, andl an asymmnetrical sinus ad joiniuuz, the stuire. Sitcl species are found in AmenlcauI fa uuias sice ELoceie time, and now live in western Atbiutic and eastern Pacific waters at low latitudes.


366






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE37


At present it is uncertain what species other than Pleurotomna ion girostropsis are to be referred to the subgenus P/euro fusia s.s. The protoconch of the type species is unknown, and some post-protoconch features appear to be unique. Eocene species of P/euro fusia have been described by Harris (1937, p. 50-53) and H-arris and Palmer (1946-4T, p. 429-431, 1947). Whether the species that have a short pillar should be referred to that genus is debatable; Pleurotoma co//aris Casey (1903, p. 270; Harris and Palmer, 19464T, p. 4301, pl. 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.
P/euro fusia servata (Conrad) (18471 (1848), p. 284; Harris, 1937, p. 50, pl. 10, fig. 4, not fig. 5; H-arris and Palmer, 1946-47, pl. 59, fig. 21, 1947), from the Oligocene Byramn marl, has an acute, 312- to 4-whorled protoconch. The later part of the last whorl bears four to six axial riblets. P/euro fusia olivia (Casey) (1903, p). 269), from the Oligocene Red Bluff clay, has a similar lprotoconcli. This type of protoconch was assumied to be typical for P/euro fusia when Fusiturricula (Aloodring, 1928, p. 165; type (orthotype) :Turrns (Surcula) fusing el/a Dali, 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 Fyusiturni cu/a, which has a blunt or moderately acute, 112- to 23/4whorled, smooth protoconch (last half whorl rounded or slightly bulging), is uncertain, on account of the uncertainty concerning P/euro fusia s.s. If it is to be retained, it rates subgeneric rank under P/euro fusia. Trhe type still is the only dredged specimen of tihe type species of Fusitri wi/a. It is a small, thin-shelled, deep-water shell dredged at a depth of 153 fathoms. Though it mnay be immature (height 17.2 mm), it is not much smaller than P/euro fusia e'na3 (Bartsch) (1934, P. 13, pl. 3, figs. 1, 2, 10; height 25 mm), a similar, thinshelled, deep-water species.
P. ena, is represented by eillt specimens, taken in hiauls at five stations on the south border of the Puerto Rico Trench, off Puerto Rico, and the Virgin Islands, at depths of 240 to 360 fathoms. Three of these thin shells, includin- tile type, as shown ill Bartsch's figure 2, h ave a perfectly preserved anal sinus, showing tile wide forward sweep of the lower limb. Until this species and P. fevimorei (B~artsch) (cited under P. cf. P. fenimorei) were described, P/eurofusia was unknown in the western Atlantic Ocean.


Subgenus?
Pleurofusia species
Plate 48, figure 27
Of medium size, apex and pillar broken. Sculpture cossiigo wdlow 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, aild corresponding threads on base of body whlorl alulost as strong~ as those oil periphery. Subsutural thread and that at base of subsutural area narrower. Anal sinusi oil depressed subsutural area.
Height (incomplete) 18.7 m-m (estimated restored hieight 30) mm), diameter 9 mim (figured specimen).
The description of this unnamed species is based on an incomplete fossil, replaced by calcite, collected from tile shi(al low-water facies of the Caimito formation in the Gatin Lake area at locality 56. In general features it resembles the Gatiin species P/euro fusia f usin'1us. 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 P/euro fusia? 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 inedium size, Fushins-like in outline an~d gross sculpture, p~illar~ long and sleilder. Protoconch acute, 312-wiiorled, last two whorls carinate below middle of whorl. Post-protoconcil whorls bearing closely spaced, wide axial swellings below anal fasciole (8 on body whorl), disappearing on body whorl below shoulder. Axial swellings overridden by spiral threads (5 on penult whorl). Similar threads on body whlorl below lower end of axial swellings undulated by low, elongate nlodes, except near end of pillar, where tile threads are smooth. Anal fasciole 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.
H-eight (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 excavations, 1 mile (1.6 ki) north of Gatun Lake, Camlal Zone), middle part of Gatun. formation.
Thuhthe type of this species, which may be immlature, is the only specimens, it combines three un-


367







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. P/eurofitsia fiheila Dal] (1908, p). 261, pl. 14, fig. 7, living, Gulf of Panama), the type of Fusiturricula, has an acute, p)olygyrate lprotoconcli. Its last half whorl bulges at the periphery, but is not sharply carinate. The retractiN-e subsutural swellings and the faintly noded spiral threads on the lower patrt of the body whorl of P. aera appear to be unique among American fossil and living species of Pletro fusia.
Occurrence: Middle part of Gatun formation (middle Miocene), eastern area, locality 155c.

Pleurofuasia cf. P. fenimorei (Bartsch)

Two incomplete shells from the Chiagres sandstone represent an undescribed species of Pleuro fusia. The pr'otoconcl i, early lpost-protoconcll whorls, and end of th~e pillar are missing on bothi. The periphery of the whorls is strongly inflated and the anal fasciole corresponding1 y 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 th reads. Thle estimated restored hieight is between 25 and 30 m-m, the diameter 10 mm.
Trhe whorl outline and its few, strong, widely spaced swellings suggest Pie urofusia fenimorci (B~artsch, 19r34, p). 7i, p1. 2, figs. 4, 5), a large species (height 71 mmn) dredged on the south border of the Puerto Rico Trrenlch 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 stron(yer. The sam-e dredge haul yielded another shell of P. fe)?nioiei that is smaller than the type, but in better condition than the somewhat worn type.
1P. feimore; is the type of Fusisyi-inx (Bartsch, 1934, p). Ti). The only essential difference between Fusi8yiias now known, and the type of F'usitutriecuia (cited in the discussion under Pleurofusia) lies in the number, width), and spacing of the axial ribs or swellBUNs. The protoconch) of Fusi yinvx, however, is unknown.
Occurrence: Chiagres sandstone (late Miocene), locality 208.

Subgenus Cruziturricula Marks

Marks, Bull. Ain. Paleontology, v. 33, no. 139, p). 129, 1951. Type (orthiotype): qurrieula (Plciirofusia) cruziana Olsson,
Miocene, Perd, Ecuador.
('rzitu ienlais distinguished from other groups of species assigned to Pleuiro ftsia lby its blunt or acute protoconchi, thle last wvhorl or less of which is carinate below the middle of the whiorl and bears fine arcuate axial riblets abov-e the carina. Trle axial sculpture consists of swellings rather than ribs and is present or absent, onl late whorls. As shown in the following tall thle fossil species have a lirate outer lip, whereas the lip) of the living species is nolirate. Also as shown in the table, (ritrrdahas an age range of Oligocene to present time and survives in the eastern Pacific Ocean.


Species of Pleurofusia, sub genus Cruziturricula


Ago and locality



Living,
Panamnic province


Middle and late
Miocene, Trinidad

Middle Miocente,
IPauaind

Middle Mioeene,
Ecuador

Early Miocene,
Ecuador; middle
Miocene, Per6

Early Mioeene,
Florida

Oligocene,
Mississippi


Species


Picurofusia panthca (Dali) P. arcuata (Reeve) P. fusinus sanctidavidis
(Mau ry)

1). fusinus fusinus (Brown and
Pilsbry)

P1. latira (Olssott) (=P. fusinu s fusinus) 1'. cruziana (Olsson)


P. glypta (Gardner) P. vicksburgensis (Casey)


11rotoconch


Unknown, except later part of last whorl; carinate, axial riblets above marina. Blunt, 2-whorled; last whorl earinate, axial riblets
above catitta.

Blunt, 1 '- to 2-whiorled; last 11 to I% whiorls marinate;
axial riblets above marina.

Blutnt, ib-r to 2-wliorled; last h4 to I whorl carinate;
axial riblets above Caritna.

Binunt, 2-whiorled; last, half wh-lorl carittate, axial riblets
above Carina.

Trip) brukettn, evidettl v bluint, about, 2-NNliorled; last 14
whiorl carinat e, axial riblet8; above Cartina.

Blun11t, i%2-wliorled; last /' whiorl caritiate, axial riblets
above caritta.
Acute, 3- to 4-whiorled; last )~to 4I whiorl carinate,
axial riblets above marina.


Interior of outer
lip

Nonlirate. Noillirate. Lirate. Lirate. Lirate. Lirate. Lirate.

_L ir at e._


368






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO IHELMINTHOGLYPTIDAE36


Marks assigned the two living Panamic species to Crazituri-icula, but under erroneous names. His "Turricila" arcuata Dali evidently is ain inadvertent error for "Reeve": the species described by Reeve (1843, pl. 3, fi.15, and accompanying text) as Pleurotoma aqcuata. For "Turricula" laviqiia Dali, Marks intended to mean Turr icula (Surcida) pantliea Dali (1919, p. 4, pl. 1, fig. 6 [not fig. 5 as printed]), not Turi~cula (Saua) lavi'nia Dali (Iclem, p. 4, pl. 1, fig. 5 [not fig. 6 as printedd]. Marks was misled by errors in Dali's publication, involving transposition of figure numbers on page 4 and in the explanation of plate 1. Turr cula lavim .a is a clavine species, not turriculine. Marks was not the first paleontologist to be misled by these errors. Olsson (1922, p. 55, p1. 4, fig. 6) named a Miocene Costa Rican species as Turricula lavioides on account of the supposed relation to the alleged Turiciula lavinua. The Milocene species, however, is a species of the living- Panamic grenus Ifnefastia. I am indebted to Dr. K. V. AV. Palmer for forwarding Marks' illustratecd specimen of Cruziturricula craziava, and to Dr. L. G. Hertlein for forwarding the shell illustrated by Marks as Cruziturricula arcuata (Dali).

Pleurofusia (Cruziturricula) fusinus fusinus (Brown and Pilsbry)
Plate 57, figures 16, 21; plate 65, figure 4
Driflia fusinus Brown and Pilsbry, Acad. Nat. Sci. Phila. Proc.,
v. 63, p). 344, pl. 23, fig. 7, 1911 (Mliocene, Canal Zone). Fusitatrricula latira Olsson, Neogene mollusks f rom northwestern Ecuador, Paleontolog-ical Research Inst., p). 94, pl. 20,
fig. 4, 4a, 1964, (-Miocene, Ecuador).
Moderately large, Fusinu8-like in outline and sculpture; pillar long and slender. Protoconch blunt, almost flat-topped, 134- to 2-whorled, last 14 to 1 whorl carinate below middle of whorl, bearing fine, arcuate axial riblets above carina. End of protoconch marked by strengtheingi, of carina to form lower peripheral spiral thread, and appearance of upper peripheral and subsutumral spiral threads, and closely spaced, wide axial swelling-s. Upper p~eriphleral 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 pinlary 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 whiorls. 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,


asymmietric; upper limb short, lower limb long. Interior of outer lip lirate.
He-Tit (incomplete) 45.71 mm, diameter (incomp)lete) 13.5 mim (larger figured specimen). Estimated restored height 60 mm, estimated restored diameter 16 mm11- (largest specimen).
Type: Acad. Nat. Sci. Phila. 1693.
Type locality: Gatun Locks site, Canal Zone, middle part of Gatuni formation.
Variation, on a minor scale, affects the strength of pimiary spiral threads with reference to the two on the periphery and the strength of axial swellings Onl 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 iDari~n (USGS 8433) and Chiriqui (UTSGS 71955).
TuisI alb ida saitctidanvdis (Mlaury, 1925, p. 188, 1)1. 32, fig-s. 6, 7), which is widespread in the late middle Milocene Sami Jos6 calcareous silt member of the i\anzanilla formation of Trinidad, is classified as a scarcely distinguishable subspecies of P. jusinus. Fifty specimens are represented in 14 USNMI collections. It IS considerably snialler than the nominate subspecies, as the miaximumiii height ranges from 40 to an estimated 45 mmn. 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 ninti whorl, are indistinguishable from immature shells of the nominate subspecies. P. fusinus sancti(aiacdis occurs also in the late Milocene Melajo clay mnemiber of the S pring-vale formation of Trinidad (USGS 21178, 18634). On the four specimens that show the early sculpture the swellings disappear on the third or fourth whorl.
Thle specimen illustrated by Mi~aury in her figure (Pal eontol ogi cal Research inst. 1008) is herewith selected as the lectotype.
Except -for its blunt, irotocomlch of fewer whorls, larger size of shell, and larger apical angle, P/euro-


