Cretaceous Actaeonellid Gastropods from 

the Western Hemisphere

GEOLOGICAL SURVEY PROFESSIONAL PAPEU 1304
Cretaceous Actaeonellid Gastropods from 

the Western Hemisphere

By NORMAN F. SOHL and HEINZ A. KOLLMANN

GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304

A description of 28 species of 

actaeonellid gastropods, with a discussion 

of the evolution, life habits, ecology, 

and Cretaceous world distribution of 

the family

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON: 1985
DEPARTMENT OF THE INTERIOR 

WILLIAM P. CLARK, Secretary

U.S. GEOLOGICAL SURVEY 

Dallas L. Peck, Director

Library of Congress Cataloging in Publication Data

Sohl, Norman Frederick, 1924-

Cretaceous actaeonellid gastropods from the Western Hemisphere.

(Geological Survey professional paper ; 1304)

Bibliography: p.

Includes index.

Supt. of Docs, no.: I 19.6:1304

1. Actaeonellidae. 2. Paleontology Cretaceous. 3. Paleontology America. I. Kollmann, Heinz A. II. Title. III. Series.

QE809.A18S63 1984 564'.37 83-600250

For sale by the Distribution Branch, U.S. Geological Survey, 

604 South Pickett Street, Alexandria, VA 22304
CONTENTS

Abstract 

Introduction 

Acknowledgments - 

Page 

1

1

2

2

2

2

4

4

______________________________ 4

Substrate preference 6

Salinity tolerance 7

Early ontogeny and dispersal potential of actaeonellids 9

Reflections of soft parts on the actaeonellid shell 

Columellar retractor muscles 

Character of the mantle edge 

Musculature of the head-foot mass 

Exhalant area 

Distribution and diversity -

Evolution of the Actaeonellidae -

Stratigraphic summary of Western Hemisphere actaeonellid

occurrence -

Early Cretaceous record -

Late Cretaceous record -

Locality register 

Systematic paleontology - 

Catalogue of actaeonellid species9

20

22 

24 

24 

35 

46 

74

Genera and species excluded from the Actaeonellidae 87 

References cited 90 

Index 97

ILLUSTRATIONS

[Plates follow index]

PLATE 1. Polished section of oriented rock surface from roadcut, Erath County, Tex.

2. Trochactaeon, Actaeonella

3-16. Trochactaeon

17. Actaeonella, Trochactaeon

18-23. Actaeonella

Page

FIGURE 1. Morphologic features of the actaeonellids 3 

2. Postulated salinity ranges for various Cretaceous faunal assemblages in which actaeonellid gastropods occur 8 

3-4. Histograms of 

3. Species diversity of the subgenera of Trochactaeon during the Cretaceous 10

4. Actaeonella species diversity during the Cretaceous 10 

5-9. Occurrence of Trochactaeon 

£) ^Duriiisr thG BsnT^mi&n ______ - ______________ ___________________________ _______ 11

6. During the Aptian and Albian 12

7. During the Cenomanian and early Turonian 13

8. During the late Turonian to Santonian 14

9. During the Campanian and Maastrichtian 15 

10-13. Occurrence of Actaeonella 

11. During the Cenomanian and early Turonian 17

12. During the late Turonian to Santonian 18

13. During the Campanian and Maastrichtian 19

14. Phylogeny of the Actaeonellidae as presented by three previous authors 2 1

15. Proposed phylogeny of the Actaeonellidae 23

16. Correlation of middle Cretaceous formations of Puerto Rico and Texas 25 

17-18. Correlation of the Upper Cretaceous rocks of 

17. Puerto Rico 26

18. Central and western Jamaica 27

19. Stratigraphic ranges of the Western Hemisphere actaeonellid gastropod species 28

m
IV CONTENTS

FIGURES 20-25. Index maps showing 

20. Collection localities in Puerto Rico 29

21. Collection localities and occurrence of Cretaceous rocks in Jamaica 30

22. Collection localities and areas of Cretaceous outcrop in Cuba 31

23. Collection localities in Texas and New Mexico 32

24. Collection localities in Mexico and Guatemala 33

25. Collection localities in California 34 

26-45. Scatter diagrams comparing height with width among selected specimens of 

26. Trochactaeon (Trochactaeon) woodsi (Rennie) 49

27. Trochactaeon (Trochactaeon)packardi (Anderson) 52

28. Trochactaeon (Neocylindrites) cumminsi Stanton 53

29. Trochactaeon (Neocylindrites) nelsoni Sohl and Kollmann, n. sp. 55

30. Trochactaeon (Mexicotrochactaeon) conformis (Bose) 57

31. Trochactaeon (Mexicotrochactaeon) burckhardti(B6se) 58

32. Trochactaeon (Mexicotrochactaeon) wrighti Sohl and Kollmann, n. sp. 59

33. Trochactaeon (Mexicotrochactaeon)palmeri Sohl and Kollmann, n. sp. 60

34. Trochactaeon (Mexicotrochactaeon) granthamensis Sohl and Kollmann, n. sp. 61

35. Trochactaeon (Mexicotrochactaeon) rossi Sohl and Kollmann, n. sp. 62

36. Actaeonellapecosensis Stanton 65

37. Actaeonella delgadoi Choffat 66

38. Actaeonella browni Sohl and Kollmann, n. sp. 67

39. Actaeonella borneensis Nuttall and Leong - 68

40. Actaeonella duckettsensis Sohl and Kollmann, n. sp. 69

41. Actaeonella jicarensis Sohl and Kollmann, n. sp. 70

42. Actaeonella coquiensis Sohl and Kollmann, n. sp. 70

43. Actaeonella marchmontensis Sohl and Kollmann, n. sp. 71

44. Actaeonella briggsi Sohl and Kollmann, n. sp. 72

45. Actaeonella cubensis Sohl and Kollmann, n. sp. 73
CRETACEOUS ACTAEONELLID GASTROPODS FROM 

THE WESTERN HEMISPHERE

By NORMAN F. SOHL and HEINZ A. KOLLMANN1

ABSTRACT

Actaeonellid gastropods constitute one family of the cephalaspidean 

opisthobranch superfamily Acteonacea. During the Cretaceous they 

were one of the most common biotic elements in the shallow warmwater faunas of the Tethyan Realm. They were common associates of 

the rudist framework ("reef) tracts that in the Western Hemisphere 

ranged throughout the Mexico-Central America-Antillean-Gulf 

Coast region. Maps are presented showing the worldwide distribution of these gastropods during five time intervals of the Cretaceous 

Period. Their greatest latitudinal extent was during Albian to 

Cenomanian times; subsequently, they were restricted to narrowed 

Tethyan and marginal Tethyan areas. One paleolatitudinal anomalous occurrence in Alaska may be related to current concepts of 

suspect allocthonous terranes.

Actaeonellids show a progressive increase in numbers of species 

from their first appearance in the Barremian Stage to a high in the 

late Turonian to Santonian interval. Subsequently, their diversity 

declined during the Campanian and Maastrichtian, and, like their 

common associates the nerineid gastropods and rudist bivalves, they 

became extinct at the end of the Cretaceous. Their record in the 

Western Hemisphere is at some variance with that of actaeonellids 

in the rest of the world. During the latest Cretaceous, they maintained a high level of diversity in the Caribbean to Mexico region 

and showed a tendency toward taxonomic provincialism. This latter 

feature is demonstrated by the appearance in the Maastrichtian of 

species of Actaeonella having only two instead of three columellar 

plications and by the presence of an endemic subgenus Trochactaeon 

(Mexicotrochactaeon) in rocks of Coniacian to Maastrichtian Age.

Evidence is presented to indicate that actaeonellids possessed two 

distinct adult life habits. At postlarval metamorphosis, both 

Actaeonella and Trochactaeon were initially epifaunal. Subsequently, 

Actaeonella assumed an infaunal habit, whereas Trochactaeon remained epifaunal to adulthood. In addition, studies of associated 

biotic elements suggest that the two genera possessed different 

salinity tolerance; Trochactaeon (Trochactaeon) had the broader 

range of tolerance, ranging well into environments of low salinity.

Only two genera, Actaeonella and Trochactaeon, are recognized as 

present in the Western Hemisphere. Trochactaeon is subdivided as 

follows: the nominate subgenus, T. (Neocylindrites), T. (Sevanella), 

and T. (Mexicotrochactaeon). Of these, only T. (Sevanella) lacks representation in this hemisphere. The earliest occurrences in the 

Western Hemisphere of species of Actaeonella and Trochactaeon 

(Neocylindrites) are in rocks of Albian Age. The reason for their 

absence in rocks of suitable pre-Albian facies is unknown. In total, 14

'Naturhistorisches Museum in Wien, Geologisch-Palaontologische Abteilung, Vienna, 

Austria.

species of Trochactaeon are described, seven of them new. In addition, three more species are distinguished but not formally proposed 

because of insufficient material. Actaeonella is represented by 14 

described species (nine new), and one species is distinguished but 

unnamed because of inadequate information.

A final section of this report consists of a catalogue of the described 

Cretaceous species of actaeonellids. This listing is the data base used 

for the construction of the maps of distribution and the histograms of 

diversity with time.

INTRODUCTION

Knowledge of the Cretaceous molluscan faunas of the 

Western Hemisphere is very unequal with respect to 

geographic areas, biologic group, and stratigraphic interval. For example, mollusks of the late Late Cretaceous of the Coastal Plains of eastern North America 

are quite well documented, whereas those of the Caribbean region to the immediate south are, overall, poorly 

known. In the Caribbean region, only one molluscan 

group, the rudist bivalves, is at all reasonably known. 

In general, knowledge of the Western Hemisphere 

faunas of the Cretaceous temperate marine seas is 

much greater than that of faunas of the Tethyan 

(warm-water tropical) biogeographic province. The 

present work is intended to give detailed information 

about the Actaeonellidae, one of the major molluscan 

groups distributed throughout the warm-water Tethyan 

Realm.

The Actaeonellidae constitute but one family of the 

cephalaspidean opisthobranch superfamily Acteonacea. 

As here conceived, they were restricted to the Cretaceous Period. Their extinction at the end of the Cretaceous coincided with that of the rudist bivalves, 

nerineid gastropods, and other major faunal associates 

of the Tethyan warm-water biota. Although the actaeonellids were of low generic diversity, the almost 

200 species recorded in the catalogue section of this 

report show that though their geologic history was 

relatively short, they were a very viable group.

The actaeonellid gastropod fossils described herein 

lived in the tropical Tethyan and marginal Tethyan
CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

seas. They were common faunal elements associated 

with the Cretaceous carbonate platform communities 

most noted for their abundance of rudist bivalve framework-building organisms (Kauffman and Sohl, 1974). 

In many parts of the Tethyan Realm they were sufficiently abundant to constitute principal constituents of 

rock bodies (Kollmann, 1965, fig. 23, Sohl, 1971, fig. 1). 

In the Eastern Hemisphere, their taxonomy and distribution has received considerable attention by paleontologists (Kollmann, 1967; Hacobjan, 1972, 1976, 

Djalilov, 1972; Pchelintsev, 1953); in the Western 

Hemisphere, however, no such synthesis has previously 

been attempted for the group. One major reason for 

this lack of knowledge has been the reconnaissance 

level of geologic work in much of the region that contains outcrops of Tethyan-belt Cretaceous rocks. In recent years, knowledge has increased because of increased interest in the geology of the outcropping 

Cretaceous rocks of Mexico, Central America, and the 

Caribbean area. This interest has been engendered in 

part by the common occurrence of hydrocarbon resources in rudist bivalve framework structures and 

associated rocks and in part by the role the Caribbean 

region plays in global plate-tectonic theory.

Much of the material that forms the basis for this 

paper was collected by Sohl (1956-1974) during a U.S. 

Geological Survey cooperative program with the Commonwealth of Puerto Rico. The aim of this program 

was to provide total geologic mapping of that island. 

Concurrent studies were carried on by Sohl and others 

on the Cretaceous rocks of Jamaica because of the presence there of less structurally disturbed sequences that 

afforded better opportunity to establish stratigraphic 

ranges of taxa. The remainder of the material described 

has been assembled from collections housed at various 

institutions throughout the United States.

ACKNOWLEDGMENTS

Many individuals and institutions facilitated our 

work by loan of specimens contained in their collections or by providing other tangible aid. The Smithsonian Institution is due special thanks for support of 

Kollmann during his tenure as a Postdoctoral Fellow. 

Christopher Burden of the Texas Memorial Museum in 

Austin; Peter A. Rodda of the California Academy of 

Sciences, San Francisco; the late Katherine Van Winkle 

Palmer of the Paleontological Research Institute at 

Ithaca, N.Y.; and Keith Young of the University of 

Texas at Austin arranged for loans of specimens housed 

at their respective institutions. Ted Robinson of the 

University of the West Indies in Kingston, Jamaica, 

provided Sohl with maps, hospitality, and aid in shipping collections from that island. Anthony G. Coates,

George Washington University, Washington, B.C.; 

J. E. Hazel, U.S. Geological Survey, Reston, Va.; and 

E. G. Kauffman, University of Colorado, Boulder, aided 

in collection of fossils and in pertinent discussion of 

stratigraphic problems in the field and office.