369






370 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


f usia fusinws fusnuts resembles the Oligocene species lPlettrotoina 'eicksburgensis Casey (1903, p. 268, Harris, 1937, p. 51, pl. 10, fig. 3; Harris and Palmer, 1946-47, 1)1. 59, fig. 22, 1947; height 27 mm), from the Byram. mnarl. The acute, polygyrate protoconch of Pleurofusia ,viclcsburgeivsis was described by Casey, and is shown in Harris and Palmer's illustration, and also in Harris' (1937, 1)1. 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. vicksburgevsis and P. fusmnus fusinus are considered to be closely related. P. latira is indistinguish able from P. fusinus fusin us. The axial swellings of P. crzzaina (Olsson, 1932, p. 150, 1)1; 15, figs.* 6, 10; Mark~s, 1951, P). 131, pl. 8, fig. 6) are more subdued than those of P. fusinus fusinus.
The early Miocene P. glyjfta (Gardner) (1926-47, p. 293, pl. 38, fig. 34, 1938; Chipola formation, height 25.3 mm) is not closely related to any of the other species of (iruziturricula. Its last protoconch whorl merges into the first post-protoconch whorl. The first three post- lprotoconch whorls are unicarinate, anguilated 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. Trle lprotoconcli of the type is worn. TJhle preservation is better on two minute shells (USNMT 644754) in C. R. 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, 138di 138e, 138f. Middle part, eastern area, localities 139b, 139c, 142, 146, 147b, 147ifI 147g, 151, 155, 155b, 155c, 157, 159, 159d; western area, locality 161c. Upper part, eastern area, localities 173a, 175, 176, 177b. Middle Miocene deposits, Darien and Chiriqui areas, Panamd. Angostura formation (middle Miocene), Ecuador.
Genus leucosyrinx Dali
Dail, Harvard Coil. Mus. Comp. Zoology Bull., v. 18, p. 75,
1889.
Type (orthiotype) :Pleuro totnclla verrilffj (Dall) (Picurotoma
(Picurotomclla) cerrifiji) North Carolina andi ealsterii
Gulf of Mexico.
Leucosyrinx xenica Woodring, n. sp.
Plate 57, figure 15
Small for the genius, very slender, anal fasciole barely constricted, forming a steep ramp. Protoconch


b~lunlt 134-whorled, last whorl carinate, the carina, gradually retreating toward base, end marked by gradual applearanice of sculpture. First post-protoconch whorl sculptured with 10 swollen axial ribs, strong only on p)erip)hery, fadncg out, on anal fasciole, subdued toward base of wh-lorl. Nine ribs on antepenult whorl. Ribs lowver, wider, and gradually fading- out on penult, absent on body wNhorl, except a slight swelling at beginning of whorl. Spiral sculpture obscure on first postprotoconch wlhool, gradually strengthened on succeedinlg whiorls, consisting of closely spaced, narrow threads of unequal width, subdued on anal fasciole. Outer lip broken back. As shown by growth lines, anal sinus wide, adjoining sutture, upper limb short, lower limb long. Aperture long, narrow. Siplional canal not notched, according to g-rowth lines. Siphional f asciole not inflated.
1-eight 15.4 min, diameter 4.9 mm. (type).
Type USNM 645768.
Type locality: 185 (USGS 8383, Caribbean coast, west of Rio Miguel, station 26 plus 100 feet (30 in), Panamd), 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 constrcte anl fsciledisappearing axial sculpture, and fairly strong spiral sculpture is unique in the grenus. Only two other species are known in the Tertiary Caribbean province: Leucosyrinx cloi Olsson (1922, 1). 59, pl. 5, fig. 19; Lim6n formation, IBocas del Trrlo area, Panam6.) and L. nicoya Olsson (1942, p. 57, 1)l. 12, fig(. 2; Chiarco Azul formation, southwestern 1Panam6.), and none in southeastern United States.
Leucosyi-ivx is basically a deep-water genus. It has beeii dredged in western Atlantic waters at depths of 0() to 1,180 fathoms, for the most part between 300 and 800. It is improbable, however, that the fossils fromt the type locality of L. xenica represent a depth greater than 100 fathoms, if that much.
Occurrence: 17p)Jer part of Gatuii formation (middlle Miocene), western area, locality 185.

Genus Cochiespira Conrad
Conrad, Ain. Jour. Conehiology, v. 1, p. 19, 1865. Type (monotype) : Picurotorna cristata Conrad, Oligocene,
Mississippi.
During recent years (/ochle8pi?'a engonata Conrad generally has been accepted as the type species. That species and Pleiwotoina cistata were assigned to Coch/e81)na by Conrad. In 1.913, however, Vincent (1913, 1). 2,5) pointed out that Cochldepira engonata was a nudle name when (Ioch/espira was proposed, and was not validated until three (actually a little more than four) months later (Conrad, 1865, p. 142; Conrad,


370






GASTROPODS: EIJLIMIDAE, MAIIGINELLIDAE TO 1{ELMINTHOGLYPTIDAE37


1865a, p. 210, pl. 21, fig. 12). Pleurotoina cristata therefore is the mi-onotype. Cocidespira engonata is the type of Coclesphropsi8 (Casey, 1904, p. 143; logotype, designated by Cossmann, 1906, p. 221).

Cochiespira? species
A f ragment of 212 incomplete whorls, found in the moderately deep-water facies of the Caimito formation on Barro Colorado Island, probably represents Cochlespbra. Trle peripheral frill is slightly upturned. It is faintly noded, but if it had been serrate, the spines aire gone. A faint spiral thread lies in the concavity between the frill and the suture, closer to the frill than to the suture. The penuilt whorl has a height of about
1. 5 mm.
Occurrence: Caimnito formation, Gatun Lake area, (late Oligocene), locality 54g.

Subgenus Ancistrosyrinx Dali
Dali, Harvard Coil. Mus. Comp. Zoology Bull., vol. 9, p. 53,
1881.
Type (monotype): Ancistrosyrinx elcyans Dali, living, West
Indies.
The protoconch of Aqiistrosyrinx is about 11/2whorled and bluntly tipped, whereas the protoconch of Cocidespira is about 212-whorled and has an acute tip. The peripheral frill of Aqicistrosyinx is more strong,(ly upturned than that of Cocidespira, producing adeepr 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 A'ncistrosyrivx as a subgenus of Cocidespira is adopted.

Cochiespira (Ancistrosyrinx) cedonulli (Reeve) Plate 57, figures 9, 10
Picurotoma cedo-nulili Reeve, Zool. Soc. London Proc., p. 185,
1843 (Recent, Panamd6). Reeve, Conchologia iconica, v. 1, Picuro toma, species 117, figs. 117, 117a, 1843 (living,
Panam6.).
Ancistrosyrin x ccdonulli rccvi Olsson, Bull. Am. Paleontology,
v. 27, no. 106, p. 53, pl. 10, fig. 4, 1942 (Pliocene, Costa Rica). Oisson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst., p. 111, p1. 15,
fig. 5, 1964 (Mliocene or Pliocene, Ecuador).
Picurotoma miranda Guppy, Sci Assoc. Trinidad Proc., v. 2,
no. 4, pt. 12, p). 178, 1)1. 7, fig. 19, 1882 (Miocene, Jamaica); reprint, Bull. Am. Paieontoiogy, v. 8, no. 35, p. 99, p1. 5,
fig. 19, 1921.
Ancistrosyrinr mniranda (Guppy), Woodring, Carnegie Inst.
Washington Pub. 385, p. 165, pi. 6, figs. 1-3, 1928 (Miocene, Jamaica). Perriiiiat Montoya, 111xico Univ. Nac., Inst. Geologfa Paleontologia Mexicana, no. 8, p). 29, p1. 4,
figs. 11, 12, 1960 (Miocene, TA1xico).


Ancistrosyrix (lall! Oisson, Bull. Am. Paleontology, v. 9, no.
39, p. 60, pi. 4, fig. 16, 1922 (M.Niocene, Panam;!).
Of medium size, moderately inflated. Protoconch about 112-whlorled, 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 on intermediate and late whorls. A spiral thread, appearing near upper base of frill on third p)ost- -protoconech 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.
H-eight (incomplete) 19.3 mm, diameter (spines broken) 10.2 mmn (figured specimen).
Type: British Museum (Natural History).
Type locality: Panam-i Bay, living.
A miinute shell from the middle part of the Gatun form-ation and the figured specimen from the upper part in the eastern area aire referred to Cochiespira cedognuii.
Three living Amierican species of the subgenus An cistrosyi2'fl aire recognized: Coo/des pira elegans Dall (1881, p. 541; 1889, p. 718, p1. 38, fig. 3) and C. 'radiata Dall (1889, p. T8, 1)1. 12, fig. 12) in the Caribbean region, amnd C. cedomudli in the Panamic region. C. elegavs (height 32 mim, depth range 260 to 805 fathomis) is sculptured below the frill with strong, noded slpirals. Thle corresponding spiral threads of C. radiata (heighit 25 mmn, depth range 22 to 170 fathoms) are nionnioded and moderately strong. C. cedonulli (height 23 mmu depth range 10 to 153 fathoms) lacks spiral threads between the frill and the pillar, except that on some slpecimieis, both living and fossil, an obscure thread em-erges at the upper end of the aperture.
These three species appeared abruptly in the middle Miocene sea of the present Caribbean region. The Mliocene representatives of both living Caribbean species, however, are smaller than living shells. Whether time small fossils are to be considered ais a, small predecessor race or as inmature is uncertain -iuntil larger samples are available. The occurrence of only small slpecimenis of C. ce(100idli at some fossil localities sugg-ests that, the small specimens of all three species are immature. Four small specimens of C. elegans (mnaximnum height 17 mmi) have been found in Costa RicaOlsson's (1922,1P. 60, 1)1. 4, fig. 17) Avcistrosyinx elegaiis, variety-and in the Cercado and Gurabo formations of the IDoiminican Republic. The Gurabo formnation yielded also a small specimen (height 18 mm) of


371






372 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


C. radiata. A small shell (height 12 mm) in Spencer's collection of deep-water Mliocene 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 aI smooth space. Should this type of sculpture prove to be consistent, treatment as a subspecies of C. radiata would be warranted. l3L'se's (in B16se and Toula, 1910, p). 251, pl. 13, fig. 26) Pleurotoma (A'nci8tI'o,/rivx) aff. radiata, from the same locality, evidently is not a species of Cooblespira.
C. cedon alli. is fairly widespread inI middle Mliocene deposits: Jamnaica, the 1'eliuantepec area of M16xico, Costa Rica, and Panault. Small Jamaican specimens (maximum height 13 m-m) were named P/euro torna ,iraunda. Trley are slender, but not more slender than ain immature living shell of 0. cedonudli (USNM 96801). T1hle fossils from the Tehuantepec area are comparable in size to Jamaican specimens; those from Costa Rica and Panamat are of miediumi size (20 nm and about 21 mmi respectively). Thle largest fossils eightt 35 to 43 mim), larger than any available living shell, occur in the present Panamic region: in the Esmeraldas formation of Ecuador and the Chiarco Aziil formation of southeastern Costa Rica.
Through the p)atterin of former distribution and survival shown by C. oedona//i is unusual at the specific level, it is common at subgreneric and generic levels.
Occurrence : Middle and upper parts of Gatun formation middlee Miocene). Middle part, eastern area, locality 14'(1. Upper part, eastern area, locality 177b. Bowcden formation (middle Mliocene), Jamaica. Middle Miocene deposits, rreliuitepec. area, ,6xico, Costa Rica. Lim6mi formation (late Mliocene) llocas dlel Toro) area, Panami. Esmnera1las formation (late Mliocene or Pliocene), Ecuador. ('larco Aztil formation (Pliocene) Costa Rica. Living-, Mlazatlani, Mf6xico, to Panam~i Bay.

Subfamily BORSONIINAE Genus Scobinella. Conrad
Conrad, Acad. Nat. Sci. Phila. J.our., 2d1 ser., vol. 1, p. 120, 1848. Type (monotype) :Scobinelia cwclata Conrad, Oligocene, Miss.
In southern IUited States Seobinel/a, an extinct endemic American genuls, has an age range of0cn to Oligocene, but in the Tr1tiar1y~ Caribbean province it surviv-ed until the end of Miocene time. The Eocene and Oligocene species heretofore described have a height of l(0 to 35r mmn, whereas the Milocene species are larger : height 42 to 70 mm. rThe i\Iiocene species have been reviewed recently by Olsson (1964, p). 114117).


Scobinelia morierei (Cossmann)
Plate 58, figures 26, 27
EBuch ilodoni 1[oricrci Laville miss., Cossmnann, Jour. Conchlyliologic, v. 61, p. 34, 1)1. 3, figs. 6, 7, 1913 (Miocene, Canal
Zone).
SCobbnclla Mforicrci (Laville) Cossrnann, Olsson, Bull. Am. Paleontology, v. ), 110. 39, p). 79), 1)1. 4, figs. 3, 4, 1922 (Miocene,
1Panamd).
Scobinclla fliorjcrci (?) (Laville) Anderson, Calif. Acad. Sdi.
Proc., -1th ser.. v. 18, no. 4, p. 131, p1. 8, figs. 6, 7, 1929
(Mliocene, Colombia).
Scobilla moricei (d'ossmnann) Olsson, Neogene mollusks from
northiwestern Ecuador, 1). 115 (in discussion) pl. 19, figs.
4, 4a, 1964 (Miocenie, Panaind).
Scobincila ioriori !lailiw~lnsis Rutsch, Eclogie Geol. Heivetlie,
v. 23, 1). 611, p1. 17. fig-s. 8, 9, 1930 (M.Niocene, Venezuela).
Rutsch, Schweitzer. Paheorit. Gesell. Abh., v. 55, no. 1,
pl. 8, fig. 17, text fig. 13, 1934 (Miocene, Venezuela).
Not 1Seobiclla nioricrci (Laville) Cossniann, Maury, Bull. Am.
Paleontology v. 10, no. 42, p. 193, pl. 34, fig. 1, 1925
(Miocene, Trinidad).
Not SCobin(11(I noricr('i (Lavilie), Perrilliat Montoya, Mkxico
TI niv. Nac. Inst. (leologia Paleontologia 'Mexicana. no. 8,
1). 28, p1. 4, figs. 7, 8, 1960 ('Miocene, M&Xico).
Moderately large, slender, turreted, anal fasciole strongly constricted. Protoconch not preserved. Sculptune of earliest lpreservNed whorls consisting of a noded suitural b)and-, arcuiate axial threads on) anal fasciole; close]l spa(ced lprotractiv-e axial ribs, noded by two p~eiph)eral spiral cords and a narrower cord at base of whlorl. Sculpture of later whorls consisting of retract ive nodes on stitiiral band ; two closely spaced rows of small nodes on anal fasciole (three on-body whorl) arirangredl in axially arcutate pattern ; coarsely no0led shoulder cord, bilirate on last 212, whorls, upper part wider and more coarsely noded than lo-wer p~art; and up to body -whorl two or three noded spiral cords siimmilar to narrow low-er part of shoulder cord. Nodes arranged in protractive p~atterni. On pillar nodes 1)rogr1essively sill)pressed toward si phonal fasciole. Anal slinus and siplional notch moderately dleep), as shown by g-row\th lines. Interior of outer lip lirate. Columella b~earing- two wi-dely spacedl upper -folds and two closely a(edl, weak lower folds. Siphonal fasciole slightly iiitlnteol, learliig_ a few widely spaced, weak spiral threiads.
I feiglit almostt coinplete) 37- mmn estimatedd height 412 inun), diameter ( Incomplete) 12.5 min.
T e EOle oles ANIinesi Paris.
11,ype lcalty Alnd, Pnaa Canal, Canal Zone, iupp ier part, of Gatlin formation.
Thme descipiltioni is based on an almost complete sl)eciieii ol moderate size -from the lower part of the Gatini formulatiomi--tme onlyN Specimen in the collections now lbeing- studied. The type (heighndt 42 mm) was col-


372






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO I{ELMINTHOGLYPTIDAE37


]ected at an outcrop of the upper part of the Gatun in the Mmndi area near locality 172.
Scobiiiella mvorierei, occurs also in the late Miocene Lim-on formation at Lim6n, Costa, Rica, and on islands of the nearby Bocas del rToro Archiipelago, Panam6. The change from arcuate axial threads to small nodes on the anal fasciole takes place two or three whorls earlier on these late Miocene shells than on the Gatun specimen. Shells from Costa Rica (Olsson, 1964, pl. 19, fig. 4a) are decidedly less turreted than the Gatun form; those from the IBocas del Toro area (Olsson, 1922,11)1. 4, figs. 3, 4; 1964, pl. 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. morieiri gavilavensi8 indicate no features to substantiate that subspecies. If Anderson's Scobinella is S. movierei the age range of the species is early to late Miocene.
Though the early and middle Miocene Ecuadorean S. on;?ola Olsson (1964, p. 116, pl. 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. 'rnorerei.
The fiat-sided, biconic, Mexican Scobinella, recorded as S. 91noriereil 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 9norierei is to be attributed to Cossmann. According to his statement (1913, p. 36), there is no indication that hie 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. Lim6n formation (late Mliocene), Costa Rica, and Bocas del Toro area, Panamn. Punta Gavihin formation (late Miocene), Vedjezeula.

Scobinella aff. S. morierei (Cossmann)
Three defective specimnens establish the presence of Scobinella in the moderately deep-water facies of the Caimito formation on IBarro Colorado Island. So far as the mneager material groes, this Oligocene Scobinella is closely allied to S. morierei, but the shoulder nodes are more widely spaced. The ]argest of these incomplete specimens has a height of 30 mm. and an estimated total height of about 40 mm; that is, compar-


able to S. inoierei and larger than any other known Oligocene species.
Occurrence: Caimito formation (late Oligocene), IBarro Colorado Island, localities 54h, 54m.
Scobineila ecuadoriana Oisson
Plate 62, figures 33, 34
Scobincula eceuadoriana Olsson, Neogene mollusks from northwestern Ecuador, 1). 117, p1. 19, figs. 1, la, 1b, 1964 (Miocene, Ecuador).
The Chiagres sandstone yielded ain incomplete large Scobinela identified as S. ecuado~iana. It shows the most distinctive feature of that species: large nodes on the shoulder, immediately below the anal fasciole, even on the penult whorl, as well as on the body whorl. Thne interior of the outer lip is lirate and the columella bears four folds, arranged like those of S. morierei. The height of the body and penuilt whorls is 43.4 mim, indicating a total height of about 63 mm.
S. ecitadoriana is a large species, but not as large as the appropriately named middle Miocene S. magnifica (Gabb) (Olsson, 1964, p. 115, pl. 19, figs. 3, 3a), which occurs in the Dominican Republic and Jamaica (height 70o.5 mm).
Occurrence: Chagres sandstone (late Miocene), locality 208. Esmeraldas formation (late Miocene), Ecuador.
Genus Paraborsonia Pilsbry
Pilsbry, Acad. Nat. Sdi. Phila. Proc., v. 73, p. 326, 1922. Type (ortliotype) : Borsonia (Paraborsonia) varicosa (Sowerby) (Mitra varicosa Sowerby), Miocene, Dominican
Republic.
Parab oironia 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 all'. 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 lbody 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 p~illarin issing. Apex of anal sinus on wide noded band, as shown by growth lines. Interior of outer lip and columella inaccessible.


373







374 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Height (incomplete) 8.5 mmn (estimated height 12 mmn), diameter (incomplete) 5.5 mm.
Ani incomplete, crushed specimen from the moderately deep-water facies of the Caimoto formation onl Iarro Colorado Island is identified as the earliest known species of Paraborsonia. If it is mature, it is also the smallest species. Though the shell is too fragile to attemlpt to expose the columella, in other features it is a Paraborso??ia closely allied to the middle Miocene Trinidad species P. brasso?)isis (Mansfield, 1925, p. 30, pl. 5, fig. 8). The wide band of that species Is bilirate at an earlier stage. Like 2liicrodrillia trina (p. 389), P. brassomsis was based on float material, but is represented by some 40 outcrop specimens from USGS locality 21234 onl Mayo River: the same locality that yielded an adequate sample of 2licrodrillia trina. P. bra, soswnsis occurs also in the San Jos6 calcareous silt member of the Manzanilla formation (USGS 21231).
Heretofore Paralborsovia was known in the middle Miocene deposits of tbe Dominican Republic and Trinidad, and in late Miocene in the lBocas del Toro area, Panamd. Borson ia (Parab orsonia) eo taurana (Jung, 1965, p. 572, p1. 77, figs. 11-14; middle Miocene, Venezuela) is not a Paraborsonia.
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 B~artsch (1943, p. 82, pl. 7, fig. 5, p1. 10, fig. 7).

Subgenus Neodrillia Bartsch
Bartsch, Soc. Cubana Historia Nat. Felipe Poey, Mem., v. :17,
p. 83, 1943.
Type (orthiotype) :Nec drillia eydia Bartschi, living, North Carolina to Florida and West Indies.
The smaller size, more slender outline, lack of ain umbilicus, and finer spiral sculpture of Neodilia distinguish it from I) ril/a s.s. Leptadrillia (Woociring, 1928, p). 159; type (orthotype) : TuNsi (Surczla) parkein (iabb, Mliocenle, D~om inican Republic) is more slender than Neodq 7/ia and has narrow axial ribs and no spiiral sculpture, except onl the pillar.
According to Pilsbry and Hlarbison (1933, p. 112), Leptad i/Va "differs from ('ymatos ilin by its more slender shape and longer interior canal, but there seems to be no other structural dlifference." Leptadill/ia, however, lacks a constricted anal fasciole, deep stromboid notchi, deep) siphional notch, and sharp thread


limliting the siphional fasciole all shown by Cyrnato8y1ivx (IDall, 1889, 1) 95; type (orthotype) : P/eurotoma liunata I-I. C. Lea, Miocene, Virginia). Cymatos8/JJ/x is the telperate counterpart of Clathrodrillia (p. 381). Onl jiiorphiologic grounds it may be better to
asi ( (maosyiinx lunata ac/mvica Tucker and Wilson (19P13, 1). 13,pl. 4, Jigs. 6,7Ti; Olsson and Petit, 1964, 1) 537, 1)1. 82, fig-s. 5-5b; assigned specific rank), a late Milocene Florida turrid, to a subgenius under
( iat ,dri//ia that has no spiral sculpture. A middle Mtiocene ally of (fyvmatovy)pinx ac/mi;ca is found in the rrel1itelpec area, where it has been recorded as Cymafo.syi)bhx lunata (Perrilliat Montoya, 1963, p. 29, pl. 5, fig(s. 17, 18).