REFLECTIONS OF SOFT PARTS ON THE 

ACTAEONELLID SHELL

Certain anatomical features of the Actaeonellidae 

and their relative placement can be deduced from shell 

characters.

COLUMELLAR RETRACTOR MUSCLES

Longitudinal grooves on the upper quarter of the 

parietal area of the body whorl of Trochactaeon have 

been described by Kollmann (1967). They occur about 

one-half of a whorl behind the aperture and in some 

species may be divided by a series of small ridges. We 

note herein similar structures in Mexicotrochactaeon. 

These longitudinal grooves are depressions of the hypostracum, marking the position where soft parts adhered 

closely to the shell. Positioning of this longitudinal 

band of grooves on the body whorl well behind the 

aperture suggests that this feature marks the position 

of the columellar retractor muscles. The work of Fretter 

and Graham (1954) supports this interpretation. They 

figured a specimen of the cephalaspidean Actaeon 

tornatilis with the shell removed, which shows the 

columellar muscle occurring as a band on the upper 

part of the whorls. Lemche (1956) recorded muscles in 

the same location on Cylichna. His anatomic investigations, however, suggest that these muscles function as 

retractors of the mantle.

CHARACTER OF THE MANTLE EDGE

The form of the aperture provides some information 

on the nature of the mantle margin. For example, in T. 

(Trochactaeon), T. (Neocylindrites), and T. (Mexicotrochactaeon), a parietal inductura extends over the 

body-whorl surface outside the aperture and over the 

columella, with a margin bending anteriorly toward 

the concave basal lip. The boundaries of this inductura 

indicate the limits of the extension of the mantle outside the aperture (fig. 1O. In contrast, no inductura 

has been observed on shells of Actaeonella. The basal 

lip of Actaeonella is somewhat reflected (pi. 19, fig. 8), 

suggesting a thickening of the mantle, which may 

have served as a device against intrusion of sediment 

into the mantle cavity.
REFLECTIONS OF SOFT PARTS ON THE ACTAEONELLID SHELL

6 and C modified from Kollmann, 1967

FIGURE 1. Morphologic features of the actaeonellids. A, View of a Trochactaeon shell as seen from the anterior; arrows indicate 

postulated directional trends of (1) upper columellar muscle, (2) middle columellar muscle, (3) lower columellar muscle. 

B, Shell as viewed in cut section, showing shell layers and internal features: (1) ostracum, (2) first-formed hypostracum, 

(3) last-formed hypostracum, (4) palatal or outerlip plication or fold, (5) columellar plications or folds. C, External shell features: (1) posterior or anal sinus, (2) parietal lip with inductura, (3, 4) columellar plications, (5) columellar lip with inductura. 

(B and C modified from Kollmann, 1967.)
CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

MUSCULATURE OF THE HEAD-FOOT MASS

All actaeonellids possess columellar plaits (figs. IB 

and 1O, but strength and extent of both the plaits and 

the excavated areas between plaits may vary. The upper, or posteriormost, columellar plait invariably extends out of the aperture onto the body-whorl surface. 

The second or next anterior plait normally terminates 

at the aperture. That is, it ends on the columellar surface precisely at the intersection of the outer lip and 

apertural margin plane. When present, a third and 

anteriormost plait never reaches the aperture margin 

plane. The plaits themselves represent the position of 

hypostracum crests that separate parallel muscle 

bands. The differing lengths of the plaits, plus the 

slight difference in their angle of inclination relative to 

the shell axis, suggest that the muscle bands served 

different functions relative to the head-foot mass. The 

interplait areas trend in an anterior direction overall. 

The incision above the first, or posterior, columellar 

plait, is the most strongly depressed. In Actaeonella, T. 

(Neocylindrites), and T. (Trochactaeon), it is a rather 

narrow incision, but in T. (Mexicotrochactaeori) the 

incision is broad and terminated posteriorly by a parietal plait. According to the trend of the incision, the 

muscle must have had a basic horizontal action function (fig. 1A). The incision between the posterior and 

second plait trends more toward the left side of the 

shell as viewed from the anterior. The incision between 

the second and third, or anteriormost, plait trends 

downward and to the right (fig. 1A). The fact that the 

upper (posteriormost) muscle-insertion area extends 

outside the aperture suggests that the soft parts could 

not be completely retracted. When these fossils are 

compared with some living neogastropods having columellar plaits (Ponder, 1973), one may postulate that 

the actaeonellid interplait muscle bands were used to 

control head and foot movements during burrowing or 

ploughing through the substrate.

EXHALANT AREA

Many authors (Cossmann, 1895; Delpey, 1939; 

Termier and Termier, 1952; Pokorny, 1959; and Kollmann, 1967) have pointed out the importance of the 

structure of the posterior aperture margin of the 

actaeonillid shell for interpreting life habits of the 

group. On shells of Trochactaeon, the posterior edge of 

the outer lip bends back in a sinus curve for approximately one-sixth of a whorl before it becomes attached 

to the penultimate whorl. This posterior slit is closed at 

the attachment point by a subsutural ramp that is 

formed by hypostracum layers. This suggests that shell 

material was deposited by the mantle surface and not 

at the mantle edge in the vicinity of the sutural ramp.

The posterior sinus is in a position analogous to the 

right posterior end of the mantle chamber, an area that 

probably also bore the anal aperture. Obviously, the 

posterior sinus (fig. Ic) marks the position where currents carrying fecal wastes were extruded. This interpretation is consistent with similar structures and 

functions in such living Cephalaspidae as Actaeon, 

Philine, Scaphander (Fretter and Graham, 1954), and 

Cylichna (Lemche, 1956), as well as in certain neogastropod groups that possess columellar plaits and have a 

burrowing habit (Ponder, 1973).

ECOLOGY

Actaeonellids were common members of Cretaceous 

faunal assemblages that are widely distributed in a 

belt paralleling the Equator, a belt commonly called 

the Tethyan province or realm (Sohl, 1971, and figs. 

5-13). The actaeonellid pattern of distribution is similar not only to that of such other gastropods as the 

nerineids Trajanella, Pileolus, and Discotectus but also 

to that of the corals that Beauvais and Beauvais (1974) 

have termed "building corals" and to that of the rudist 

bivalve framework builders (Kauffmann and Sohl, 

1974). These faunal associations have long been considered to delimit a tropical biotic realm. The distribution 

shown by some actaeonellids suggests that they may 

have had a wider temperature tolerance than others. 

For example, node-bearing forms of Trochactaeon are 

found not only in normal warm-water assemblages in 

the Caribbean and in Transcaucasian Armenia but 

also from both northern and southern extremes of 

actaeonellid distribution. The noded species T. 

(Trochactaeon) woodsi (Rennie) occurs in the Cretaceous 

of Pondoland, South Africa, associated with a fauna 

otherwise similar in composition to that of the North 

American Ripley Formation that Sohl (1964) judged as 

a temperate fauna. North of the Tethyan belt, noded 

species such as T. (Trochactaeon) beyrichi (Drescher, 

1863) occur in Coniacian-Age rocks in Poland and 

Czechoslovakia (Fric, 1911; Andert, 1934), where the 

associated fauna contains only one other species, 

Nerinea bicincta Bronn of Andert (1934), of presumably 

warm-water affinities.

LIFE MODE

Direct evidence of life mode of Actaeonella is found in 

an occurrence near Morgan Mill in Erath County, Tex., 

mentioned in the section on substrate preference and 

illustrated on plate 1. In the lower part of the sectioned 

slab is a bioturbated zone about 6 cm thick. Parts of 

both vertical and diagonal burrows having a circular
ECOLOGY

cross section can be recognized in this zone. Cut sections occasionally intercept specimens of Actaeonella 

within the burrows (see arrow "A" on pi. 1). The specimens sectioned are oriented with the anterior part of 

the shell inclined downward. Both the form of the burrows and orientation of the shells therein agrees well 

with Fretter and Graham's (1954) observations of the 

burrowing behavior of Actaeon tornatilis (Linne). These 

opisthobranch gastropods begin burrowing into the 

substrate in a vertical or diagonal direction. After a 

few centimeters of penetration they turn to a lateral or 

horizontal mode of travel and then eventually move 

back toward the sediment surface. During burrowing, 

the head lobes and pedal gland are pressed against the 

sand; secretions then agglutinate the sand particles 

and cement the burrows, preventing collapse as the 

animal moves on.

The close similarity of the burrows of Actaeonella 

and Actaeon and the similar orientation of the shells of 

both genera within the burrows is strong evidence for 

postulating an infaunal habit for Actaeonella. In addition, the general shell form of Actaeonella and T. (Neocylindrites) suggests an infaunal life habit. Not only 

does the spindle-shaped outline suggest such a habit, 

but the funnel-shaped posterior siphon in Actaeonella 

and the sutural slit of T. (Neocylindrites) are analogous 

to similar sanitary devices in other infaunal gastropods (Kollmann and Yochelson, 1976). Ponder (1973) 

has pointed out that infaunal neogastropods have both 

a similar siphonal canal system and columellar plaits.

Although Trochactaeon possesses a posterior siphonal 

slit, other characters suggest that it had a life mode differing from that of Actaeonella. In contrast to the 

smooth streamlined spindle-shaped shells of Actaeonella, Trochactaeon has a relatively high spire and 

a proportionally broad shell; some species even bear 

coarse nodes on the whorl shoulder. Overall, these 

characters do not appear to be conducive to an infaunal 

habit, but infaunal habit is found in other living snails 

of similar form. The infaunal tonnacean Cassis madagascarensis spinella Clench, is large, possesses noded 

whorls, and superficially resembles Trochactaeon. As 

Moore (1956) noted, this species plows through the substrate in search of its prey, the echinoid Plagiobrissus. 

Though grossly similar to Trochactaeon, Cassis differs 

significantly in its widely expanded aperture that 

reflects a great broad muscular foot, a feature not present in Trochactaeon.

The great variability in shell character within individual species of Trochactaeon may also suggest a primarily epifaunal existence. This variability is especially well expressed in proportional height of the spire 

and in maximum diameter of the body whorl compared 

with total height. Such variability far exceeds any

comparable variation in Actaeonella. The work of 

Levinton (1973) and others provides an explanation. In 

studying extant bivalves, Levinton has demonstrated 

that below the sediment-water interface, temperature 

and salinity variations are at a minimum. Similarly, 

as water depths increase, there is a correlative increase 

in environmental stability. Levinton concluded that 

species living in variable environmental settings 

tended toward polymorphism, whereas those living in 

the more stable situations tended toward isomorphism. 

Struhsaker (1968) has demonstrated a similar relationship in the Gastropoda. She found that polymorphic 

species of Littorina were better adapted to withstand 

environmental extremes than were isomorphic species. 

Thus, Actaeonella fits the model of an isomorphic form 

living in a stable situation such as that provided by 

an infaunal habit, in contrast to the polymorphic 

Trochactaeon.

We can see no direct correlation between a gastropod 

having a proportionally higher spire or greater maximum diameter and the ability to withstand greater 

environmental variation. Thus, we assume that these 

characters were genetically linked with other characters that together provided some advantage.

Other lines of evidence suggest that Trochactaeon 

was dominantly an epifaunal dweller. For example, 

many shells show the effects of mechanical wear and 

either repair of shell or internal secondary shell deposition (pi. 15, fig. 8). In many shells, numerous additional 

shell layers are deposited in the spire and internally 

separate the body or later whorls from those formed 

earliest. Kollmann (1967) has noted that the hypostracum shell layers are exposed near the suture. These 

layers are not as resistant as those of the ostracum and 

are the first affected by mechanical wear such as wave 

action. Thus, probably in response to irritation of the 

mantle, secondary lamellae were deposited in the posterior part of the whorl (pi. 15, fig. 9). The fact that 

apical whorls could be sealed off indicates the ability to 

withdraw the visceral mass from the earlier parts of 

the shell, that is, those parts most endangered. Such 

withdrawal logically could cause a shell-space problem 

that was not totally alleviated by continued growth. 

This problem may have been countered by resorbtion 

of some shell wall and columellar structures (pi. 15, fig. 

9). Both secondary deposition of lamellae and resorbtion phenomena have been observed in T. (Trochactaeon) but have not been noted in Actaeonella and T. 

(Neocylindrites).

The foregoing lines of evidence suggest that Trochactaeon lived in shallow, well-agitated water and that if 

it ever was a dominantly infaunal organism, it became 

increasingly epifaunal in habit during the Late Cretaceous. Such a view would be consistent with the fact
CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

that species such as T. (Mexicotrachactaeon) coniformis 

(Bb'se), which is one of the stratigraphically youngest 

species, shows the highest degree of shell polymorphism.

Not all workers accept such a view of the life habits 

of Trochactaeon. For example, Herm and Schenk(1971) 

concluded that, on the basis of the proportionally narrow aperture and comparatively large and thick shells, 

Trochactaeon was not very mobile and possessed a 

rather stable life position. This interpretation has several weak points. For example, such a sedentary mode 

would be compatible only with a filter feeding or 

mucous-string feeding habit. Both these feeding modes 

are unknown among other opisthobranchs (Purchon, 

1968). The closest analogues, the Cephalaspidea, are 

all collectors of organic detritus. In addition, one may 

also cite both the Cypraeidae and Conidae as groups 

that have shells of at least moderate size, possessing a 

proportionally narrow aperture and yet not being sedentary. Herm and Schenk (1971) have also applied 

Vogel's (1968) suggestion that the curvature or growth 

direction of a radiolitid on a gastropod shell in a direction opposite that of the growth direction of the outer 

lip indicates that it grew upon the shell of a live snail. 

Among the specimens investigated by Herm and 

Schenk, 72 percent show radiolitids attached in a 

fashion consistent with the growth of both rudist and 

Trochactaeon; 28 percent do not. Fretter and Graham 

(1954) mentioned that in all cephalapidean gastropods, 

the shell aperture is to the right of the midline of the 

shell. Even if the aperture is rather narrow, which, in 

actaeonellids, it appears to be only when the edge of 

the outer lip is broken, an empty shell has greater stability relative to overturning on the aperture side. During the growth of a radiolitid that settled upon a dead 

shell, the probability of rotation or overturning toward 

the less stable side that is, opposite the aperture is 

highest. Thus, most radiolitids would show a growth 

direction that Herm and Schenk believe is characteristic for growth on a shell of a living gastropod. Finally, 

the high concentration of shells at the locality they 

studied suggests accumulation after death. If this is 

true, such a shell pavement would provide a prime 

hard substrate for settlement of rudist spat and the 

subsequent growth of radiolitids. In summary, we do 

not see that the evidence of narrow aperture, size, and, 

rarely, radiolitid attachment provides a compelling 

case for interpreting Trochactaeon as a sedentary epifaunal gastropod.

On the basis of the evidence and interpretations presented above, we maintain that Actonaeonella and T. 

(Neocylindrites) were infaunal burrowers much in the 

fashion ofActaeon tornatilis and that T. (Trochactaeon) 

and T. (Mexicotrochactaeon) were primarily epifaunal 

vagrant benthos elements.

SUBSTRATE PREFERENCE

Direct evidence for substrate preference of actaeonellids is rare. Most such evidence points to a preference for finer grained bottom sediments. For example, 

in Cretaceous exposures in roadcuts immediately 

northwest of Morgan Mill in Erath County, Tex., 

Actaeonella sp. was found in living position (pi. 1, fig. 1, 

and preceding discussion on life habits). The rock is a 

calcareous, silty fine sand. Turritella is the only associated larger fossil. We interpret the Actaeonella as an 

infaunal burrower and believe that the Turritella probably paralleled its living counterpart T. communis by 

living apex down in the muddy sand substrate in its 

filter-feeding mode (Merriam, 1941; Yonge, 1946; 

Thorsen, 1965; and others). Low-diversity associations 

of this type are not uncommon today in relation to 

cephalaspidean opisthobranch gastropods. For example, Parker (1975) has noted that in the Woods Hole, 

Mass., vicinity, Cylichna oryza (Totten) and Retusa 

canaliculata (Say) occur together in clayey silt substrates where other gastropods are rare. Similarly, 

Herm (1977), in his studies on the faunal assemblages 

of the northern calcareous Alps, has emphasized the 

low diversity in his "Trochactaeon association." These 

"associations" are divided first into a RadiolitesTrochactaeon group occurring in a well-sorted, mediumgrained sand that is interpreted as a high-energy beach 

deposit. Laterally, these deposits grade into "Trochactaeon sands," in which shells are concentrated by wave 

action. Herm also described obviously autochthonous 

Trochactaeon from poorly sorted silty sands that contain beds of pebbles, a deposit that he interprets as 

tidal flat.

Actaeonellid-dominated Cretaceous deposits are 

sporadic throughout the Caribbean Antilles and in 

Central America. In general, these occurrences parallel those from Europe described above. Actaeonella 

occurs in some deposits to the virtual exclusion of other 

molluscan taxa. Chubb (1959, p. 743), in describing the 

uppermost sandy limestone of the Nuevo beds of 

Chiapas, Mexico, stated that "A. cf. laevis occurs in incredible profusion in all exposures; other fossils are 

generally comparatively rare." These Actaeonella are 

referred to A. borneensis herein (pi. 19, figs. 16-21) and 

occur in a matrix of fine calcareous quartz sand containing large Foraminifera, rounded concentric-growth 

algal spheres (5 mm diameter), and echinoid debris. 

The large (8 cm) slender actaeonellids are well preserved, showing little evidence of transport. The combination of biotic elements suggests a shallow nearshore 

deposit of moderate energy level. Another Actaeonelladominated assemblage from the Melones Limestone of 

southwestern Puerto Rico has been figured by Sohl
ECOLOGY

(1971, fig. 1). The actaeonellas are in a calcareous 

tuffaceous limestone containing abundant angular 

coarse clasts of volcanic rocks and generally finer 

rounded bioclastic fragments of echinoids and rudists.

Not all Actaeonella occur in sand. For example, 

Actaeonellajicarensis Sohl and Kollmann, is found in a 

muddy limestone in Puerto Rico (Smith and others, 

1968, p. G2), in which tall slender nerineids and Turritella are the other dominant species. Nerineids are 

usually the most abundant, but locally Actaeonella sp. 

may dominate.

T. (Trochactaeon) woodsi (Rennie), a noded form, in its 

occurrences in Puerto Rico is found both in a mechanically concentrated bed at locality 39 and as the dominant constituent in limestone at localities 41 and 47 

(fig. 20). The latter limestones indicate that the substrate was basically a lime mud, but allocthonous 

rounded coarser shell debris is not uncommon.

T. (Mexicotrochactaeon) is most common in shaly intervals interbedded with rudist-dominated limestones 

in such sections as that along the Rio Minho in the 

Central Inlier of Jamaica (localities 64 and 65, fig. 21). 

The matrix is generally a silty clay containing fine 

shell and plant fragments. The associated fauna is 

moderately diverse, including a few species of infaunal 

bivalves, the thin-walled small rudist Distefenella, and 

trochiform and strombid gastropods. Kauffman and 

Sohl (1974) have viewed these as lagoonal associations.

The above observations can only vaguely suggest 

that Trochactaeon and its allies preferred calcareous 

muds, whereas Actaeonella was more commonly associated with somewhat coarser clastic sediments.

SALINITY TOLERANCE

The well-studied Upper Cretaceous sequences in the 

Alps provide us with the best information relative to 

salinity tolerance of the actaeonellids. Earlier authors 

(Zittel, 1881-1885; Zapfe, 1937; Pokorny, 1959) usually 

considered the fossils to have lived in brackish water. 

This was suggested to explain the concentrated occurrence of Trochactaeon in the northern calcareous Alps. 

In addition, these authors considered both Actaeonella 

and Trochactaeon to belong to the same genus. On the 

basis of a comparison of the accompanying fauna, Kollmann (1965), in contrast, demonstrated that Actaeonella 

and Trochactaeon possessed differing salinity tolerance. Although Kollmann found no suggestion that 

salinities higher than Polyhalinikum (classification of 

Remane, 1963, see also fig. 2) could be tolerated, 

Schenk (1972) demonstrated that both genera could 

tolerate the higher concentrations of the Euhalinikum. 

In figure 2, occurrences of Actaeonella, T. (Neocylindrites), and T. (Trochactaeon) have been plotted against

Remane's salinity zones. As indicated on this figure, T. 

(Neocylindrites) and Actaeonella do not pass into salinity zones below 18 ppm. In the Alpine Cretaceous, 

these two genera occur in association with rudist and 

coral faunas, which are commonly quite taxonomically 

diverse. Similar occurrences may be documented from 

numerous Western Hemisphere localities. Such limitations of distribution support the view that the habitats 

of T. (Neocylindrites) and Actaeonella have a closer 

affinity than do the habitats of either of them to that of 

T. (Trochactaeon).

T. (Trochactaeon) presumably had a broader salinity 

tolerance, ranging from Meiomesohalinikum (5-10 

ppm) to Euhalinikum (30-40 ppm). The association in 

shales of Trochactaeon with Glauconia, Neritopsis, 

Pirenella, and Pseudomelania led Kollmann (1965) to 

postulate a low-salinity occurrence. Similar associations are documented elsewhere in the Alps, such as 

Herm's (1977) Polymesoda-Glauconia association. In 

addition, similar faunas are found near Dreistaetten in 

lower Austria and Vorderstoder in upper Austria. The 

constancy of the composition of these assemblages is 

strong support for the view that Trochactaeon was a 

component of many brackish-water faunas.

In many sections, sandstones contain densely packed 

Trochactaeon shells. In discussion of such occurrences, 

Nuttall and Leong (1972, p. 6) stated: "It seems likely 

that many concentrations of shells of one species occur 

as a result of differential sorting after death. The most 

acceptable explanation is that actual actaeonellid limestones were formed of current-sorted shells probably in 

very shallow water." We agree that many such paucispecific occurrences are the result of transport and size 

sorting. However, some occurrences of Trochactaeondominated, low-diversity assemblages appear to us to 

represent a biocoenosis. For example, Herm (1977) 

reported sands containing abundant Trochactaeon of 

various size grades (growth stages) and associated with 

a few specimens of the coral Astrocoenia, the gastropod 

Patella, and the bivalves Pedalion and Tellina. The 

sands are highly bioturbated by Callianassa, but the 

assemblage is deemed autochthonous. This sand 

assemblage grades laterally into other sands that contain size-sorted Trochactaeon. Herm (1977) also recorded Trochactaeon in an assemblage dominated by 

the bivalve Cucullaea and the gastropod Ampullina. 

Because rare specimens of Glauconia and Polymesoda 

are also associated in this assemblage, it is assigned to 

the Pleiomesohalinikum (10-18 ppm) salinity zone.

Present knowledge of the occurrences of T. (Neocylindrites), Actaeonella, T. (Mexicotrochactaeon) and T. 

(Trochactaeon), in the Caribbean region is insufficient 

to indicate salinity tolerance to the degree indicated 

for the assemblages discussed above. This insufficient
SALINITY ZONES

(FROM REMANE

1963)

Euhalinikum 

30-40 ppm

Polyhalinikum 

30-18 ppm

Pleiomesohalinikum 

10-18 ppm

Meiomesohalinikum 

5-10 ppm

Oligohalinikum

0.5-5 ppm

FAUNAL ASSEMBLAGES

All marine including: 

Anthozoa 

Cephalopoda 

Archaeogastropoda

Decreasing number of marine 

species, almost no euryhaline 

species

Marine fauna dominant, number 

of species reduced, few 

brackish species

Minimum of species, extreme 

brackish-water organisms 

dominant, no echinoderms

Limnic fauna dominant, but

low number of species

LOCALITIES

Uchaux, France (Roman and Mazeran, 1920) 

Texas, Devils River Limstone 

(Stanton, 1947) 

Saxony (Geinitz, 1874) 

Gosau, Randograben (Felix, 1908)

Brandenberg, Tyrol, TrochactoaeonCucullaea-Association (Herm, 1977) 

Gams, Noth (Kollmann, 1963)

Brandenberg, Tyrol, TrochactaeonCucullaea-Association (Herm, 1977)

Russbach, Traunwand (Kollmann, 1975)

Vorderstoder, Upper Austria (Kollmann, 

unpublished) 

Dreistaetten, Lower Austria (Ploechinger,

1961 and Kollmann, unpublished)

FIGURE 2. Postulated salinity ranges for various Cretaceous faunal assemblages in which actaeonellid gastropods oc
DISTRIBUTION AND DIVERSITY

knowledge is primarily due to the lack of study of the 

molluscan faunas in general in the area and especially 

to the dearth of quantitative analyses.

EARLY ONTOGENY AND DISPERSAL 

POTENTIAL OF ACTAEONELLIDS

Maestrichtian-age shales at several localities in 

Jamaica have yielded immature actaeonellid specimens that show the character of the protoconch and its 

relationship to the early teloconch development. These 

specimens indicate a significant difference in early life 

history between Actaeonella and Trochactaeon.

In Actaeonella, the early sequence of growth may be 

seen in the specimen figured on plate 2, figure 4. This 

specimen (USNM 305999) from the "Titanosarcolites 

limestone" of the Marchmont inlier (locality 97) of 

Jamaica, although poorly preserved, shows a complete 

heterostrophic protoconch at the apex together with 

the initial phase of the teloconch. The protoconch or 

larval stage has a diameter of 0.24 mm and is delimited from the teloconch by a narrow incision. On this 

and a second specimen (pi. 2, fig. 3, USNM 305998) 

from this locality, one may note that the first three 

teloconch whorls form a turriculate shell. These whorls 

are convex sided and show no subsutural ramp. After 

these three initial whorls, shell geometry changes; 

whorl overlap increases so rapidly that after one more 

whorl, most of the teloconch is enveloped. The change 

from an initial turriculate teloconch stage to later total 

envelopment is also shown in the illustration (pi. 23, 

fig. 5) of a section cut through the center of a specimen 

of Actaeonella cubensis.

The clear distinction between Actaeonella and 

Trochactaeon is well seen by contrasting figures 3 and 

4 with figures 1 and 2 on plate 2. The two specimens of 

Trochactaeon (Mexicotrochactaeon) sp. from the Guinea 

Corn Formation of Jamaica illustrated here show that 

the protoconch is similarly heterostrophic (see pi. 3, 

figs. 1-4), and it is clearly separated from the teloconch 

by an incision. The maximum diameter of the protoconch, however, is 0.18 mm, and the whorls are circular in cross section. The early whorls of the teloconch 

have the same elongate narrow shape with subsutural 

ramp as those of the later shell.