Drillia (Neodrillia) riogurabonis eurysoma. Gardner
Plate 58, figures, 17, 18; plate 64, figure 2
"Drilia" curysrnia Gardner, U.S. Geol. Survey Prof. Paper 142,
1). 316, 1)1. 40, fig-. 10, 1937 (1938) (M~iocene, Florida).
Of mleditm size, moderately inflated, pillar short, whiorls not constricted adjoining( suture to form an anal fasciole. Protoconch acute, 3- to 312-whorled, last 1 to 112 wvhorls bulging near base. End of protoconch marked by anl arcuiate riblet, followed by first axial rib. Axial ribs undulating suture except where they are alined and onl body whorl, where they are attenitated near suture; ali ned on first 3 to 41/2 whorls; practically straight on about first 3 whorls, slightly arcuiate onl remaining spire whorls, decidedly arcuate onl body whorl; 7 or 8 on all whorls. Last rib wide and igh-i, forming a humnp. Spiral sculpture consisting of weak, closely spaced, narrow threads, not discernible onl earliest few whorls. Outer lip broken back. Anal sinuis moderately deep, as indicated by growth lines; upper li mlb heavily bordered. Strom-boid notch very shallow, according- to growth lines. Siphional notch mfoderately deep. Si 1)1 onal fasciole slightly inflated, sculptured witli spiral threads, stronger and more widlel spaced th an those elsewhere.
hieigh~lt (practically complete) 11.8 mlm, diameter (incluiding humnp) 4.6 mmn (larger figured specimen).

TYpe locally 1 S1 86 5 to 6 illes west-northwvest of M~ossyhiead, Walton County, Florida, Shoal IBtiv~er fritol
Thie larger fig-ured specimen, Onl Which the earliest part of' the protoconcli is mlissing,. is the only practicall 11omp111lete mnature shtell amlongo the six specimens oftis (jatun tuirrid. 'I'lough it is identified as the Florida form, time axial ribs onl the early whorls of the two available specimens from the Shoal River format 'on ire muot, uniforinl alined. Gardner realized that her species is closely related to Neodi/ia iogurabonis


374







GASTROPODS: EULIMIDAE, MARGINELLIDAE TO HELMJNTHOGLYPTIDAE37


(Maury, 1917, p. 54, pl. 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, p1. 3, fig. 8) as Dr-illia afr. D). rioguraboni8 and by Rutsch (1942, p. 166) as Cymatosyr-ivx? 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 B~owden formation of Jamaica, described as "Drillia" sp. and Rellaspira? sp. (Woodring, 1928, p. 162, 163, respectively) represent Neod7rillia. Though the so-called Bellaspira? is poorly preserved, they probably are the same species-a species that is larger and more inflated than D. riogztrabonis.
The species so far mentioned, and also early Miocene species from the Tampa limestone of Florida and the Thomonde formation of Haiti (USGS 9945), are more similar to the living Panamic, species D. cybele (Pilsbry and Lowe (1932, p. 46, pl. 2, fig. 6) than to the living Caribbean species D. cydia. The sutural area of the Caribbean species is constricted, forming an anal fasciole, and the shell is sculptured with fine, submicroscopically crimped spiral threads. rThe sutural area of D. 8ubperpolita B6se) (in Bose and Toula, 1910, p. 246, pl. 13, fig. 22), a late Miocene species from the Tehuantepec area, is constricted, but not as strongly constricted as that of D. cydlia.
Abbott (1958, p. 96) was justified in placing in synonymy under D. cydia the four other living Caribbean species of Neod3 illia named by Bartsch (1943, p. 84-89) and in rejecting, on the basis of Melvill's illustration, B~artsch's identification of the Cuban species D. eupharnes Melvill (1923, p. 164, pl. 4, fig. 4). If Bartsch's identification were correct, D. oydia, the type species of Neodrilla, would be a synonym of D. euphane8.
Occurrence: Lower and middle parts of Gatun forination (middle Miocene). Lower part, locality 138c. Middle p~art, eastern area, locality 139b; western area, locality 162. Shoal River formation (middle Miocene), Florida.
Drillia? (Neodrillia?) species
The upper part of the iBohio formation of Barro Colorado Island yielded an incomplete minute clavine turrid consisting of the protoconch and four post1)rotoconch 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 allied. The anal fasciole is not constricted and spiral. sculpture is not discernible. The height, ininus time p~illar, is 4 nmn, and the diameter, minus the outer lip, 2mm.
Despite the absence of spiral sculpture, the outline, protoconch, and style of ribbing point to Neodi Wlla. 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 Dali
Dali, U.S. Natl. MNus. Proc., v. 56, p. 17, 1919. Type (orthotype) : Clathrodrillia halis Dali, living, Gulf of
California to Manzanillo, A16xico.

Subgenus Carinodrillia s.s.
Carinodrillia, like Pleurofustia, has Fusinus -like sculpture. It appeared in early Miocene time and survives in the western Atlantic and eastern Pacific oceans.
The subgenus Caiivodrillia s.s. embraces species of meduni to moderately large size. The type and other species, including C. zooki, have an acute, polygyrate lprotoconch (three to four whorls), the last 1/4 to / of which bears arcuate axial riblets. The protoconch of other fossilI and Recent species, however, is blunt, paucigyrate (about two whorls), and near the end bears a few peripheral axial swellings. The siphional canal of all the species described iii the present report is slightly notched and the siphional fasciole 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) : Ca? modi' Wa (Buchemta) taivoa Corea, living, Puerto Rlico).

Carinodrillia (Carinodrillia) zooki (Brown and Pilsbry)
Plate 58, figures 9-12; plate 65, figure 9
Dritlia zooki Brown and Pilsbry, Acad. Nat. Sd. Phila. Proc.,
v. 63, p). 345, pl. 23, fig. 18, 1911 (Miocene, Canal Zone). Drillia (Crassispira) zooki Brown and Pilsbry, Cossmann, Jour.
Conchyliologie, v. 61, p. 23, pl. 3, figs. 8, 9, 1913 (Miocene,
Canal Zone).
Not Drillia zoo1ki Brown and Pilsbry, Li, Geol. Soc. China Bull.,
8% 9, p). 275, 1)1. 8, fig. 71, 1930 (Miocene, Panam6. Bay)
= Clathrodrillia alccstis Dali, fide Pilsbry, Acad. Nat.
Sei. Phila. Proc., v. 83,1p. 434, 1931 (Living, 1Panaini Bay).


368-278 0 70 6


375






376 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Of medium size, slender, F'usinus-like, aside from short pillar. Protoconch acute, 3- to 4-whorled, last 1/4 to 1/,2 whiorl bearing- 3 to 7i arcuiate riblets terminating~ on or below the swollen, almost marinate periphery. End of 1)rotoconch marked by abrupt appearance of a -xial and spiral sculpltulre. First post- protoconch whorl scullptured with 6 to 8 closely spaced, wide, swollen axial ribs and 3 spiral threads : 1 sutural and 2 periphera]. Axial ribs on late whorls moderately swollen and fading out on lower part of anal fasciole, or strongly swollen and modifying(1 outline of lower part of anal fasciole; 8 to 10 ribs on p~enullt whorl. Spiral sculpture of late spire whorls consisting of a sutural thread and generally ") other widely spaced primary threads overriding axial ribs; a fourth thread exposed at base of whiorl on some specimens. Sutural thread strong on all whorls or relatively weak. In addition to sit ural thread, 15 to 1T primary threads on body whorl, those on pillar more closely spaced and weaker than elsewhere. Microscopic secondary spiral threads and exaggrerated 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 limib somewhat thickened.
Estimated restored heighit 25 mm, dliameter (including hump) 8 mmn (largest specimen). H-eight 19.8 mmi, diameter (Including hump) 6 mim (figured strongly sculptured specimenn.
Type: Acad. Nat. Sci. Phila. 1695.
Type locality: Gatun Locks excavation, Canal Zone, middle part of Gatun formation.
Trlhe 20 specimens from the three parts of the Gatun formation in the eastern area identified as Cai inodrillia zoole show a wide range of variation, chiefly affecting the strengthi of the axial ribs and sutural spiral thread. The~~ type is an incomplete specimen that has subdued sculpture, like thiat shown on plate 58, figures 9, 10, whereas plate 58, figures 11, 12, shows strong sculpture. Other shells, however, are more or less intermediate. Trhe supplression of axial ribs on the lower part of the anal fasciole of specimens thiat have subdued sculpture produces the effect, of a wide fasciole.
This slender species is more similar to C. haiostrcplli8 (1)all) (1889, p). 86, pl. 13, fig. 3), dredged northwest of Alacran Reef off Yucatdin at a depth of 84 fathomns, than to any of the described fossil species. TIhe primary, spiral threads of the living( species are more delicate. Th'le lprotocoic of the type (the only available specimen) is blunt and )aucig-yrate : 13/t whorls. It shjowNs a suggr~estion of a few axial riblets, but is worn. Thie primary sp~iral threads of C. fcli8 Olsson (19634, p. 102, pl. 19, fig. 6) a inidldle 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 Gratun formation (middle Miocene). Lower part, localitY 3a Midldle part, eastern area, localities 139b, 139e, 146, 14Thl 147f, 147g, 1,55c, 1571, 159d. Upper part, eastern ar1ea, localities 173, 1 75.
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 preservedl whiorls partly corroded. Axial ribs wide, swollen; 6 on p~enult and body whorls. A sutural primary sp)iral thread and 4 others on penult whorl ; 19 others on body whiorl, one of which is replaced by 2 weaker threads. A strong- secondary spiral thread at lower edge of anal fasciole. Subicroscopic secondary spiral threads on noncorroded whorls; those on late half of body wI ion st rengtlIicued. E xaggeratecd growth threads here and there especilly on late half of body whorl. Last axial rib widened to formn a hump. Outer lip broken back. Upper limib of anal sinus heavily bordered. Aperture moderately long.
Ifeight almostt complete) 26.8 mim, diameter (inchiding hiump) 9.8 mmit (figured specimen).
One specimen of this species is In a collection from the middle part of the Gatun formation. It is more slender than Caiiodi!1lia elocata (Pilsbry, 1922, p.
31,?1.1,1i.9 Gurabo formation, Dominican Republic) and hias stronger secondary spiral sculpture.
It has been claied that C'. wIon chesterce Pilsbry (1922, p). 318, pl. 16, figs. 7i, 8), also from the Dominican Rlepublic, is an usually slender, long- canaled formi oI' c ocaa (Woodring, 1928, p). 155), but it dloubltless is a distinct species, as Pilsbry thought.
()ccurrence : M[iddle part of Gatun formation (mid(Ilie Milocenie), eastern area, locality 139c.

Carinodrillia (Carinodrillia) species Plate 63, figure 5
M odh'rate Iy large, moderately slender, pillar moderately long,. anial fasciole strongly constricted. Protoc(1(] a ml early post-protoconchi whorls missing. Axial 111)8 iu()deniltel 'vN widle, swNollen ; 8 on lpenullt and body wh1orls. Sutural primary spiral thread closely hugging sutture. Iiour other pliwimrv spiral threads on penult wh'lorl; 11 on body whiorl. Submicroscopic secondary spiral thlreadls on noncorroded parts ; those on anal huasciole Thiuter thian elsewhere. A strong secondary tllread(i t lower edge of anal fasciole. Last axial rib


376






GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE37


higher and wider than others, forming a narrow hump. Outer lip broken back. Upper limb of anal sinus heavily bordered. Aperture moderately long.
H-eight (almost complete) 27.6 mm, diameter (inclucding hump) 10.3 mm.
Ani unnamed clavine turrid from the Chagres sandStonle is represented by anl imperfect specimen. The entire apertural face of the shell and part of the adalpertural face are corroded. The relatively narrow axial ribs and strongly constricted, weakly sculptured anal fasciole 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. Cohln., v. 91, no. 2, p. 22, 1934. Type (ortliotype) : Darbya lira Bartsch, living off north coast
of Puerto Rico.

Subgenus Buridrillia Olsson?
Gisson, Bull. Am. Paleontology, v. 27, no. 106, p. 51, 1942. Type (orthiotype): C~lathrodrillia pan arica Olsson, Pliocene,
Pacific coast of Central America straddling Costa RicaPanamd. boundary.
Though the protoconcli of B'uridrillia is unknown and Dauibya 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 p~laits of the borsonines and does not belie the clavine features.
A worn corroded shell, dredged ait a depth of 134 fathoms near Cocos Island (USNM 106888), is indistinguishable from the type species of Bunridrillia, so far as it goes. Cocos Island is off Coiba, Island, in the Pacific coastal waters of Panamk.