The major differences thus indicated between Actaeonella and Trochactaeon early growth stages are: 

1. The diameter of the protoconch is larger in specimens of Actaeonella than in Trochactaeon (Mexicotrochactaeon). Kollmann (1967) also found a similar relationship in actaeonellids from Gosau, 

Austria; the protoconch of T. (T.) lamarcki consisted of two volutions having a maximum diameter of 0.1 mm, whereas that of A. laevis exceeded 

0.2 mm.

2. The early teloconch whorls of Trochactaeon (Mexicotrochactaeon) have a subsutural ramp and are 

almost exactly like those of the later teloconch. 

Kollmann (1967) noted the same character for T. 

(Trochactaeon). In contrast, Actaeonella undergoes a two-stage change in shell geometry, first, a 

change in coiling direction between larval and 

neanic stages (postprotoconch) and second, a 

change in overlap of the whorls after the third 

convex-sided teloconch whorl culminating in an 

involute shell.

Although the protoconch size differs in Actaeonella 

and Trochactaeon, it may have little significance. In 

both genera the ratio between the number of protoconch volutions and the maximum diameter places 

them in the same category as those prosobranchs having a long to moderately long pelagic larval stage 

(Shuto, 1974). More information is needed to determine 

whether protoconch size is consistent in the group, but 

the distribution of the family during the Cretaceous 

supports a high larval dispersal capability.

The second noted contrast between Actaeonella and 

Trachactaeon is more significant. The incision separating protoconch from the teloconch certainly marks the 

metamorphosis or change from a pelagic life stage to a 

benthonic life mode for both. We suggest elsewhere 

that Trochactaeon was epifaunal as an adult, and the 

outogeny would suggest that this mode was assumed at 

metamorphosis. By contrast, we have presented evidence that the adult Actaeonella was of infaunal habit. 

The early teloconch turriculate-form developmental 

stage, however, may mark a short epifaunal crawling 

stage that preceded the assumption of the adult infaunal mode that is marked by the adult Actaeonella 

involute shell form.

DISTRIBUTION AND DIVERSITY

Figures 3-13, showing species diversity and distribution of the Actaeonellidae, have been derived from the 

information presented in the accompanying species 

catalogue. The data are divided into five time categories that to some degree crosscut stage boundaries. 

These five intervals are: Barremian; Aptian to Albian; 

Cenomanian to lower Turonian; upper Turonian to 

Santonian; Campanian to Maastrichtian. We realize 

that this breakdown into several large time intervals 

may obscure some important fluctuations in diversity 

and distribution. This objection is offset by the fact 

that precise age assignment of many many taxa is suspect. Acceptance of worldwide time equivalency of such
10 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

60

50

UJ 40 

U

UJQ_

LL. 

O

QC 

UJ 

CO

^

=> 

Z

30

20

10

PROPORTIONAL STAGE DURATION 

IN MILLIONS OF YEARS (M.Y.)

EXPLANATION

Trochactaeon (Trochactaeon) 

Trochactaeon ( Neocylindrites)

Trochactaeon ( Me.ricotrochactaeon) 

Trochactaeon (Sevanella)

FIGURE 3. Histogram of species diversity among the subgenera of 

Trochactaeon during the Cretaceous. Width of the columns is proportional to the duration of the stages represented. 1, Barremian 

(4 m.y.); 2, Aptian-Albian (18 m.y.); 3, Cenomanian-lower Turonian 

(6 m.y.); 4, upper Turonian-Santonian (7 m.y.); 5, CampanianMaastrichtian (19 m.y.).

assignments is variable, and construction of maps and 

graphs on that refined a basis would introduce many 

errors of fact. We have confidence that in accepting 

these broader time intervals, the minor "noise" fluctuations that the erroneous age assignments would cause 

is smoothed out, and the concommitant potential for 

misinterpretation is thus reduced.

Figures 3 and 4 show diversity of Actaeonellidae 

with time. The patterns of peak diversity for Trochactaeon and Actaeonella show a remarkable consistency. 

Both show progressive increase in number of species 

from their first appearances to a high in the late 

Turonian to Santonian, followed by a decline in CamLi.

O 10

QC 

UJ 

CO R

PROPORTIONAL STAGE DURATION 

IN MILLIONS OF YEARS (M.Y.)

FIGURE 4. Histogram of Actaeonella species diversity during the 

Cretaceous. Width of the columns is proportional to the duration of 

the stages represented. 1, Aptian-Albian (18 m.y.); 2, CenomanianTuronian (6 m.y.); 3, upper Turonian-Santonian (7 m.y.); 4, 

Campanian-Maastrichtian (19 m.y.).

panian to Maastrichtian time. This decline, however, 

does not reach as low a level as that during the virtually equally long Aptian to Albian time. One might 

argue that the maximum is a reflection of provincial 

taxonomy, biased by the unusually large numbers of 

described species from the Transcaucasus (Pchelintsev, 

1953, 1954; Hacobjan, 1972; Djalilov, 1972).

If these species from the Transcaucasus are discarded 

from consideration, a large number of species in the 

Mediterranean region and elsewhere remain. Thus, 

although the peaks of diversity might be less accentuated if a more stringent species concept was used, the 

diversity peaks do seem to have objective validity.

The Cenomanian to Santonian interval also marks a 

time of wide distribution of Actaeonellidae in the 

Tethyan realm and in such marginal areas as northern 

California, Brazil, and East Africa (figs. 7, 8, 11, and 

12). The general lack of actaeonellids at this time in 

the Caribbean is striking. This absence coincides with 

what Khudoley and Meyerhoff (1971, p. 116) consider 

the major stratigraphic break in the Cretaceous sequence of the Antilles, that is, "Cenomanian or early 

Turonian below and the Campanian or Maastrichtian 

above." They envision this as a period of erosion during which few basins received sediments. Where sediments are present, they are deeper water mudstones 

and volcaniclastic rocks. In summary, this period was 

one of little or no carbonate-platform sedimentation. 

Locally in Mexico and California, a few actaeonellids 

are present.

In three aspects, the Campanian through Maastrichtian interval stands in marked contrast to the upper 

Turonian to Santonian interval. The first aspect is a 

lesser diversity, the second, a longer time interval, and 

the third, a declined proportional geographic representation. Few Campanian and Maastrichtian species
150° 120° 90° 60° 30° 30° 60° 90° 120° 

60°

30°

0° -

30°

60C

EXPLANATION

Occurrence of Trochactaeon during

the Barremian 

T. (Neocylindrites)

J_______L

Base modified from Smith and others, 1977

FIGURE 5. Geographic distribution of Trochactaeon during the Barremian. The reconstructed position of landmasses follows Smith and Bride

shelf areas. Plus signs represent the arbitrary line of separation of one continent from another.
30°-

60°-

150' 120° 

Occurrence of Trochactaeon during

the Aptian and Albian 

T. (Neocylindrites)

Base modified from Smith and others, 1973

FIGURE 6. Geographic distribution of Trochactaeon during the Aptian and Albian. The reconstructed position of landmasses follows Smith a

deletion of shelf areas. Plus signs represent the arbitrary line of separation of one continent from another.
DISTRIBUTION AND DIVERSITY 13

e

I

«

TJ

ca

43 «j

1

cc 

I

41

.o -5

^ S 1-3

8 2 B&0 CO

r T .p-i 

O 03

-5 

c

g .2

o
150° 120° 90° 60° 30° 30° 60° 90° 120° 

60°

30° -

0° -

30°

60°

EXPLANATION

Occurrence of Trochactaeon during 

the late Turonian to Santonian

 T. (Trochactaeon)

 T. (Neocylindrites)

 T. ( Mexicotrochactaeon) 

o T. (Sevanella)

Base modified from Smith and others, 1973

FIGURE 8. Geographic distribution of Trochactaeon during the late Turonian to Santonian. The reconstructed position of landmasses follows S

with deletion of shelf areas. Plus signs represent the arbitrary line of separation of one continent from another.
150° 120° 90° 120° 

EXPLANATION

Occurrence of Trochactaeon during 

the Campanian and Maastrichtian

 T. (Trochactaeon)

 T. (Neocylindrites)

A T. (Mexicotrochactaeon)

60° ! -

Base modified from Smith and others, 1973

FIGURE 9. Geographic distribution of Trochactaeon during the Campanian and Maastrichtian. The reconstructed position of landmasses foll

(1977) with deletion of shelf areas. Plus signs represent the arbitrary line of separation of one continent from another.
150'

Occurrence of Actaeonella during 

the Aptian and Albian

30°-

60°,-

Base modified from Smith and others, 1973

FIGURE 10. Geographic distribution of Actaeonella during the Aptian and Albian. The reconstructed position of landmasses follows Smith an

deletion of shelf areas. Plus signs represent the arbitrary line of separation of one continent from another.
DISTRIBUTION AND DIVERSITY 17

o3 o3

f I C L, 03 <*-i

i i .jj

<~ C

O CD

11

O O

2 c

tf -2

O iCD o.

a >.

>> 

S &

.

.£ §

I!

O |£j

If

-Q £

IE

ii i*

o
150'

Occurrence of Actaeonella during 

the late Turonian to Santonian

30'

60° '-

Base modified from Smith and others, 1973

FIGURE 12. Geographic distribution of Actaeonella during the late Turonian to Santonian. The reconstructed position of landmasses follows S

with deletion of shelf areas. Plus signs represent the arbitrary line of separation of one continent from another.
150° 120° 

Occurrence of Actaeonella during 

the Campanian and Maastrichtian

60° -

Base modified from Smith and others, 1973

FIGURE 13. Geographic distribution of Actaeonella during the late Campanian and Maastrichtian. The reconstructed position of landmasse

Briden (1977) with deletion of shelf areas. Plus signs represent the arbitrary line of separation of one continent from anoth
20 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

occur in the old diversity centers of Mediterranean 

Europe and the Transcaucasus, but the CaribbeanMexico region possesses the highest diversity of both 

Actaeonella and Trochactaeon. By Maastrichtian time, 

the Caribbean can almost be deemed a refugium.

Among the Actaeonellidae, the subgenus T. (Neocylindrites) was the first to appear. It is recorded from 

the Barremian of southern France by two species, 

Trochactaeon (Neocylindrites} barremicus Sayn and T. 

(N.) boutillieri Cossmann (fig. 5). T. (Neocylindrites) 

never attained the diversity of the other subgenera 

during any unit of time considered here (fig. 4), but 

representatives maintained a presence throughout the 

remainder of the Cretaceous. T. (Neocylindrites) attained its greatest distribution during Albian time (fig. 

6), its range extending from Central Asia through 

Mediterranean Europe to Angola and westward to 

Texas and Baja California. By Cenomanian to early 

Turonian time (fig. 7), T. (Neocylindrites) was restricted 

to the area from the Transcaucasus to central Europe. 

The Campanian Maastrichtian species are restricted to 

the Western Hemisphere and are represented by T. 

(N.) nelsoni Sohl and Kollmann in Puerto Rico and T. 

(N.} punctatus (Maldonado-Koerdell) in the State of 

Chiapas, Mexico.

The nominate subgenus T. (Trochactaeon) shows a 

rapid and continuous increase in species from its first 

appearance in the Cenomanian through Turonian time. 

This increase is followed by a decrease to the Campanian to Maastrichtian interval. This viable group, however, maintains the highest diversity of any of the subgenera throughout its stratigraphic range (fig. 4). 

Although widely distributed in Europe, Africa, and the 

Middle East during Cenomanian time, no Western 

Hemisphere representatives are known until the Turonian when T. (T.) frazierensis and T. (T.)packardi were 

present in California (fig. 8). Maximum geographic 

range of Trochactaeon was probably attained in Campanian time (fig. 9), species being reported from such 

disparate areas as Pondoland, South Africa; the 

Trichinopoly area of southern India; and the Assam 

Province of northeastern India. Maastrichtian occurrences appear restricted to the Caribbean region in the 

form of (T.) (T.) woodsi (Rennie).

The first record of T. (Mexicotrochactaeon) is in the 

Coniacian of Mexico (fig. 8). The subgenus subsequently 

spread throughout the Caribbean region and is to be 

found in the highest Maastrichtian beds of the Antillean 

islands. It thus appears to be the last survivor of the 

genus.

T. (Seuanelld) is a short-lived Coniacian to Santonian 

endemic subgenus restricted to Transcaucasian U.S.S.R. 

(figs. 4, 8).

Actaeonella first occurs in beds of Aptian Age; it is

represented by A. fusiformis Coquand from Spain and 

Lebanon and A. baconica (Benkb-Czabalay) from Hungary. By the end of the Albian, species of Actaeonella 

are to be found eastward from this center to the Transcaucasus, southward to West Africa, and westward 

through the Caribbean and Texas to Baja California. 

Actaeonella never attained as high a species diversity 

as did Trochactaeon (compare figs. 3 and 4), but it is 

found in areas not reached by the latter (figs. 10-13). 

Most of these extensions are latitudinal, into marginal 

Tethyan areas. For example, in the Cenomanian and 

early Turonian (fig. 11), Actaeonella is found well north 

of Tethys in eastern Siberia, and in North America, it 

occurs well outside the normal Caribbean track on 

Martha's Vineyard, Mass. Perhaps the most unusual 

occurrence is that of Actaeonella sp. C, which is found 

in the Yukon River valley of Alaska. This far-north 

occurrence is so anomalous that it may best be explained within the framework of "suspect terranes" 

(Coney and others, 1980). That is, the host rock is part 

of an allocthonous or transported mass accreted to the 

North American Cordilleran margin after the middle 

Cretaceous. Maastrichtian distribution of Actaeonella, 

like that of Trochactaeon, was much reduced. Actaeonella was represented in Europe only by A. oliviformis in France and A. agricolai Vidal from the 

Garumnien of Spain. Despite its restricted world geographic range, Actaeonella remained diverse and abundant in the Maastrichtian deposits of the Caribbean 

region into the youngest Cretaceous rocks known in 

the area (fig. 13).

EVOLUTION OF THE ACTAEONELLIDAE

Kollmann (1967) has discussed the relationships between Cylindrites and the Cretaceous Actaeonellidae. 

He suggested that Neocylindrites was a direct descendant of Cylindrites that differed only by possessing a 

larger number of columellar plaits. He considered 

Neocylindrites (Barremian to Maastrichtian) to be the 

conservative stem from which all other actaeonellids of 

middle and Late Cretaceous age descended (fig. 14).

This phylogenetic relationship has been modified by 

Hacobjan (1972, 1976) and Djalilov (1972) (see fig. 14 

for comparison). Djalilov (1972), dealing only with 

Actaeonella and other involute forms, derived them 

directly from Ptychocylindrites (Jurassic to Early 

Cretaceous), a nearly involute form with a depressed 

apical part. Among these involute genera, Omphactaeonella and Eotrochactaeon have been removed from 

the Actaeonellidae by Kollmann and Sohl (1979) and 

assigned to the Itieriidae. Hacobjan (1972, 1976), on 

the other hand, is in favor of separate phylogenetic
Hacobian, 1972 and 1976

Kollmann, 1967

FIGURE 14. Phylogeny of the Actaeonellidae as presented by three previous authors.
22 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

origins for the spired and involute forms of Actaeonellidae. He derived Actaeonella from Actaeonellina 

Pchelintsev, 1963, and T. (Neocylindrites) from Cylindrites. Although morphologic agreement is good among 

the genera Ptychocylindrites, Actaeonellina, and Actaeonella, we do not agree with the direct descendance 

of Actaeonella from one of the two other genera, as was 

proposed by Djalilov and Hacobjan. The paleontological 

record shows that T. (Neocylindrites) occurred much 

earlier in the Cretaceous than did Actaeonella. Both 

Cylindrites and T. (Neocylindrites) are found in Barremian rocks, but neither Actaeonellina or Ptychocylindrites occur in Lower Cretaceous rocks. For this 

reason, we maintain the phylogenetic position of T. 

(Neocylindrites) as the older and conservative stem in 

the family from which all other, mostly Late Cretaceous forms, have descended (fig. 15). Actaeonella arose 

from this stem in the late Aptian by an increase in 

overlap of the whorls, leading ultimately to an involute 

shell form. Evidence for this evolutionary transition 

stage is shown by Actaeonella fusiformis Coquand from 

the Aptian of Spain and Lebanon. In this species, the 

overlap is still not total, so that the early whorls are 

still visible.

The normal base plan of three columellar plaits is 

present throughout the geographic and stratigraphic 

range of Actaeonella. The species described in this 

paper that bear only two plications occur in the 

Maastrichtian of Cuba, Jamaica, and Mexico (fig. 15) 

and are considered only a congeneric Caribbean endemic offshoot of the primary Actaeonella lineage.

Trochactaeon (Trochactaeon) is proposed to have 

evolved from T. (Neocylindrites) during the upper part 

of the Cenomanian. The transition involved an increase 

in proportional breadth of the shell and an increase in 

spire height. In general, Cenomanian species such as 

T. (Trochactaeon) matensis (Fittipaldi) and T. (Trochactaeon) pseudocylindraceus (Pschelintsev) remain morphologically close to T. (Neocylindrites), as the whorls 

are only slightly inflated. In the Western Hemisphere, 

this form group, which was named Mesotrochactaeon 

by Hacobjan (1972, 1976), is represented by the California Turonian species T. (Trochactaeon) packardi 

(Anderson) and T. (Trochactaeon) frazierensis (Anderson) (pi. 8).

Trochactaeon (Mexicotrochactaeon) makes its first appearance in the Coniacian rocks of Mexico (fig. 15). It 

differs from T. (Trochactaeon) in possession of a parietal 

fold or swelling. T. (Mexicotrochactaeon) is restricted in 

distribution to the Western Hemisphere, but in this 

area it diversified rapidly, and, with the exception of 

the noded T. (Trochactaeon) woodsi, it replaced 

Trochactaeon s.s. completely. The fact that Actaeonella 

and T. (Neocylindrites) both maintained worldwide distribution through the Late Cretaceous may suggest 

that typical Trochactaeon had a lower dispersal potential. This factor, compounded by the progressive separation of the Caribbean area from Europe during postCenomanian time, may have created an interrupted 

gene flow leading to endemism. To some degree, as discussed previously, this view is supported by the smaller 

protoconch size in Trochactaeon as compared with that 

in Actaeonella. In addition the only Trochactaeon to occur in post-Turonian rocks are forms related to T. (T.) 

woodsi, distinctive by being noded Trochactaeon as 

well as being related to forms from South Africa rather 

than to those from Europe.

Trochactaeon (Sevanella) Hacobjan (1972) evolved 

from Trochactaeon (Trochactaeon) and consists of a 

short-lived (Coniacian to Santonian) group of species 

endemic to the Armenian SSR. They are all heavily 

noded shells with laterally depressed whorls.

Actaeonellids became extinct at the end of the Cretaceous. This extinction parallels that involving many 

other groups such as nerineid gastropods and rudist 

bivalves that were dominant components of the Tethyan 

shallow-water Cretaceous marine assemblages. Whatever cause one proposes for these extinctions, the fact 

remains that the Actaeonellidae were declining in 

diversity during the latter part of the Late Cretaceous 

prior to extinction (figs. 3 and 4). Parallel to this decline 

in the diversity of actaeonellids was a rise in the diversity of gastropod groups of similar habit. Cephalaspidean gastropods and neogastropods are dominantly 

carnivores and both increased markedly in diversity 

during Late Cretaceous time. The early evolution of 

the neogastropods was restricted to the boreal and 

temperate seas (Kollmann, 1979). During the Albian, 

the neogastropods invaded the Tethyan seaways in 

which the actaeonellids were already established. During the Late Cretaceous, the neogastropods increased 

significantly, occupied increased numbers of ecologic 

niches, and in essence must have come into direct competition with the Actaeonellidae. Also during this 

time, new feeding habits in other groups such as the 

Naticidae (Sohl, 1969) suggest a concomitant effect on 

existing food chains and disruption of prevailing biological equilibrium on the existing surface and substrate of the sea bottom. That the decline of the 

actaeonellids coincides with these events seems more 

than mere coincidence.

STRATIGRAPHIC SUMMARY OF WESTERN 

HEMISPHERE ACTAEONELLID OCCURRENCE

Actaeonellid gastropods occur sporadically but are 

locally abundant in Cretaceous rocks of post-Aptian
STAGES

MAASTRICHTIAN

CAMPANIAN

SANTONIAN

CONIACIAN

TURONIAN

CENOMANIAN

ALBIAN

APTIAN

BARREMIAN

HAUTERIVIAN

RANGE AND RELATIONSHIPS

Actaeonella

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EVOLUTIONARY EVEN

-0 

Actaeonellid Extinction

n First occurrence Actaeonella with two

plications (Endemic to the Caribbean) 

 First occurrence of Trochactaeon with

noded whorls in Western Hemisphere

Q Replacement of T. (Trochactaeon) bv

T. (Mexicotrochactaeon) in Western Hemisph

O First occurrence of T. ( Trochactaeon)

_ First occurrence of fully involute Actaeonell

O Species transitional between T. (Neocylind

and Actaeonella

O First occurrence of T. (Neocylindrites)

FIGURE 15. Proposed phylogeny of the Actaeonellidae. Solid bars indicate ranges in Western Hemisphere; outlined bars indicate ranges in 
24 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Age throughout the Western Hemisphere (figs. 16-25). 

Their absence in rocks of Aptian Age is unexplained, 

as seemingly suitable facies were present in such areas 

as southern Mexico (Allencaster de Cserna, 1956) and 

the Benbow inlier of Jamaica (Sohl, 1979), where their 

common associate the nerineid gastropods are abundant. Figure 19 shows that actaeonellids are most 

diverse during the Albian and the Campanian to 

Maastrichtian time in the Western Hemisphere. This 

might be predicted from the fact that this time coincides with widespread carbonate-shelf deposition in the 

Caribbean to Middle America region, thus providing 

optimum environmental conditions for the establishment of actaeonellids. This phenomenon finds a parallel in the episodic rudist-framework-building activity 

noted by Kauffman and Sohl (1974,1979) for the Caribbean region.

In this region, correlation of the units within which 

actaeonellids occur is based primarily upon a rudist 

bivalve biostratigraphy. The endemic nature of the 

rudist faunas, however, precludes their utilization for 

intercontinental correlation. Age assignments are 

therefore based upon the conventional biostratigraphic 

tools of ammonite, inoceramid, or planktonic foraminifer associations. For some areas, such as Texas, the formations are firmly dated because of the intensive work 

that has been done and the abundance of biostratigraphic information. In other areas, such as the Antillean islands, critical dating fossils are rare in the dominantly volcaniclastic sequences. Thus, the correlations 

of formations presented on figures 17 and 18 rest 

mainly on summaries provided by Pessagno (1979) for 

foraminifers, Kauffman (1979) for inoceramids, and 

Sohl (1979) for ammonites and other macrofossils. The 

rudist bivalve ranges presented on these figures are 

based primarily on the studies by Dommelen (1971) 

and MacGillavry (1977) and on unpublished observations of Sohl.

Precise information on the localities at which the 

actaeonellids discussed here were found is provided in 

the locality register (p. 48). These localities have been 

given numbers keyed to general locality maps (figs. 20 

to 25) and to the sections on occurrence in the species 

descriptions. Undoubtedly some new taxa and many 

new finds of actaeonellids will be made as work proceeds beyond the existing reconnaissance level of investigation, especially in Hispanola, Central America, 

and parts of Mexico.

EARLY CRETACEOUS RECORD

The record of Early Cretaceous actaeonellids in the 

Western Hemisphere is sporadic and restricted to rocks

of Albian Age (fig. 16). In the Antillean islands, 

reported occurrence of actaeonellids is restricted to the 

eastern and central parts of the island of Puerto Rico.

Where found, as in the Barrancas Limestone Member 

of the Magueyes Formation in Puerto Rico (fig. 16), 

Actaeonella is a rare constituent in muddy lagoonal 

limestone associated with Caprinuloidea rudist bivalve 

framework sequences (Kauffman and Sohl, 1974, fig. 

23, C) of early to middle Albian Age. Elsewhere, 

actaeonellids are locally abundant in the area from 

Guatemala northward to Texas and thence westward 

to Baja California. In all these areas they occur in rock 

sequences containing intervals in which faunas are 

dominated by rudist bivalves. Trochactaeon (Neocylindrites) cumminsi Stanton and Actaeonella peconsensis 

Stanton are especially common in the middle to upper 

Albian biosparites of the Devils River Limestone in 

southwestern Texas (fig. 23, localities 134, 135, 138, 

139), where they are associated with abundant nerineid 

gastropods. Trochactaeon (Neocylindrites) cumminsi, 

however, is not restricted to biosparites, as it also occurs in tuffaceous siltstone of the Alisitos Formation in 

Baja California (Allison, 1955).

In Guatemala, Actaeonella browni is abundant and is 

associated with a diverse mixed assemblage of intertidal to shallow subtidal grass-flat mollusks and clusters of small rudist bivalves of probable late Albian 

Age.

LATE CRETACEOUS RECORD

Occurrence of actaeonellids in Upper Cretaceous 

rocks of the Western Hemisphere is better known. The 

occurrences are, overall, similar to the faunal associations of the Lower Cretaceous and parallel the track of 

the rudist-framework facies. Several notable exceptions to this pattern exist. The first is the occurrence of 

a few specimens of Actaeonella sp. (pi. 22, fig. 7) in 

Cenomanian rocks of Martha's Vineyard, Mass. Here 

(Sohl, 1977, p. 203), their association with cerithiid 

gastropods, Brachydontes, and other bivalves suggests 

a marginal marine, lagoonal to estuarine assemblage. 

This occurrence is latitudinally well north of rudistid 

distribution and probably represents an incursion of 

the genus north into more temperate or at least marginal Tethys waters during middle Cenomanian time.

The other anomalous Late Cretaceous occurrence is 

that of Trochactaeon (Trochactaeon) frazierensis and T. 