Darbya? (fluridrillia?) species
Plate 63, figures 6, 7
Small, moderately slender, whorls strongly inflated, pillar of moderate length. Protoconch and early postprotoconch whorls missing. Anal fasciole constricted. Axial ribs wide and widely spaced, attenuated and bent onl anal fasciole, 10 on penult whorl. Spiral sculpture consisting of narrow bands, separated by grooves, in which microscopic axial threads are conspicuous. Spiral bands closely spaced onl anal fasciole. A hump on body whorl near aperture. Outer lip broken back


to humip. Aperture and siphonal canal of moderate length. As shown by growth lines, anal sinus moderately deep; border of upper limb thin. Siphonal fasciole sculptured with closely spaced spiral threads.
H-eight (incomplete) 15 mim, diameter 6 mmn.
A second unnamed clavine species from the Chagres sandstone, like the preceding species, is represented by anl 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 siphional canal and other apertural features, so far as they are preserved, suggest a small species of B'aridrillia 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 onl the Pacific coast of Central America straddling the Costa Rica-Panam~i boundary (Olsson, 1942, p. 51-53). rTle spiral sculpture of the Chiagres species is stronger than that of the described species.
Occurrence: Chiagres sandstone (late Miocene), locality 208.
Genus Crassispira, Swainsan
Swainson, A treatise on mnalacology, p. 313, 1840. Type (logotype, Ilerrmiannseni, Indicis generum malacozoorum,
v. 1, p. 318, 1847): Plcurotoma bottw Valenciennes in Kiener (cited by Swvainson as Plcurotoma Bottwv Auct.),
living, Gulf of California.

Subgenus Crassispira, s.s.
(Irasispira henweeni has a narrower aperture and less inflated siphonial fasciole than the type species, as well as fewer and wider axial ribs. In number and width of ribs (C. agis (Woodring, 1928, p. 151, pl. 4, lig. 12; B~owden formation, Jamaica) is more similar to the type species, but, like C. henekeni, has a narrower alperture.

Crassispira (Crassispira) henekeni leptalea Woodring, n. subsp.
Plate 58, figure 19; plate 65, figure 5
Of miediumi size, slender, aperture long and narrow. Protoconch acute, conical, 3-whorled; end marked by fain-t arcuate axial riblet. Sculpture consisting of slightly p)rotractive, swollen axial ribs (8 on body whorl), overridden by strong, narrow primary spiral threads (7 or 8 oil penullt whorl). Subsutural thread strong; anal fasciole below subsutural thread bearing faint secondary spiral threads. Outer lip broken back. Anal sinus wide, apex below subsutural thread. Silplonal fasciole scarcely inflated.
H-eight (practically complete) 30.6 mm, diameter 9.7 inun (type). Height (almost complete) 34 mim, diameter 10.5 mil.


377







378 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Type: USNM 645777.
Type locality: 147h (USGS 6442, '/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, bo0th 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 Mliocene Caribbean Sea. The nominate subspecies (Sowerby, 1850, p. 50, pl. 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. Trle blunt, cylindrical protoconcli is not thought to belie the affinities of the Panam6. form.
Crass ispira i wqitistrata (Li) (Pilsbry, 1931, p. 437, p1. 41, fig. 2), living in 1Panamni. Bay, is the most similar living( species. No comparable species is known in the western Atlantic Ocean. The deep-water western Atlantic species I1'leotoina (Drilia) horrenda Watson (1886, p. 308, pl. 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 Milocene), localities 147h, 157.

Subgenus Hindsiclava Hertlein and Strong

Hertlein and Strong, Amn. Mus. Nat. H-istory Bull., v. 107, p.
227, 1955.
Type (orthotype) : Picurotorna militaris Hinds in Reeve, living,
Baja California to Colombia.
Trle more slender outline, more numerous and narrower axial ribs, stronger spiral sculpture, narrower aperture, and less inflated siphonal fasciole distinguish Hindsiclava from Ciassis'pira s.s.
The 01ligocene Species Plemirotoina ab andcans Conrad (1848,1p. 115, 1)1. 11, fig. 25; Byramn marl, Mississippi) is the earliest Ilidsiela'ra now known. Conrad's illustration, however, leaves much to be desired.

Crassispira (Hindsiclava) consors consors (Sowerby)

Plate 58, figures 1, 22; plate (35, figure (3

Picurotorna consors Sowerby, Geol. Soc. London Quart. Jour.,
v. 63, p. 50, 185(0 (M.Niocene, D~ominican Republic). Guppy, Idem, v. 32, p. 527, 1p1. 28, fig. 7, 18763 (Miocene, Dominican Republic).
Drillia, consotrs (Sowverby) Brown and Pilsbry, Acad. Nat. Sci.
Phila. 1'roc., v. (63, p. 345, 1911 (Miocene, Canial Zone).
Maury, Bull. Ain Paleontology, v. 5, no. 29, p. 53, pl. 8, figs. 15, 163, 1917 (MNiocene, Dominican Republic). Maury, N.Y. Acad. Sci., Scientific Survey of P~orto Rico and Virgin Islands, v. 3, lit. 1, 1). 71, 1920 (-Miocene, Puerto Rico).
Pilsbry, Acad. Nat. Sci. Phila. Proc., v. 73, p). 319, pl1. 163,


fig. 3, 1922 (Miocene, Dominican Republic). Olsson, Bull.
Am. Paleontology, v. 9, no. 39, p. (31, pl. 4, figs. 8, 10, 13, 1922 (Miocene, Costa Rica, Canal Zone, Panaid) Paliner, Idem, v. 10, no. 40, p. 11, pl. 2, figs. 7, 8, 1923 (Miocene, Costa Rica, Dominican Republic). Maury, Idem, v. 10, no. 42, p). 190, pl. 32, fig. 10, 1925 (Miocene, Trinidad). MNaury, Brosil Servi:o Gleol. -Mineral. Mon. 4, p).
203, 1)1. 12, fig. 5, 1925 (M.Niocene, Dominican Republic). IDrillia consort, ((huppy) Cossinann, Jour. Conchyliologie, v. 631,
p. 20, pl. 2, figs. 8-14, 1913 (Miocene, Martinique, Domninican Republic, Canal Zone).
Tn rris (Crassisp ira) con sors (Sowerby), Rutscb, Schweizer
Paeomit. Gesell. Abb., v. 55, p. 99, p1. 8, figs. 13-163, 1934
(Miocene, Venezuela).
Crassispira consors (Sowerby), Pflug, Acta Humboldtiana, Geol.
Paheontologica ser., no. 1, p. 67, 1)1. 19, figs. 4, 7, 10, 1961
(_Miocene, Dominican Republic).
Crassispira aff. consors (Sowerby), Jung, Bull. Am. Paleontology, v. 49, no. 223, p. 5635, pl. 763, figs. 14, 15, 19635 (Miocene, Venezuela).
?Crassispira? cf. Crass isp ira?" consors (Guppy), Marks, Idem,
v. 33, 110. 139, p). 135, 1951 (Miocene, Ecuador).
?Drillia consors portoricwnsis Hubbard, N.Y. Acad. Sdi., Scientific Survey of P'orto Rico and Virgin Islands, v. 3, pt. 2, 1). 158, pl. 24, figs. 8, 9, 1920 (Mliocene, Puerto Rico; not IDrillia, portorioxensis Hubbard, Idem, p). 159, pl. 24, figs.
11, 12).
?Drillia, con sors bulibrooki Mansfield, U.S. Natl. 'Mus. Proc., v.
66(,1p. 163,1)1. 3, fig. 10, 1925 (Miocene, Trinidad).
.,Drillia consors trinitatcnsis -Mansfield, Idem, p. 17, p1. 3, figs.
12, 13, 1925 (M-Niocene, Trinidad).
Turris (iDrillia) militaris (Hinds), Gabb, Am. Philos. Soc.
Trans., nm. ser., v. 15, p). 207, 1873 (Miocene, Dominican Republic). Gabb, Acad. Nat. Sd. Phila. Jour., 2d ser.,
v. 8, p. 350, 1881 (Pliocene, Costa Rica).
Picurotonia sp. aff. 1P. alcsidota macilcuta Dali, Toula, K. k.
Geol. Reiclisanstalt Jamrb., v. (31, p. 50(3, p)1. 30, fig. 11,
1911 (Miocene, Canal Zone).
Plcwrotoina (Drillia ) J)Ulli T0oula, Idenii, p). 5063, p1. 30, fig. 12,
1911 (Miocene, Canal Zone).
i'ot Piurotonia (1 lii Verrilt and 'Smnimtli, Connecticut Acad.
Trans., v. 5, p. 451, 1)1. 57, figs. 1, la, 1882.
IDrillia, macilcnta rcetaxis Pilsbry, Acad. Nat. Sdi. Phila. Proc.,
v. 73, p. 319, pl. 1(3, figs. 1, 2, 1922 (Miocene, Dominican
Republic).
Aslurcitla liob5ofli H-anna, California Acad. Sdi. Proc., 4th ser.,
N% 13, no. 10, p. 181, 1924 (substitute name for Picurotonma, (uli Toula).

Moderately large, slender, aperture long and narrow. Protocoll] acu1te (atpex almost invariably brok en), 21/.2- to el-wliorled, last half whorl slightly or
(lecide("I b I enid marked by appearance of first axial rib1. First 3 to .5 axial ribs narrow, arcuate, extendilig practically from suiture to suture. Later ribs wider, slightly p~rotractive, cut off at anal fasciole. Ribs prwressiNvelv narrower and more closely spaced. Scul1ptuire of late who0rls consisting of skewed rectangrles forimmed bv strong(, slightly p)rotractive axial ribs (22 to 30 on Mature penult whorl) and narrower primnary spliral threads (4 or 5 on nature penult whorl),


378







GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO I{ELMINTHOGLYPTJDAE37


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.
H-eight (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 large shell frorm the upper part of the Gatun illustrated by Cossmaiin (1913, pl. 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 all'. P. alesidota maci7en ita is split and the lower part itself is doubled. The type of his Pleuro tonsa dalli is an immature shell (height 26.6 mm).
Crasspira 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 iiamed Drillia macilenta rectaxis by Pilsbry. C. con sors consors reaches a height of about 75 mmn in the middle Miocene deposits of southeastern Costa Rica. Trhe axial ribs on the last two whorls of that large specimen and another almost as large, illustrated by Gisson (1922, pl. 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 mnicroscopic growth threads. With few known exceptions, shells from the type region and many other localities have microscopic growth threads, 1)lt lack microscopic spiral threads. A late Miocene specimen, collected on a cay oil' Bluefields Point at the west end of Valiente Peninsula (in the Bocas del


Toro area of northwestern Panamd), 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 Blueflelds Point. The last two or three whorls of C. comsors bulibrooki and C. con sors trinitatensi82, both from the middle Miocene part of the Brasso formation of Trinidad, bear one to three microscopic spiral threads, like those of the Panamd fossils, or a little stronger. Both subspecies probably are to be suppressed. All the specimens are small, doubtless immature, and C. consors trinitatensis' 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 h obso0ni, which was used by Gardner (19264T, 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 1-aiti, the IDominican Republic, and Venezuela are like small replicas of middle Miocene specimens (height 15 to 25 mm). The Brazilian C. cons o9s pen'na3 (Maury), (1925a, p. 205, 1)1. 12, figs. 1, 2, 4, 6), however, is large (height 40 mm) and has more ribs than C. consors
covw~. heslight~ly arcuate risshown inMaury's illustrations presumably are due to inaccurate drawing. iPlettrotoina (Th'ilia) alesidota lnaggia Thise (1906, p. 4T, 1)1. 5, figs. 30, 31, 33, 45) may be treated as a middle Miocene Mexican subspecies of D. condors that has spiral bands instead of narrower threads.
Tme status of the lineage in the Miocene of Florida has not yet been carefully evaluated: Drillia eupora Dall (1915,1p. 42, pl1. 5, fig. 3), Crassispira prceco'nsors Gardner (19201-47, 1). 299, pl. 39, fig. 3, 1938), C. caligonoide8 Gardner (Idem, p. 300, pl. 39, figs. 4, 9), C. blouvtensis Mansfield (1935, p. 22, pl. 1, fig. 11), and C. anitealesidota Mansfield (1930, p. 39, pl. 2, fig. 10). At least some of the Florida fossils have a shorter, wider aperture and more inflated siphonal fasciole than C. eo) so's. Trhis tendency culminates in the unillustrated Pliocene species C. perspirata (Dali) (18901903, p. 31, 1890).
C. con sw's, left descendants on both sides of Central America: C. alesidota (Dall) (1889, p. 84) and C. macilenta (Dail) (Idem, p. 85, pl. 36, fig. 1) in western Atlantic waters, and C. mnilita? 4s (H-inds in Reeve) (1843, species 55, pl1. 7, fig. 55), the type of Hindsiclava, in eastern Pacific waters. C. alesidota ranges from Cape H-atteras to southern Florida at depths of 35 to 110 fathoms. It has a somewhat less constricted anal