(T.) packardi (Anderson) in sandstones in northern 

California (fig. 25) in rocks of Turonian Age, well north 

of normal rudist distribution. The association here of 

Trochactaeon species with such genera as Trajanella 

suggests a northward incursion of warm-water species 

into a generally temperate province.
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FIGURE 18. Correlation of the Upper Cretaceous rocks of central and western Jamaica.
STAGES

STRATIGRAPHIC RANGES 

OF 

SELECTED RUDIST BIVALVES

Trochactaeon (T.) tvoodsi (Rennie) 

T. aff. T. (T.) woodsi (Rennie) 

T. (T.)frazierensis (Anderson) 

T. (T.) packardi (Anderson) 

T. (Neocylittdrites) cumminsi Stanton 

T. (N.) punctatus Maldonado-Koerdell 

T. (N.) nelsonin. sp. 

T. (N.) sp. A. 

T. (N.)sp. B. 

T. (Mexicotrachactaeon) coniformis 

(Rose) 

T. (A/.) burckhardti (Buse) 

T. (M.) wrighti n. sp. 

T. (M.) palmer/ n. sp. 

T. (M.) teaensis n. sp. 

T. (M.) granthamensls n. sp. 

T. (M.) rossi n. sp. 

T. (M .) melonensis n. sp.

Actaeonella pecosensis Stanton 

A. delgadoi Choffatt 

A. cf. A. sublaevis Briart and Cornet 

A. broivni n. sp. 

A. borneensis Nutall and Leong 

A. oviformis Gabb 

A. duckettsensis n. sp. 

A. jicarensis n. sp. 

A. coquiensis n. sp. 

A. cf. A. coquiensis n. sp. 

A. marchmontensis n. sp. 

A. briggsi n. sp. 

A. robin son i n. sp. 

A. n. sp. 

A. cubensis n. sp.

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FIGURE 19. Stratigraphic ranges of the Western Hemisphere actaeonellid gastropod species. Numbers above stratigraphic range line represent 

as follows: 1, Jamaica; 2, Puerto Rico; 3, Cuba; 4, Brazil; 5, Guatemala; 6, Mexico; 7, Texas; 8, New Mexico; 9, California.
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FIGURE 20. Index map showing collection localities in Puerto Rico. Numbers on this map refer to those given in the far left-hand column of 

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FIGURE 21. Index map showing collection localities and occurrence of Cretaceous rocks in Jamaica. Numbers on map refer to those given 

in the far left-hand column of the locality register. Letters refer to the following Cretaceous inlier names: A, Green Island; B, Jerusalem Mountain; C, Sunderland; D, Marchmont; E. Maldon; F, Central; G, St. Ann's Great River.

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STRATIGRAPHIC SUMMARY OF WESTERN HEMISPHERE ACTAEONELLID OCCURRENCE 31

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STRATIGRAPHIC SUMMARY OF WESTERN HEMISPHERE ACTAEONELLID OCCURRENCE 33

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NICARAGUA

FIGURE 24. Index map showing collection localities in Mexico and Guatemala.
34 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

PACIFIC \» Santa Ana 

OCEAN

FIGURE 25. Index map showing collection localities in California.

In the Western Hemisphere, post-Turonian occurrences of T. (Trochactaeon) are restricted to Puerto 

Rico, where the earliest occurrence is in the basal part 

of the Rio Bauta Member of the Pozas Formation of 

north-central Puerto Rico (fig. 20, locality 25). Here, 

specimens of Trochactaeon aff. T. (T.) woodsi (Rennie) 

occur in a calcareous tuffaceous sandstone some 3 m 

below a bed containing an abundance of Barrettia 

coatsi (Chubb). This hippuritid rudist bivalve in 

Jamaica is associated with inoceramids of latest Santonian to earliest Campanian Age (Kauffman, 1966).

The Puerto Rico occurrences of T. (Trochactaeon) 

woodsi (Rennie) represent the last appearance of the 

typical subgenus anywhere in the world. The species is 

especially common in upper Campanian rocks of the 

southwestern part of the island in the Cotui Limestone 

(fig. 17). For example, at locality 39 near Palmarejo 

(fig. 20), T. (T.) woodsi occurs in a zone of concentration 

about 0.3 m thick. Underlying units are dominated by 

Toucasia-rich limestone and nerineid-dominated shale 

and limestone that contain a few actaeonellids. The

overlying massive biomicritic Cotui Limestone contains numerous, but scattered, large specimens of T. 

(T.) woodsi together with large nerineid gastropods. A 

similar successional alternation of nerineid-dominated 

shale overlain by T. (T.) woodsi-bearing limestone is 

found at locality 40. The species is found in similar 

limestone in central Puerto Rico in the Botijas Limestone Member and Reves Member of the Pozas Formation (fig. 17) immediately below shale and limestone 

bearing Barrettia species C and D of middle Maastrichtian Age (fig. 20, localities 5-9).

Species of the Late Cretaceous Trochactaeon (Neocylindrites) are restricted to Campanian and Maastrichtian 

rocks in Mexico and Puerto Rico. T. (Neocylindrites) 

nelsoni occurs as small specimens in a tuffaceous shale 

in the uppermost part of the Rio Bauta Member of the 

Pozas Formation (fig. 17), in the Orocovis area of central Puerto Rico (fig. 20, locality 28), in beds of early 

Campanian Age above Barrettia coatsi, and immediately below limestones bearing Barrettia rusae?. Nearby, 

in the Florida Quadrangle (localities 22-24), but in 

rubbly limestones associated with abundant specimens 

of the rudist bivalves Barrettia monilifera and Durania 

nicholasi, larger specimens of T. (N.) nelsoni occur 

sporadically. This association appears to be of late 

Campanian Age. Trochactaeon (Neocylindrites) punctatus (Maldonado-Koerdell) although of very local 

occurrence in the Maastrichtian of Chiapas, Mexico 

(fig. 24, locality 148), marks the youngest occurrence of 

the subgenus in the Western Hemisphere.

Mexicotrochactaeon. Of all subgenera of Trochactaeon present in the Cretaceous of the Western Hemisphere T. (Mexicotrochactaeon) has the greatest diversification of species and widest geographic distribution. 

The oldest record is that of T. (M.) burckhardti (Bb'se) 

from Coniacian rocks (Imlay, 1944, p. 1102) in the 

State of Guerrero, Mexico (Bbse, 1923), and in Brazil 

(Maury, 1930).

No Santonian or Campanian records are known, but 

Maastrichtian species are numerous and spread 

throughout the Caribbean and middle America (fig. 9). 

Three of these species show narrow geographic spread. 

T. (M.) melonensis is restricted to the Melones Limestone of southwestern Puerto Rico in the range zones of 

Barrettia sp. C and sp. D of Dommelen (1971). T. (M.) 

coniformis is known only from the Cardenas Formation 

of San Luis Potosi, Mexico; it is exceptionally variable 

in form and abundant in the shales of this unit (Myers, 

1968). Associated mollusks such as Arctostrea aquilerai 

and Sphenodiscus sp. suggest at least a middle 

Maastrichtian Age, but T. (M.) coniformis may range 

into beds of late Maastrichtian Age. Trochactaeon 

(Mexicotrochactaeon) granthamensis Sohl and Kollmann 

is endemic to Jamaica (fig. 21, localities 79,106), occur-
LOCALITY REGISTER 35

ring in calcareous shale interbedded between rudistrich limestones. In both the Guinea Corn Formation of 

the Central inlier and in the Marchmont inlier it 

occurs within the lower part of the range zone of 

Titanosarcolites that equates with the middle 

Maastrichtian.

The three remaining species of Trochactaeon (Mexicotrochactaeon) have a wider geographic range. The first, 

T. (M.) wrighti, is restricted to the Antillean Islands of 

Jamaica, Puerto Rico, and Cuba, but T. (M.) teaensis 

and T. (M.) rossi range from Mexico through the Antilles. All three species range through most of the middle and perhaps lower part of the upper Maastrichtian.

Actaeonella. Cenomanian through Turonian representatives of Actaeonella are rare in the Western 

Hemisphere and are unknown in rocks of Coniacian 

and Santonian Age. In contrast, they are locally abundant and diverse in rocks of Campanian and Maastrichtian Age. In general, this pattern parallels that of 

Trochactaeon; however, individual species are slightly 

longer ranging and few are as widely distributed.

Cenomanian distribution is restricted to questionable range extensions of two species known to occur in 

the Albian of Texas and Guatemala (fig. 19) and to an 

unnamed Actaeonella (pi. 22, fig. 7) that occurs with 

late Cenomanian mollusks in erratic boulders on 

Martha's Vineyard in Massachusetts. Actaeonella 

oviformis Gabb demonstrates the presence of the genus 

in California, but its precise stratigraphic range is 

unknown. Although Coniacian and Santonian occurrences are unknown at present, Actaeonella may well 

be found in rocks of these ages in geologically unexplored parts of Middle America. Only one species currently unnamed is present in lower Campanian rocks; 

it occurs in shales of the Rio Bauta Member of the 

Pozas Formation in Puerto Rico. Late Campanian 

through Maastrichtian time saw a great increase in 

Actaeonella species diversity. Of special interest among 

these species is A. borneensis, whose distribution, 

which includes occurrences in the limestones of the 

Rancho Nuevo beds of Chiapas, Mexico, and in central 

Cuba in addition to its type locality in Borneo, stands 

in sharp contrast to the endemic Caribbean-Middle 

America distribution of the other coeval species. This 

affinity with Indo-Pacific faunas parallels that pointed 

out by Kollmann and Sohl (1980, p. A5) for Western 

Hemisphere Vernedia gastropods.

Three other species of Actaeonella A. cubensis, A. 

robinsoni, and A. jicarensis are of special note. They 

are especially distinguished by possessing two columellar plaits, which are best shown on plates 22 and 23. 

The first two are rare and all are limited in distribution. Actaeonella cubensis is known only in calcareous 

shales of probable Maastrichtian Age from Oriente

Province, Cuba. The second species, A. robinsoni, is 

known from middle Maastrichtian rocks in the Guinea 

Corn Formation of Jamaica and the Cardenas Formation of Mexico. A. jicarensis is known from but a single 

locality; it occurs in exceptional abundance in limestones of the middle Maastrichtian El Rayo Formation 

of Puerto Rico, cooccurring with abundant nerineids.

A. duckettsensis, A. marchmontensis, and A. briggsi 

are endemic to single Antillean Islands. The first is the 

longest ranging (Campanian to middle Maastrichtian) 

and, although restricted to Jamaica, is found widely 

there. Actaeonella marchmontensis is the most common species in the Titanosarcolites-bearing limestone 

and shale of Jamaica. It occurs in both the Maldon 

Limestone and Woodland Shale of the Maldon inlier, in 

unnamed shales of the Marchmont inlier, and in calcareous shales of the Guinea Corn Formation of the 

Central inlier. Actaeonella briggsi is restricted to the 

Maastrichtian Reves Member of the Pozas Formation 

and to the El Rayo Formation, both of central Puerto 

Rico, and to the El Rayo Formation and Cotui Limestone of the southwestern parts of the island.

The remaining named species, A. coquiensis, ranges 

from upper Campanian to middle Maastrichtian and is 

known from the Coamo Formation of south-central 

Puerto Rico and from the Maldon Limestone and 

Stapleton Formations in Jamaica.

Many specimens of undetermined specific affinities 

are widespread in Campanian and Maastrichtian rocks 

of the Caribbean, Mexico, and Central America. 

Although poorly preserved, they do serve as a record 

for occurrence in such units as the Inoceramus Shales 

of the St. Ann's Great River inlier of Jamaica and the 

Maravillas Formation of Puerto Rico.

LOCALITY REGISTER

The number in the far left-hand column of the locality 

register (1-170) is that cited under the section labelled 

"occurrence" in each species description; this number 

is also placed in the appropriate geographic position on 

the locality index maps (figs. 20-25). The locality number is followed in sequence by the USGS Mesozoic (or 

other identified institution) collection number, the geographic locality, the stratigraphic assignment, and the 

collectors and date on which the collection was made.

Locality No. 

(fig. 201

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collectors),

year of collection

1 29368 -

PUERTO RICO

Barranquitas Quadrangle 

 Reves Member of the Pozas Formation. From 0.5-m-thick bed of gray
36 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Locality No. 

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collectors),

year of collection

2 29367

3 29366 -__ 

4 29365

5 29364, 30388 

6 29861 

7 30506 

 129382

tuffaceous limestone 4 m above the 

base of an 8-m thick, dominantly 

greenish-gray to brownish-red tuffaceous shale unit. Roadcuts of 

Puerto Rico Route 772 on southfacing valley wall of Rio Canabon, 

1.85 km east-southeast of road intersection in Botijas, Barranquitas 

Quadrangle, Barrio Canabon, 

Municipio de Barranquitas. Puerto 

Rico meter grid 44.500N x 

160.150E. N. F. Sohl and W. O. 

Ross, 1965.

Reves Member of the Pozas Formation. From gray-brownish calcareous, tuffaceous, cobbly siltstone at 

top of a 7-m-thick, gray to brownish, tuffaceous shale, about 10 m 

above the preceding collection 

(29368) at the same locality.

Reves Member of the Pozas Formation. From 0.5-m-thick bed of finegrained cobbly limestone immediately overlying the preceding collection (29367) at the same locality.

Reves Member of Pozas Formation. 

From 1.5-m-thick bed of reddishbrown shale overlying collection 

29366; locality is the same as that 

of collection 29368 (loc. 1).

Reves Member of Pozas Formation. 

From 0.25-m-thick bed of rudistbearing cobbly limestone immediately overlying shale of preceding 

collection: locality same as for 

29368 (loc. 1). N. F. Sohl and W. O. 

Ross 1965; N. F. Sohl and W. O. 

Ross, 1973.

Reves Member of Pozas Formation. 

Limestone float on hill slopes above 

roadcuts of Puerto Rico 772 described under USGS colln. 29368 

(loc. 1). N. F. Sohl and W. O. Ross, 

1973.

Reves Member of Pozas Formation. 

Cuts of Puerto Rico Route 772, 

1.85 km east-southeast of road 

junction in Botijas, Barrio Botijas, 

Municipio de Barranquitas. Puerto 

Rico meter grid 44,725N x 

161.000E. N. F. Sohl, H. Kollmann, and W. O. Ross, 1974.

Botijas Member of Pozas Formation. 

Massive bioclastic limestone exposed on hill slopes immediately 

above Rio Canabon at meander 

2.25 km east-southeast of the road 

junction in Botijas, Barrio Canabon, 

Municipio de Barranquitas. Puerto 

Rico meter grid 44,290N x

Locality No. 

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collectors),

year of collection

9 29381 

10 29394 

11 26802,31489 

12 29862, 31363 -

13 30565

14 30566 

161.310E. N. F. Sohl and W. O. 

Ross, 1965.

Botijas Limestone Member of Pozas 

Formation. Massive, whitish, bioclastic limestone exposed on hill 

slopes north of Rio Canabon 2.74 

km east-southeast of road junction 

in Botijas, Barrio Canabon, Municipio de Barranquitas. Puerto Rico 

meter grid 44,100N x 161,650E. 

N. F. Sohl and W. O. Ross, 1965.

Botijas Limestone Member of Pozas 

Formation. Roadcuts of Puerto 

Rico Route 568 between Orovcovis 

and Botijas about 50 m from the 

small settlement at a sharp road 

bend 0.56 km (airline) a little 

north of immediately west of Botijas, Barrio Botijas, Municipio de 

Orocovis. Puerto Rico meter grid 

45.450N x 158,880E. N. F. Sohl 

and W. O. Ross, 1965.

Botijas Limestone Member of the 

Pozas Formation. Limestone exposed in roadcusts of Puerto Rico 

Route 568 between Orocovis and 

Botijas at a small settlement on 

sharp bend of road 0.56 km (airline) just north of west from Botijas, Barrio Botijas, Municipio de 

Orocovis. Puerto Rico meter grid 

45,500N x 158.880E. N. F. Sohl 

and P. A. Gelabert, 1957; N. F. 

Sohl and W. O. Ross, 1965.

Barrancas Limestone Member of the 

Magueyes Formation. From shaly 

interbeds below main dark-gray to 

blackish limestone exposed in 

quarry near Barrancas, 5.06 km 

northeast of Barranquitas, Barrio 

Barrancas, Municipio de Barranquitas. Puerto Rico meter grid 

44.200N x 164,100E. N. F. Sohl, 

1958.

Cabo Rojo Quadrangle

Melones Limestone. Lower part of 

15-m-high bulldozer scraping on 

north-northwest-facing slope of 

arcuately east-trending ridge 1.53 

km northeast of Corozol, Barrio 

Boqueron, Municipio de Cabo Rojo. 

Puerto Rico meter grid 18.125N x 

73,675E. N. F. Sohl and W. O. 

Ross, 1974.

Melones Limestone. Upper Part of 

the 15-m section exposed at locality 13 above. Puerto Rico meter 

grid 18.125N x 73,675E. N. F. 

Sohl and W. O. Ross, 1974.
LOCALITY REGISTER 37

Locality No. 

(fig. 20)

USGS Mesozoic 

colln. No. 

(unless otherwise 

identified)

Geographic locality, stratigraphic assignment, 

description of locality, collectors), 

year of collection

15 26822

16 26393, 29090

30358

17 30359

18 30541 

19 26834 -

Cayey Quadrangle

Rio Mato'n Limestone Member of the 

Robles Formation. Dark-gray to 

blackish bioclastic limestone exposed on hill slopes about 30 m 

above and 61m east of Puerto 

Rico Route 1, 2.74 km (airline) 

west-southwest of Cayey Plaza, 

Barrio Pedro Avila, Municipio de 

Cayey. Puerto Rico meter grid 

29,920N x 178,300E. N. F. Sohl 

and P. A. Gelabert, 1957.

Central Aguirre Quadrangle 

Coamo Formation. Shale exposed in 

gullies above abandoned quarry 

about 10 to 14 m above road level 

at southeastern end of elliptical 

hill 0.88 km north-northwest of 

Puerto Rico 3 and the CoquiCentral Aguirre road, Barrio 

Aguirre, Municipio de Salinas. 

Puerto Rico meter grid 16,075N x 

173,375E. H. L. Berryhill, 1956; 

N. F. Sohl, P. A. Gelabert, and 

H. L. Berryhill, 1957; N. F. Sohl 

and E. G. Kauffman, 1964; N. F. 

Sohl and W. O. Ross, 1973. 

Coamo Formation. Bulldozer track 

taversing lower part of hill immediately above the intersection of 

the road circumscribing the hill 

and the canefield road leading 

south to Route 3, Barrio Aguirre, 

Municipio de Salinas. Puerto Rico 

meter grid 16,480N x 173,875E. 

N. F. Sohl and W. O. Ross, 1973. 

Coamo Formation. Shales exposed 

in bulldozer track along crest of 

hill 1.45 km northwest of intersection of Coqui-Central Aguirre road 

and Puerto Rico Route 3, near the 

76-m summit. Barrio Aguirre, 

Municipio de Salinas. Puerto Rico 

meter grid 16.410N x 173,100E. 

N. F. Sohl, H. A. Kollmann and 

W. O. Ross, 1974.

Coamo Quadrangle 

Maravillas Formation. Exposures of 

conglomerate in upper 106.75 m of 

formation. Roadcut of Puerto Rico 

Route 14, about 1.609 km a little 

north of east of Coamo and immediately south of Rio Cayon, Barrio 

Palmarejo, Municipio de Coamo. 

Puerto Rico meter grid 27.900N x 

162,OOOE. N. F. Sohl and P. A. 

Gelabert, 1957.

Locality No. 

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collectors),

year of collection

20 26841

21 26845 

22 28629

23 28742

24 29372

25 28830, 29099 

26 29383,30532-

Maravillas Formation. From cobbly, 

shaly limestone exposed in lower 

27.4 m of hill slopes 1.01 km (airline) west-northwest of Sabana 

Hoyos and about 2.57 km westsouthwest of the road junction at 

Las Flores, Barrio San Ildefonso, 

Municipio de Coamo. Puerto Rico 

meter grid 23,260N x 162.260E. 

N. F. Sohl and P. A. Gelabert, 

1957.

Maravillas Formation. Cobbly Limestone (about 800 m below top of 

formation) exposed at base of hill 

0.27 km a little north of east of 

preceding (26841) locality. Puerto 

Rico meter grid 23,300 x 161,960E. 

L. Glover and N. F. Sohl, 1957.

Florida Quadrangle 

Flor de Alba Limestone Lentil of 

Pozas Formation. Roadcut near 

top of hill 0.48 km east of Hacienda Penonales, Barrio Pesas, 

Municipio de Ciales. Puerto Rico 

meter grid 51.500N x 144.400E. 

N. F. Sohl and Arthur Nelson, 1962. 

Flor de Alba Limestone Lentil of 

Pozas Formation. Exposure along 

secondary road 0.77 km west of 

Hacienda Flor de Alba, Barrio 

Cialitos, Municipio de Ciales. 

Puerto Rico meter grid 28,742N x 

143,025E. N. F. Sohl and Arthur 

Nelson, 1962.

Flor de Alba Limestone Lentil of 

Pozas Formation. Limestones exposed in roadcuts and hill slope, 

north of sinkholes west of Hacienda Flor de Alba, Barrio Cialitos, 

Municipio de Ciales. Puerto Rico 

meter grid 50,300 to 50,500N x 

142,900 to 143,300E. N. F. Sohl 

and W. O. Ross, 1965.

Orocovis Quadrangle 

Rio Bauta Member of the Pozas Formation (near base of member). 

Roadcuts of Puerto Rico Route 

157, 1.29 km (airline) from crossing of the Rio Bauta, exposures of 

a calcareous, tuffaceous sand below a sequence of welded tuff. Barrio Damian Abajo, Municipio de 

Orocovis. N. F. Sohl, 1963; N. F. 

Sohl and E. G. Kauffman, 1965. 

Rio Bauta Member of Pozas Formation. Collections from float in 

scraped areas along Puerto Rico 

Route 157 derived from the same
38 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Locality No. 

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collectors),

year of collection

beds as 28830 at the same location. N. F. Sohl and W. O. Ross, 

1965; N. F. Sohl, H. A. Kollmann, 

and W. O. Ross, 1973.

27 28831 Rio Bauta Member of the Pozas Formation, (near base of member). 

Roadcuts of Puerto Rico Route 

157, 1.29 km (airline) from crossing of the Rio Bauta, 0.5-m-thick 

tuffaceous limestone rubble bed 

containing Barrettia, 3 m above 

the Trochactaeon-bearing tuffaceous sand bed of 28830. Barrio Damian Abajo, Municipio de Orocovis. 

N. F. Sohl, 1963.

28 30534 Rio Bauta Member (upper part) of the

Pozas Formation. Collection from 

brown, tuffaceous, silty shale exposed in lower part of a bulldozed 

track on the east-facing slope of a 

north-northeast-trending ridge 

overlooking the Rio Bauta and 

just north of bridge of Puerto Rico 

Route 158. Puerto Rico meter grid 

44.900N x 150,300E. N. F. Sohl, 

H. A. Kollmann, and W. O. Ross, 

1974.

29 28617 Sabana Hoyos Limestone Member of

the Maravillas Formation. Exposures in roadcuts on Puerto Rico 

Route 155 (at bench mark 595.3) 

4.75 km (airline) south-southeast 

of junction with Rico Route 143, 

Barrio Pedro Garcia, Municipio de 

Coamo. Puerto Rico meter grid 

32.725N x 157.250E. N. F. Sohl 

and R. P. Briggs, 1962.

30 28618 Sabana Hoyos Limestone Member of

the Maravillas Formation. Roadcuts of Puerto Rico Route 151 

about 300 m south of Ortiga, Barrio Hato Puerco Arriba, Municipio 

de Villalba. Puerto Rico meter 

grid 36.700N x 150,700E. N. F. 

Sohl and R. P. Briggs, 1962.

Puerto Real Quadrangle 

- Melones Limestone. West-facing 

slopes of low hill just north of secondary road, 3.54 km (airline) 

south-southwest of Las Arenas, 

Barrio Boqueron, Municipio de 

Cabo Rojo. Puerto Rico meter grid 

18,525N x 74,750E. N. F. Sohl 

and R. P. Briggs, 1957; N. F. Sohl, 

1963; N. F. Sohl, H. A. Kollmann 

and W. O. Ross, 1974.

32 30537 Melones Limestone. Crest and slopes

of 40-m-high hill, 3.78 km (airline) southwest of Las Arenas,

31 27014,29087 

30535

Locality No. 

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collectors),

year of collection

33 29079,29354 

34 27611

35 27612

36 2866428748 

29075, 29088

37 30544 

38 30510 

Barrio Boqueron, Municipio de 

Cabo Rojo. Puerto Rico meter grid 

18,700N x 73,275E. N. F. Sohl, 

H. A. Kollmann, and W. O. Ross, 

1974.

Melones Limestone. Low exposures 

along cattle run (abandoned farm 

road) 3.22 km south-southeast of 

Las Arenas, Barrio Llanos Costa. 

Puerto Rico meter grid 18,580N x 

74.875E. N. F. Sohl, 1963; N. F. 

Sohl and W. O. Ross, 1966.

Rio Descalabrado Quadrangle

Maravillas Formation (upper part). 

Exposures near secondary (field) 

road, 2.17 km (airline) northnorthwest of Dscalabrado, Barrio 

Rio Canas Abajo, Municipio de 

Juana Diaz. Puerto Rico meter 

grid 20.450N x 151,480E. Peter 

Mattson, 1959.

Maravillas Formation (upper part). 

Exposures of limestone low on hill 

slopes, 2.33 km (airline) northnorthwest of Descalabrado, Barrio 

Rio Canas Abajo, Municipio de 

Juana Diaz. Puerto Rico meter 

grid 20.650N x 151.640E. Peter 

Mattson, 1959.

Sabana Grande Quadrangle

Limestone within El Rayo Formation. Saddle and gullies below 

crest of 125-m-high hill east of 

Quebrada Jicara, 4.75 km southwest of Sabana Grande, Barrio 

Lajas Arriba, Municipio de Lajas. 

Puerto Rico meter grid 23,900N x 

93,400 to 93,600E. N. F. Sohl and 

Peter Mattson, 1963; N. F. Sohl, 

1963; N. F. Sohl and E. G. Kauffman, 1964.

Limestone within El Rayo Formation. Near crest of 147-m-high 

hill 0.88 km north-northwest of 

Las Guaras School and west of 

Puerto Rico Route 328, Barrio 

Rayo, Municipio de Sabana Grande. 