379






380 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


fasciole than C. COflsor; 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. ,macilen ta, dredged in the northeastern Gulf of Mexico (depth I11 fathoms) is the only specimen of that species in the collections of the U.S. National Museum. It is more slender than C. consort and C. alesidota, and has a less constricted anal fasciole than either of those species. Trhe secondary threads on the anal fasciole and the swelling-s on the axial ribs are strong. The lprotoconch is like that of C. alesido [a. A specimen of 0. a/esidota, dredged off Cape H-atteras, was identified by Dali. as C. macilenta. It remains to be seen whether Hlivd8ic/ava actually has withdrawn from the Caribbean Sea since Pliocene time. C. militari8 has strong swellings on the axial ribs and fairly strong undulations onl the subsutural spiral thread. It occurs in the Esmeraldas formation, of late Miocene age, of Ecuador (Olsson, 1964, 1) 97, pl. 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, 139el 146, 147b, 147f, 147g, 1471i, 151, 155, 155a, 155b, 155c, 156, 157, 159d; western area, locality 161a. Upper part, eastern area, localities 172, 173, 175, 176, 176a, 177b; western area, locality 182a. Thomonde formation (early Miocene), Haiti. Bl~atoa formation (early Miocene), Dominican Republic. Ayinam6n. limestone (early Miocene), Puerto Rico. La, Rosa, formation (early Miocene), Venezuela. Early Miocene part of Uscari shale, Costa. Rica. Gurabo formnation (middle Miocene), Dominican Republic. Middle Miocenie deposits, Martinique, Trrinidad, Venezuela, Costa, Rica, Colombia, 1)ari6n area, 1Panain-u (USGS 8477). Punta, Gavila'n formation (late M1iocene) Venezuela. Lin16ii formation (late Miocene), Costa, Rica, and lBocas del Toro ar-ea, Panamai. Deposits of late Miocene age, Tehuantepec (ire, M6xico. Pliocene deposits, Cumank area, Venezuela, Costa. Rica.

Crassispira. (Hindsiclava) pyrgoma Woodring, n. sp.
Plate 62, figures 21, 27
Of medium size, slender, distinctly turreted, apertire longr and narrow. Protoconch and early postjprotoconicli whorls missing. Sculpture consisting of slightly p~rotractive axial ribs (15 or 16 on penult whorl) and low primary spiral bands (4 or~ 5 onl penlilt whorl), swollen onl crest of ribs. Secondary threads of varying width between primary bands, some almost as wide as the bands, others microscopic. Anal fasciole wide, strongly constricted. Subsutural thread strong,


undulated secondary spiral threads between it and base of fasciole. Anal sinus wide, shallow, apex on lower pa~rt of fasciole.
H-eiglit (incomplete) 35.5 mm, diameter 10.5 mm (typ~e).'Type: TTSNM\ 645867. Paratype, USNM 645866.
Type locality: 208 (USGS 8437, Caribbean coast at mouth of Rio Indio, Panami), Chiagres sandstone.
Thongh) the type and p)aratype-the only specimens
-are incomplete and partly worn and corroded, this species is named, as the turreted outline, and wide and strongly constricted anal fasciole are distinctive. In general the sclplture resembles that of C. consors mnagna, mentioned under that species.
C. pyigoma doubtless is an offshoot from the lineage of (.co)?sor8. A middle Miocene Mexican turrid, from the Sayula district, Chiapas (USGS 5886), has aimian ledr turreted outline, a few less ribs (13 on p~enult), and somewhat weaker primary spiral sculpture.
Occurrence: Chiagres sandstone (late Miocene), locality 208.

Subgenus Crassispirella Bartsch and Rehder
Bartsch and Rehider, U.S. Natl. Mus. Proc., v. 87, p. 135, 1939. Type (orthotype) : (Jra.sispira (Crassispircelu) rugitccta (Dali)
[Turrns (Crassispira) rugitecta Dali], living, Baja California.
(!rassispirella is adopted as the name for species of (!ra. .ispira 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 thian other threads on the anal fasciole.
Thle earliest species aire found in formations of early MNiocene age: the Tampa formation of Florida and the 'Ilioionde formation of H-aiti. The subgenus now lives in bo0th 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
v~ie La, Boca, formation yielded a, small incomplete (r.a. ispprell and a m-inute shell that evidently is a, tip) of the same species. The axial ribs are closely spaced (about 15 onl last whorl). The subsutural thread is strong, undulated, and occuplies most of the anal fasciole. The spiral sculpture below the anal fasciole is Weak. Thle height, almost complete, is 8.8 mm and the diamleter 3.6 mi-m.
Occurrence : La Boca formation (early Miocene), locality 116a.


380






GASTROPODS: EULIMIDAE, MARGINELLIDAE TO IIELMINTHOGLYPTIDAE


Crassispira (Crassispirella) cymiation Woadring, n. sp.
Plate 58, figures 4, 5
Small, rapidly enlarging. Protoconch acute, 3- to 312-whorled, a wide axial swelling preceding end, marked by an arciiate riblet. Sculpture consisting of slightly protractive, inverted V-shaped axial ribs, swollen at apex (10 on body whorl) and low, narrow p)rimiary spiral threads (8 or 9 on penult whorl), roughiened by microscopic growth threads. Spiral threads subdued or disappearing on swollen apex of ribs; those on lower part of body whlorl above siphonal fasciole widely spaced. Anal fasciole undulated; subsutural thread strong; fasciole below siibsttral 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. Siphonatl fasciole slightly inflated.
Height (not quite complete) 8 mm, diameter (incomplete) 3.3 min (type). Height (incomplete) 7.5 mnm, diameter 4 mmn (paratype).
Type: USNM 645781. Paratype, USNM 645782.
Type locality: 161c (USGS 8382, railroad cuts west of Gatun Dam, station 13, Canal Zone), middle part of Gatun formation.
The undulated anal fasciole, 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 Gatuin formation in the western area. In sculptural plan it is not comparable to other small species from the Caribbean region: Crassispira inao'nisriparum (Maury) (1917, p. 56, pl. 9, fig. 5; Cercado formation, Dominican Republic), C. his paniolm (Maury) (1917, p. 56, pl. 9, fig. 6; Gurabo formation, Domninican Republic), C. ritanida (Mansfield) (1925, p. 24, pl. 4, fig. 10; Matura, formation, Trinidad), and C. a'nnella Woodring (192-8, p. 151, pl. 4, fig. 11; B~owden formation, Jamaica).
Oceurreie Middle part of Gatun formation (middie Miocene), western area, localities 161c, 170a.

Crassispira (Crassispirella) tyloessa Woodring, ni. sp.
Plate 58, figures 2, 3
Of medium size, subsutural spiral thread noded. Protoconch acute, about 212-whorled (apex worn), last whorl bulging below middle, apparently merging into first lpost- protoconch whorl. Axial ribs narrow, 20 on p~eniult whorl. Subsutural spiral thread bearing nodes alined with axial ribs. Spiral threads below anal f asciole narrow, relatively widely spaced, slightly noding crest of ribs; 5 on p~enult whorl. Anal fasciole below subsuitural 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. Growthi lines showing a deep anal sinus and shallow stromiboid notch. Upper border of anal sinus thickened. Siphional fasciole slightly inflated, sculptured within spiral threads cruder than those elsewhere.
Height 13.4 mmii, diameter (including hump) 5.2 mm (type). Heighit 15.3 mim, diameter 5.8 mmn (paratype).
Type: USNMN 645783; p aratype USNM 645784.
Typ)e locality: 139c (UJSGS 22018, east side of road leading from Transisthimian Highway to refinery site on Payardi Island, Panama.; about 100 meters southwvest of refinery gate), middle part of Gatun formation.
The com-bination of noded subsutural spiral thread and relatively widely spaced other spiral threads, not observed on other species of Crassispirella, is distinctive. The 1)aiatype 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 Cra88ispi'rella, similar to Cra8ispira janimaiceiisis (Guppy) (Woodring, 1928, p). 149, pl. 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 Clatlirodrillia Dali
Dali, U.S. Natl. Mus. Proc., v. 54, p. 317, 323, 1918. Type (orthiotype, 1p. 323): Plcurotoma gibbosa Reeve (Pleurotoma gibbosa Kiener = Murex gibbosa Born), living,
south border of Caribbean Sea.
A fragment of two incomplete whorls from the La Boca form-ation is listed as Clathrodri0ia sp.

Clatlirodrillia gatunensis (Toula)
Plate 58, figures 8, 20, 21, 23-25; plate 65, figure 7
Pleurotoina (Drillia) gatunensis Tonla, K. k. Geol. Reiclisanstalt Jahirb., v. 58, p. 707, p1. 25, fig. 16, 1909 (Miocene, Canal Zone). Engerrand and Urbina, Soc. Geol. Mexicana Bol., v. 6, pl. 59, fig. 27, 1910 (reproduction of Toula's
illustration).
Plcurotomta (Drillia) gatunensis Tonla, n. var., Toula, K. k.
Geol. Reiclisanistalt Jahrb., v. 61, p. 507, p1. 31, fig. 20,
1911 (Miocene, Canal Zone).
Drillia qatuncnsis (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 venvsta (Sowerby), Gisson, Bull. Am. Paleontology,
v. 9, no. 39, p). 61, pl. 4, fig. 9, 1922 (Miocene, Costa Rica). Drillia puertocolonibiana Weisbord, Idem, v. 14, no. 54, p. 55,
p1. 5, fig. 16, 1929 (Miocene, Colombia).


381






382 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Cia throdrillja pitertocolombiana (11eisbord), Jung, Idem, v. 49,
110. 223, 1). .566, pl. 76, figs. 12, 13, 19, 1965 (Miocene,
Venezuela).
Cia throdrillia onzola Olsson, Neogene mollusks from northwestern Ecuador, Paleontological Research Inst., p. 96, pl. 15,
figs. 6, 6a, 1964.
Moderately large, moderately slender, pillar strongly constricted, ap~erture relatively short and wide. Protoc'onchlillnt, almost cylindrical, 134 to 4-whorled, generally 134 to 2; lower part of last half whlorl faintly to distinctly aiigulated. Lower part of first post-protoconch whorl or two angulateci; first whorl sculptured with 7 or 8 axial ribs, swollen on angulation, graduially pinclied off near suiture. Faint spiral sculpture ap)pearig" Onl second or th ird post- protoconch whorl; anal fasciole onl about third. At that and later stages axial ribs5 slightly priotractive, moderately narrow, separated lby spaces grreater than their wvidth, disappearing onl body whorl below periphery ; 24 to 26 onl lpenllt whlorl. Spiral sculpture below anal fasciole consisting of narrow bands (4 to 7 onl penult whorl), which widen onl late whorls of a few specimens and ,ire separated by narrow grooves. Secondary spiral threads between lbandls onl a few specimens. Anal fasciole strongly constricted. Subsutural cord of variable strenioth aind width. Wide subsuitural cord strongly wrinkled by swellingrs corresponding to axial ribs and also swellings intermediate in position with reference to rilbs. Narrow subsututral cord faintly wvrinkled. Oni shells that hav-e narrow subsutural cord, con('ave lower part of fasciole generally steeply sloping; rarely gently sloplig, p)rodu~cing (list inctly turretedi outline. Concav-e lower part of fasciole bearing exagg-erated growth threads, some as strong as wrinkles onl stibstural cord. Faint spiral sculpture visible onl entire fasciole of some shells, especially onl early whorls. Pillar sculptured with widely spaced spiral cords and faint secondary spiral threads. A massive hiump at v-arying distance fromt outer lip ; ribs subdlited andl closely spaced between hum-rp and outer lip. Earlier hum1-tps, as many as 10, bult generally 1 to 5, Onl spire whorls of some shells. Outer lip flaring forward, lbeari-ng a (deel), wide stromiboid notch at top of pillar. Anal sinus (deep), lower limbh long, upper limib bordleredl by thick pad of callus onl )ai'ietal wall. Siplional notch wide and deep. Siphional fasciole slightly inflated, bearing no spiral sculpture or weak spiral sculpture, generally limited by a strong(, narrow thread, especially onl mature shells.
1 lel(iht (almost complete) 52.-I mm11, diameter (inchiding humip) 17j.5 mm11- (largest figured specimen). hteighlt (almost coinplete) 40.5 mmi, diameterr (including humip) 12..5 mmi figuredd specimen of interinedlate size).


Type: Tech. I-ocl schulle, Vienna.
Typ.Ie locality: lPresumnably Gatuin Locks excavation, Canlal Zonie, middle part of Gatun formation.
The V6 specimens of (Iiatlirodrillia from the Gatun formation (lower and middle parts, and upper part in eastern area) closely related to C/. gi)bosa are identified as C. gluesi'.Otherwise three names, presumatbly subspecific names, would lbe needed and some speci miens that show lintermnediate features would have to lbe placed doulbtfully under one of the three nam-es. Thle three main forms are as follows: Forms' of Cl atlirocirilIi a gattunensis in Gahmn formlation
1. Subsuitural cordl narrow, weakly wrinkled.
a. C'onicavte part, of anal fasciole below subsuitural cord steeply sloping (1)1. 58, figs.
24, 25).
1). Concav-e part of anal fasciole below subsutural cord gently sloping (p)1. 58, fig. 8).
2. Subsutural cord wide, strongly wrinkled (p1. 58,
figys. 20, 21, 2:3).
Typ)e la is represented ]in the lower part of the Gratim and in strata near the base of the middle part, althoughd sonm. specimens inthat part are transitional toward typie 2. Only one shell, collected at a middle (latumn locality in tile western area, represents type lb. It is associatedl with a specimen leaning toward type 2. The remiaining specimlens from the mnidd~le part that are larger than a few whorls and those from the upper part, 65percent of thle total numb11er, are unmistakably referred to type 2. That type agrees with Toula's type spiecimuen: an incomplete shell of six whorls (height 32.6 nun,11 diameter 12.3 mmi). is Pi~uotoina (Dirillia) galtouen. i. n. var. is a(n fimmature shell (height 10.1 11un,1 diameter 4.t mnm) It shows the 2-whorled, almost cylindrical protoconch. Type lb) closely duipli(ates the Ecuiadorean fossils named C.* onzola.
These Gratun -fossils could he further subdivided onl the basis of outline and spiral sculpture. One of five sp~ecimiens from locality 139c, is as slender as Crassi8p? ia cofNi's.. Onl a fewi like that shown onl plate 58, figures 20, 21, ti e spiral cords of' late whorls are wide and~ sep~aratedl by narrow groov-es. That specimen is one of' three that hias a p)erfectly 1)reserv-ed outer lip.
As nOW identified, (Clath rodiri/lia gatunwis occurs in (deposits of middle Miocene age in Venezuela, Colollia, Costa Rica, the Canal Zonie, the lDari6n and Chiriqui areas of' IPanamia, amid Ecuador, and at least a related form ]in the M\iocene of M\artinique (Cossimiana1918N, p). 22, 1)1. 2, Ti. ). If the indlefinite type localit.A of C. pteirloco'oiuNa represents the Las
1~erdice sh lte agre range extends back to early


382






GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO HELMINTHOGLYPTIDAE


Miocene. The Venezuelan shells identified as C. puertocololnliana have a few less ribs than Gatun shells (18 to 20 on penult whorl) and lack a strong, thread at the margin of the silphonal fasciole. A middle Mliocene specimen of C. puertocoloynbiana from Colombia (USGS 11241) has 25 ribs on the penult whorl and a strong( thread at the margin of the fasciole.
C. 'cemista (Sowerby) (1850, p. 50, pl. 10, fig. 7), which was well illustrated by Sowerby, was the first Milocene species of a closely knit group of ClatluodilVia to be described. It occurs in the Gurabo formation of the Dominican Republic and is distinguished by the absence of a subsuitural cord and by the attenuated continuation of the axial ribs across the entire anal fasciole. A lectotype was designated and illustrated by Pflug (1961, p. 65, pl. 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 origfinally in the Casey turrid collection, are in USNMI collections. It is found as a Pliocene fossil in the Cabo Blanco and Cuniand. areas of Venezuela. C. mnareana (W11eisbord, 1962, p). 437, pl. 41, figs. 16, 17; Pliocene, Venezuela) is an inmature 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. aalakowssa (Gardner) (1926-47, p. 310, pl. 40, figs. 2,4 1938), a small, slender species that has a moderately coiistricted anal fasciole. The younger C. subva) cosa (Gardner) (Idem, p. 312, pl. 40, figs. 5, 6; Oak Grove sand member of Shoal River formation) also has a moderately constricted anal fasciole. The fasciole of the still younger C. em-pera (Gardner) (Idem, p. 312, 1)1. 40, figs. 7, 8; Shoal River formation prIoper), however, is strongly constricted. That species, as Gardner realized, is like a small version of typ)e 2 of C. gatneinsis, lbut, at least on intermediate whorls, attenuated wrinkles more consistently extend across the anal fasciole.
No matter whether the synonymy of C. gatunelsi8 Is modified, the past and present distribution of C. gibbosa aiid its close allies, like that of lluracypraea, Sco)?sia and Xan ens, reverses the general trend. -Close 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 iMiocene). Lower part, localities 138c, 138d, 13Sf. Middle part, eastern area, localities 139b, 139c, 139e, 139fl 147f, 147g, 151, 155, 155a, 1551), 155c, 156, 157, 159a, 159b, 159d; western


383


area, locality 16la. Upper part, eastern area, localities 175, 17 6. Deposits of early (?) Miocene age, Colombia, Ecuiador. Deposits of middle M\,iocenie age, Venezuela, Colombia, Costa Rica, 1)ari~n area, Paniamd. (USGS 8477), Chiriqui area, Panai (USGS 7955), Ecuador.

Clathrodrillia saavedrai Woodring, n. sp.
Plate 59, figures 13, 14; plate 65, figure S
Small, subsutural cord noded, anal fasciole const ricted, pillar sh~ort. Protoconch blunt, 112-whorled, lower part of last half whorl swollen. First post-protoconch whorl sculptured with 8 axial ribs, swollen oni lower part, pinched off near suture. At later stages axial ribs narrow ; 17 or 18 on lpenu~lt whorl. Subsutural cord strong, bearing elongate retractive nodes, alined with axial ribs. A spiral thread marking lower edge of anal fasciole. Spiral sculpture below anal fasciole 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 hiump on body whorl about 450 from outer lip. Axial ribs subdued and irregular in spacing between hump and outer lip. Anal sinus deep, upper limb, heavily bordered. Stromiboid notch wide and deep. Siphional notch deep. Siphional fasciole slightly inflated, bearing no spiral sculpture, limited by a fairly strong- thread.
Height 16.5 umm, dianieter 6.2 mmn (type).
Type: USNMI 645790.
Type locality 14Thb (USGS 6033c, Panama Railroad, about 3,500 feet (1,065 in) southeast of Gatun railroad station, Canal Zone), middle part of Gatun formation.
Three specimens of Ciath rodrillia saavedrai have been found in the middle part of the Gatun formation. It suggests a miniature C. gatunevsis, but is distillgishied from iiimmature specimens of that species by its noded subsutural cord, more delicate sculpture, and weaker thread limiting the siphonal fasciole. Thlough~l the outer lip) of the type was well preserved, it was damaged in handling.
Thlis species is named for Alvaro de Saavedra Ceron, who early in the 16th century suggested Cenitral American routes to open the land from sea to sea (I-owarti, 1966, 1) 56-57).
Occurrence: Middle part of Gatun formation, eastern area (mi-iddle MIiocen~e), localities 147ib, 147h.

Clathrodrillia aff. C. saavedrai Woodring, n. sp.
Plate 58, figures 6, 7
Resem-bling4 C/at/u odrilia saaredrai in general featiires of protoconch and post-protocomich whorls, but distinguished as follows. Sinaller and more slender.






GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


Axial ribs narrower (16 011 penult whorl), pinched on lower part of anal fasciole, but continuing across entire fasciole, including subsutural cord. Axial ribs on sll1sittural cord sl iglhtly arcuiate on intermediate andl late whorls. Grooves between spiral lbandls wider than those of C. saa.cediai. Siphional fasciole sculptured with 2 coarse spiral threads.
1-eight 10 .4 nmm, diaml-eter r).5 mim (figured speciineii).
The idldle part of the Gatlin formation in the western area yielded the only specimen of this mniulte (lathjv'di/lia. TI omigl it evidently rep resents a new species, the outer lip) is damagedd and the shell is further disfiguried by two b~ore holes, which are large for the size of the shell.
Occurrence: Aliddle part of Gatun form-ation (m1-id(lie Mloce~ne), western area, locality 161c.

Clathrodrillia aff. C. lelandi (Olsson) Plate 59, figures 5, 6
Of inedium size, slender, whorls slightly inflated, pillar short. iProtoconcu and first few lpost-lprotoconch whorls ilissling. Anal fasciole slightly constricted on earliest preservedl whorls, (list inctly constricted on later whorls. Axial ribs narrow, slightly lprotractive, slitly bent, backward and lpinchied at lower edge of anial f.s i ole, (ouit ilin iIV across sulbsltiural cord imarkIng anal fasciole, straight or slightly arcuate on fasciole ; 21 on p~enuilt whorl. Spiral sculpture below anal fasciole consisting of' low hands (9 or 10 on p~enult whorl ), sep~aratedl by narrow grooves, faint except oil body whorl. Faint spiral grooves visible on anal fasciole adjoining anal sinuts. A humip on body whorl opplosite aperture. Axial ribs subdued and closely spaced between hump and outer lip. Aperture short. Stromnboid notch on outer lip shallow. Anal sinuis deep, upper Ii mb heavily bordered. Sipli onal notch deep), siphjonal fasciol)e l1iited by a strong thread.
Height (incomplete) 21.8 min, dliameter 7.5 mmn (figulred specimen).
This slender species is represented by one somewhat worn specimen from time lower part of the Gatuin formation. Owing( to wear, the spiral sculpture is faint, except, ot tihe abapertitral face of the body whorl near the outer i1 p. lThe whorls are less Inflated, or sholil(leredl, below time anal fasciole than those of' the middle Miocene 11ciadoreani A qliilla it-indi (Olsson, 1964, 1). 101, pl1. 15), fig. 7) and the slpiral bands are slightly wider.
it seems to b)e more appropriate to assign AgiadrdlNa ic/and; to ('lath odl,iiiia.
Occurrence : Lower part of Gatun formation (middl~e Miocene), locality 1136a.


Clathrodrillia cf. C. islalindae (Maury) Plate 58, figures 15, 16
Sm)Ilall, slender, whorls slightly inflated, pillar short. 1Protoconicl a mud first few post -protoconlih -whorls m-issig,. Anhal fasciole slightly constricted, except on late
halfof odywhorl. A xial ribs narrow, slightly protractivye, (oit mium"4 to stuire without being pinched at lower edge of' anal fasciole, arcuate on fasciole ; 16 to 18 onl penult, whorl. Spiral scullpture below anal fasciole consisting of low bands ((6 on penult whorl), sep~aratedl by narrow g-rooves. Faint spiral sculpture visible here and there, onl anal fasciole. A hum11-p Oil body whorl near outer lip. Apertulre short. Outer lip b~rokenI back to hiump, except at andl near anal fasciole. Anal slinus (feet), upper) limib heavily bordered. End of' siplhomd caiial broken. Si plional fasciole limited by a strongo, thol01111 wvorn thread.
I I( guit (not (ftlute comle~lte) 1.5.8 nill, dliameter (inci mding- hump) .5 mini (figured specimen).
Fomur illore or less worn specimens, also from the lower himit of time Gatlin formation, are identified as mnotliei sljecies of ('lath od/'irma. n~le Illustrated shell is I lhe largest amiid the. least worn. InI general features this slpecies is siular to C'. aff. CI. iCind;. It is, however,l silmal Icr, has less inflated whorls, and its ribs are iulinteiiriltedl and decidedly arcuiate on the anal fasciole. The rihs of C. HIai/nda? (Mlaury, 1917, p). 57, pl. P, fig. 7; ( ercado and Gurabo :format ions, Dom-inican Repihi ic) are simlillar, but are more p~rotractive, fewer in niiiuh1er, and therefore more widely spaced. M\oreoverl the spiral scuilpIture of that Species is stronger.
Occuirrence : Lower part of G atuin formation (n1-iddle .Il ocenle) ,localities 138d, 13Sf.

Clatlirodrillia? species
Plate 63, figures 1, 2
Of mediiun size, mioderately slender, -whorls mioderately Inflated, pillar short. Protoconich and early 1)0t-protoconich -whorls myising,. Anal fasciole molded agaist preceding whorl,1. progressively more strongly constricted. Akxial ribs wide, bent andi attenuated on anial tasciole, subdued on anal fasciole of body whorl and absent o)iu last quarter of that -whorl, 17 on p~enullt whorl. Spiral sculpltulre. faint, ex('ept on lower twothirds of' body whorl, where it consists of narrow, ('lose] .v spaced, crimped threads. A humip on body whorl about 600 from aperture. Aperture short. Outer 1i11) and~ lower pa rt of' sipliona I canal broken back. As shown by g-rowth lines, anal sinus deep); upper limb heavily bordered. Stromboid notch wide and very shmal low, asIindiocated by growth lines. Siphional fas(Mole not, bordlered by a strong(,, thread.


384






GASTROPODS: EIJLIMIDAE, MARGINELLIDAE TO IHELMINTHOGLYPTIDAE38


Heighit incompletet) 25.9 mm, diameter (including Iiumnp) 9.71 mmn figuredd specimen).
The Chagres sandstone yielded an incomplete, corrodled clavine turrid of undetermined affinities. The anal fasciole, molded against the preceding whorl in the fashion of some species of Latiras, is a diagnostic feature. The outer shell layer is missing on the crest of most body-whvlorl ribs and on all parts of spire whorls, except on the lower part of the penuilt, where faint spiral sculpture is discernible.
Thoug-h 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 ('lathrodrillia.
Occurrence: Chiag4res sandstone (late Miocene), locality 2008.
Genus Agladrillia Woodring
Woodring, Carnegie Inst. Washington Pub. 385, p. 157, 1928. Type (orthiotype) :Agladrilla callothyra Woodring, Miocene,
Jamaica.

Subgenus Agladrillia s.s.
Thle earliest species of Agladrillia s.s. now known occur in the late early Miocene rrhoionde formation of 11aiti. The subgenus survives in the eastern Pacific Ocean, but has not been recognized in the western Atlantic.
Trle four species in the Gatun formation, all of which are small, have a hump on the body whorl 900 or less from the outer lip. Between the hiump and the outer lip) the axial ribs5 are sup~pressed, irregularly and generally closely 0.11,ad eerly oe byth sp~iral banids, or are absent. As shown by the few shells that hiave an intact outer lip and by growth liles oil other shells, the anal sinus is deep and the stromboid notch shallow, although distinct. The sipbonud notch is moderately deep, and the siphional fasciole is slightly inflated and sculptured with relatively coarse slpiral threads. These noncliaguiostic 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, mnoderately inflated, anal fasciole slightly const rictedl, subsuitural cord generally faint, rarely distinct. Protoconch acute, 134- to 3-whorled (generally 134 to 212), 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, arcuiate on the cord; 11 to 15 ribs (generally 12 to 14) on penult whorl. Spiral sculpture below subsutural cord consisting of bands separated by rela-


tively deep and relatively wide grooves; 5 to T bands on lpenult 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 645T94.
Type locality: 147ig (USGS 5899, highest fossilbearing beds, Quebrauchia H-ills, 34 mile (1.2 ki) from G3atun, Canal Zone) middle part of Gatun formation.
Thlis species is the largest of the four in the Gatun formation. Trle relatively large size and more -uniform grTeater inflation distinguish it from the others.
A. chiaracta, 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 134 whorls and the lower part of the last -whorl is subcarinate. The spiral sculpture of two agrees with that of A. (latiacta, 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.
Aqlaa)l/da m vsOlsson (1964, p. 100, pl. 16, figs. 63, 6a ; Esmieraldas 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. characla. In spiral sculpture the Gatlin species resembles A. callot7?yra (Woodringo, 19298, p. 158, pl. 5, fig. 7 ), the type species, but that species is larger O(hit 20) mmn) aind has a more constricted anal fascdole, on which the axial ribs are subdued. Aside from its larger size and fe-wer, wNider ribs, A. callothyra is sinillar to Clat~owella ])aiwnaflla Dali (1908, p. 288, 1)1. 14, fig. 1). In DalIs illustration the ribs on the anal fasciole are too strong. Powell (1966, p. 92) suspected that Dali's species is to be assigned to Agladrillia. It has been dredged in the Gulf of California at depths of 47 to 153 fathomns.
Occurrence: Lowver and middle parts of Gatun formnation (middle Miocene). Lower part, locality 138a (identification doubtful), 138c, 138d (identification doubtful). Middle part, eastern area, localities 139b, 146, 147b), 147gr, 151, 159d.
Agladrillia (Agladrillia) enneacyma enneacymia (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. Protoconchi acute 3- to 312-whorled, lower part of last whorl swollen, rarely subcarinate. Axial ribs extending to suture. or almost to suture, on first post-


385






386 GEOLOGY AND PALEONTOLOGY OF CANAL ZONE


protoconcli whorl. On remaining whorls ribs extendiii~ across entire whorl of shells that have no subsutural cord, but gradually bent on anal fasciole, or on nonconstricted, part of whorl corresponding to anal fasciole, forming elongate nodes onl anal fasciole; 9 to 12 ribs oni penult whiorl. Spiral sculpture consisting of narrow bands separated by shallow grooves; 5 or 6 bands onl penu~lt whiorl, not~ including 1 to 3 narrow bands at lower edge of anal fasciole, or onl corresponding' part of whorls that lack a constricted fasciole.
1-leiglit 7.7 mmul, diameter (including hump) 2.5 mmn (figured slender specimenn. H-eight 7.2 mmn, diameter (including huminp) 2.8 mmn (figured inflated specimen).
Type: Acad. Nat. Sci. Phila. 3848.
Type locality: (Ttun lower locks excavation, Canal Zonle, middle part of Gatun formation.
rTle little glistening shells of Agladvillia eneacymna ewnneacyma were abundant in the middle part of the (4atun formation about a kilometer southeast of Gatun ralroad station during- th~e construction period, when fresh cuts were accessible onl the present line of the 1Panamia Railroad. A hundred shells, all. of which have the lprotoconcll preserved, were collected at locality 147b, where many other small species were found to be abundant,, and 45 at locality 147g. Shells that have ain intact outer lip, however, are rare. The upper part of the (iatiin yielded only one specimen and its spiral sculpture consists of narrow threads instead of bands.
This miinute Agiadi/Vla is variable in outline and in the presence or absence of a subsuitural cord. Aside from the single upper Gatun shell, the sculpture is basically uniform, although the spiral grooves on m-lany well -preserv ed shells are faint and the crest of the axial ribs of the type is exceptionally wide. r Te illustrations onl plate 59, representing two specimens selected from the large 147b collection, show some of the range in variation. The contrast between the unpinchied early axial ribs of A. enneaoyma en'neacyina andl the pinched ribs of A. eliaracta is shownv by figures
1 and 4 onl plate 64.
A larger similar species occurs in the Thomondle formation of Haiti and a smaller species inl the Cerado and Gurabo formations of the I)oinican Republic. Thle early axial ribs of 1)0th aire lpinclied.
Occurrence: Middle andl upper parts of Gatun formnation (middle Milocene). -Middle p)art, eastern area, localities 139b, 139cl 146, 147b, 147f, 147g,~n 147h, 153a,
1 55c, 159d. Upper part, eastern area, locality 173a.

Agladrillia (Agladrillia) enneacymia (Brown and Pilsbry), sulbsp.
Fomur specimens from the lower p)art of the Gatun formation are (list ingi shed from the nominate sub-


species of Agiadvillia emneacymza by the presence of a smbsutural cord onl all except the first few post-protoconchi whiorls and by relatively dleep) grooves between the spiral bands, like those of A. (iairaeta. These shells hiave the l)1otoconlil and unlpinchied early axial ribs of the nominate subspecies. Two (hieit, 8.7 and 9.5 mm) aire larg-er than any shells of the noininate subspecies. The sample of this form is inadequate.
Occurrence : Lower part~ of Oat un formation (id(Ile Milocenle), localities 107, 138, 138c.

Agladrillia (Agladrillia) phengoides Woodring, n. sp.
Plate 59, figures 1, 2 ; pliate 64, figure 3
Very small, slender, anal fasciole constructed, sublsutural cordl narrow, moderately strong. Protoconch bl imlit, 11/.). to 134-whiorled, lower p)art of about last half whiorl subcarinate. Axial ribs pinched off near sufli re onl first, few lpost-l)1otoconcll whiorls. Subsutural cordl app)ea1rii onl fourth or fifth post-protoconch whlorl. Onl later whiorls axial ribs p~inchied at lower edlge of sti lsmtu ral cord, forming11 small, retractives nodies onl the cord, 19 or 10 rib~s onl penult whorl. Spiral 5(1111t ire b)el ow sulbsutural cord consisting) of narrow biinds separated by shallow grooves ; 15 to 7 bands on plenumlt whiorl, not including 1 or 2 minor bands present oil soiiie shells near edge of' subsutural cord. Grooves necar si lisultural cord (list inct, indistinictl, reven absent.
I 1eighmt 6.8 nun,1 diameter (includingr hump) 2.2 11-m1 (type).
Type: UISNM~ 645 799.
Tyelocality: 1471) (ITS(OS 6033c, PuanaRal
roa1.d, about 3,,500 feet (1,065 meters) southeast of (Thtunl railroad station, Canal Zone), middle p)art of Oatliun formIiatilonl.
Thie slender outline, blunt, paucisplral 1)rotoconchi, and moderately strong, noded subsutural cord onl intermlediate andi late whiorls are cliaracteristic, features of this little gl isteiiing- species. Unlike Agladrila en nea('yina einueacyiiia, it is not variable in basic features, but tie spiral grooves uuear the subsutural cord may be (list iuct, inldist iict, or absent.
A. )licihqo;dcsN was found in the middle part of the ( latun format ion. 'Ihle largest num11ber of specimens
(46) wvas collected at the type locality, where it is associ-atedl with .4. (1Irt/Y~a anid A. e)Owa(?/flia en neacellia. It is, in fact, associatedl with A. en neacyinma enneacyina ait thie six localities where it, occurs.
The, iore slender oiti me, more strongly constricted aaIascile aind more, bullgin wh vorls distinguiish Ai. 1p/tenrqoide8 from A1. e,e/lania (Olsson) (1922, p). 69, p]. 10, fig-s. 31, 32), a miiddle. iI\ iocenie Costa Rican Species.


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