Puerto Rico meter grid 25.825N x 

94.725E. N. F. Sohl, H. A. Kollmann, and W. O. Ross, 1974.

San German Quadrangle 

Basal part of Cotui Limestone. From 

a cobbly but nerineid-rich shaly 

limestone exposed on hill crest 

and southeast-facing slope to 

southeast of the Palmarejo to 

Lajas road (Puerto Rico Route 101) 

1.09 km northeast of school at Pal-
LOCALITY REGISTER 39

Locality No. 

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collector^),

year of collection

39 30512

40 30347

41 30350 

42 31488

43 30343 _-

44 30315

45 30514

marejo, Barrio Sabana Yeguas, 

Municipio de Lajas. Puerto Rico 

meter grid 23,800N x 85,025E. 

N. F. Sohl, H. A. Kollmann, and 

W. O. Ross, 1974.

Basal part of Cotui Limestone. From 

a 0.3-m-thick bed of brown silty 

shale containing a concentration 

of Trochactaeon woodsi (Rennie), 

2.4 m above the limestone of 

30510 (preceding locality). N. F. 

Sohl, H. A. Kollmann, and W. O. 

Ross, 1974.

Cotui Limestone. Collection from 

float near base of small limestone 

knob rising through the alluvium 

of the Lajas Valley and amidst a 

cluster of houses along a secondary road about 300 m east of junction with Puerto Rico Route 116, 

about 1.61 km (airline) southsoutheast of Lajas, Barrio Sabana 

Yeguas, Municipio de Lajas. Puerto 

Rico meter grid 22.790N x 

87.140E. N. F. Sohl and W. O. 

Ross, 1973.

Cotui Limestone. Locality same as 

preceding (30347), but from massive gray bioclastic limestone near 

the top of the hillock. N. F. Sohl 

and W. O. Ross, 1973.

Limestone cobbles in tuffs of the El 

Rayo Formation, 5.63 km southeast of San German, Barrio Lajas 

Arriba, Municipio de Lajas. Puerto 

Rico meter grid 22,560N x 

89,700E. Peter Mattson, 1955.

Erratic block of silicified limestone in 

brecia of the Sabana Grande Formation. Exposures along foot trail 

about 2.73 km northeast of El 

Brujo, Barrio Tuna, Municipio de 

San German. Puerto Rico meter 

grid 26,300N x 82,025E. Peter 

Mattson, 1955.

Lower part of Cotui Limestone. Roadcuts at end of paved road 2.22 km 

north of Llanos intersection, Barrio Paris, Municipio de Lajas. 

Puerto Rico meter grid 24,225N x 

81.195E. N. F. Sohl and W. O. 

Ross, 1974.

Lower part of Cotui Limestone. Hill 

slopes north of Puerto Rico Route 

312, 2.22 km north of El Brujo, 

Barrio Monte Grande, Municipio 

de Cabo Rojo. Puerto Rico meter 

grid 26,480N x 80,OOOE. N. F. 

Sohl and W. O. Ross, 1974.

Locality No. 

(fig. 20)

USGS Mesozoic

col In. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collector!s),

year of collection

46 30516

47 30518

48 30354

49 27090

49a 31488 -

50 30364 

51 30367,30526 

Lower part of the Cotui Limestone. 

Limestone exposures in field adjacent to road and hill slopes opposite intersection with dirt track to 

west, 1.89 km north-northeast of 

intersection of Puerto Rico Routes 

306 and 4 at Llanos, Barrio Paris, 

Municipio de Lajas. Puerto Rico 

meter grid 23,925N x 81,050E. 

N. F. Sohl and W. O. Ross, 1974.

Top of Cotui Limestone. Path and 

field adjacent to jeep trail, 2.57 

km south-southeast of road intersection in Monte Grande, Barrio 

Tuna, Municipio de San German. 

Puerto Rico meter grid 26,150N x 

81,980E. N. F. Sohl, R. P. Volckmann, and W. O. Ross, 1974.

Limestone within the El Rayo Formation. Northeast-facing slopes of 

46-m-high hill in canefields east of 

farm road, 1.20 km west-southwest 

of road junction in Lajas Arriba, 

Barrio Lajas Arriba, Municipio de 

Lajas. Puerto Rico meter grid 

22/725N x 90,050E. N. F. Sohl 

and W. O. Ross, 1973.

Limestone within El Rayo Formation. Slopes and quarry at base of 

triangular east-trending elongate 

hill (altitude 25-30 m), about 0.24 

km north of Parada de Senal Irizarry, Barrio Lajas Arriba Municipio de Lajas. N. F. Sohl and R. P. 

Briggs, 1956.

El Rayo Formation. From limestone 

cobbles in tuff about 5.63 km 

southeast of San German. Puerto 

Rico meter grid 22,560N x 

89,700E. Peter Mattson, 1954.

Limestone within the El Rayo Formation. Collection from a 0.3-mthick, cobbly, shaly limestone 17.4 

m above base of section exposed in 

a quarry about 300 m north of 

road that follows Antigua via del 

Ferrocarril north of Parada de 

Senal Irizarry and 1.5 km airline 

west-southwest of road junction at 

Lajas Arriba. Puerto Rico meter 

grid 22,550N x 89J50E. N. F. 

Sohl and W. O. Ross, 1973.

Locality same as preceding but from 

a thin shale interbedded with 

limestone 49.5 m above base of 

section exposed in quarry. N. F. 

Sohl and W. O. Ross, 1973 and 

1974.
40 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Locality No.

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified I

Geographic locality, stratigraphic assif

description of locality, collector^

year of collection

jnment,

i),

51a 30368

52 29866

53 30032

54 29451

55 29452,29454 

56 30039

57 30040

58 31487

Locality same as for locality 50, but 

from the lower part of a 30-mthick unit of cobbly to massive 

limestone at the top of the section. 

N. F. Sohl and W. O. Ross, 1973.

VIRGIN ISLANDS 

Caledonia Formation. Fragmental 

rudists and other mollusks from a 

conglomerate containing igneous 

and metamorphic pebbles. Buck Island (Buck Island National Monument) about 2.57 km north of Yellow Cliff Bay, St. Croix. John 

Whetten, 1968.

JAMAICA 

Central Inlier

Slippery Rock Formation. Bluffs 

along Slippery Rock River, upstream about 100-150 m from 

bridge of the Frankfield to Mocho 

road and 2.57 km airline south of 

church at Nine Turns. A. G. 

Coates, W. O. Ross, and N. F. 

Sohl, 1971.

Guinea Corn Formation. Thin bed of 

rudist limestone exposed in a low, 

1-m-thick, cut on a farm road 3.6 

km a little north of west of Chapelton. 432,OOON x 447,OOOE. A. G. 

Coates, J. E. Hazel, E. G. Kauffman, and N. F. Sohl, 1966.

Guinea Corn Formation. Fields on 

hill slope above the roadcuts at 

preceding locality (29451) but 

about 10 M higher in the section. 

J. E. Hazel, E. G. Kauffman, 

N. F. Sohl, and A. G. Coats, 1966.

Guinea Corn Formation. Logie 

Green Section; exposure in cuts of 

the Spaldings to Frankfield road 

at the 10.4-mile post. 2.09 km 

west northwest of Grantham. N. F. 

Sohl, W. O. Ross, and A. G. Coates, 

1971.

Guinea Corn Formation. Logie 

Green section; exposures in cuts of 

the Spaldings to Frankfield road 

about 50 to 75 m east of locality 

30039. N. F. Sohl and W. O. 

Ross, 1971.

Guinea Corn Formation. Exposures 

along the Rio Minho from a position 100 m downstream from 

Coates measured section in lower 

part of formation, near Grantham. 

A. G. Coates, 1964.

Locality No. 

(fig. 20)

USGS Mesozoic 

colln. No. 

(unless otherwise 

identified)

Geographic locality, stratigraphic assignment, 

description of locality, collectors), 

year of collection

59 29961

60 30042

60a 30043

61 29972

62 29973

63 29976

64 29978, 30044 

65 29979 - 

66 29980

67 29981

Guinea Corn Formation (basal beds). 

Bluffs on right side of Rio Minho 

immediately above the mouth of 

the White River to about 12 m 

above the mouth, 2.25 km N. 

54 °W. of the bridge at Frankfield, 

Clarendon Parish. A. G. Coates, 

J. E. Hazel, N. F. Sohl, and E. G. 

Kauffman, 1966.

Guinea Corn Formation. Locality 

same as preceding, but from a 

blue-gray shale 2 m above. N. F. 

Sohl and W. O. Ross, 1971.

Guinea Corn Formation. Locality 

same as preceding but from base 

of bluffs. N. F. Sohl and E. G. 

Kauffman, 1966.

Guinea Corn Formation. Rio Minho 

section; locality same as 29961 (locality 59), but downstream. Bluegray silty sandstone about 34.1 m 

above the Bullhead Volcanics. 

N. F. Sohl, E. G. Kauffman, A. G. 

Coates, and J. E. Hazel, 1966.

Guinea Corn Formation. Rio Minho 

section; locality same as 29961 (locality 59). Dark blue-gray shale, 

35 m above the top of the Bullhead 

Volcanics. N. F. Sohl, E. G. Kauffman, A. G. Coates, and J. E. 

Hazel, 1966.

Guinea Corn Formation. Rio Minho 

Section; radiolitid rudist-bearing, 

rubbly limestone 40.87 to 42.4 m 

above the Bullhead Volcanics. 

E. G. Kauffman, A. G. Coates, 

J. E. Hazel, and N. F. Sohl, 1966.

Guinea Corn Formation. Rio Minho 

Section; blue-gray, dark shale 48.8 

to 52.5 m above the Bullhead Volcanics. E. G. Kauffman, A. G. 

Coates, J. E. Hazel, N. F. Sohl, 1966.

Guinea Corn Formation. Rio Minho 

section; blue-gray shale about 52.5 

m above the Bullhead Volcanics. 

N. F. Sohl, A. G. Coates, J. E. 

Hazel, E. G. Kauffman, 1966.

Guinea Corn Formation. Rio Minho 

section; silty cobbly limestone 53.4 

to 55.5 m above the Bullhead Volcanics. A. G. Coates, J. E. Hazel, 

N. F. Sohl, and E. G. Kauffman, 

1966.

Guinea Corn Formation. Rio Minho 

section; brown shale and nodular 

limestone 67 m above top of the 

Bullhead Volcanics. J. E. Hazel, 

N. F. Sohl, E. G. Kauffman, and 

A. G. Coates, 1966.
LOCALITY REGISTER 41

Locality No. 

(fig. 20)

USGS Mesozoic 

colln. No. 

(unless otherwise 

identified)

Geographic locality, stratigraphic assignment, 

description of locality, collectors), 

year of collection

68 29982

68a 29985

69 29987

70 29988

71 29989

72 29990

73 30050

74 30054

75 30058 -

Guinea Corn Formation. Exposure of 

gray silty shale in banks of the 

Rio Minho, 67.3 m above the base 

of the formation at same locality 

(59) as 29961. A. G. Coates, J. E. 

Hazel, N. F. Sohl, and E. G. Kauffman, 1966.

Guinea Corn Formation. Limestone 

containing volcanic pebbles 70.1- 

74.7 m above the base of the formation at same locality (59) as 

29961. N. F. Sohl, J. E. Hazel, 

E. G. Kauffman, and A. G. Coates, 

1966.

Guinea Corn Formation. Massive 

limestone exposed in banks of the 

Rio Minho 76.2 m above the base 

of the formation at same locality 

(59) as 29961. J. E. Hazel, E. G. 

Kauffman, N. F. Sohl, and A. G. 

Coates, 1966.

Guinea Corn Formation. Blue-gray 

shale in banks of Rio Minho, 82.3 m 

above base of formation at same 

locality (59) as 29961. N. F. Sohl, 

E. G. Kauffman, A. G. Coates, and 

J. E. Hazel, 1966.

Guinea Corn Formation. Interbedded 

cobbly limestone and shale exposed in banks of the Rio Minho 

82.9-85.9 m above the base of the 

formation at same locality (59) as 

29961. E. G. Kauffman, J. E. 

Hazel, A. G. Coates, and N. F. 

Sohl, 1966.

Guinea Corn Formation. Chiapasellarich limestone exposed in banks of 

the Rio Minho about 87.8 m above 

base of the formation at same locality (59) as 29961. N. F. Sohl, 1966.

Guinea Corn Formation. From near 

base of 6.3-m-thick light-gray 

limestone 6.7 m below the contact 

with the volcanic rocks of the 

Summerfield Formation exposed in 

banks of the Rio Minho, 2 km N. 

30 °W. of Frankfield Post Office. 

N. F. Sohl and W. O. Ross, 1971.

Guinea Corn Formation. From middle 

part of the 19.8-m unit of interbedded cobbly limestone and massively bedded radiolite limestone, 

59.8 to 79.6 m below the Summerfield Formation at same locality 

(73) as 30050. N. F. Sohl and 

W. O. Ross, 1971.

Guinea Corn Formation. From 0.6-m 

shale about 82.6 m below the 

Summerfield Formation exposed

Locality No. 

(fig. 20)

USGS Mesozoic 

colln. No. 

(unless otherwise 

identified)

Geographic locality, stratigraphic assignment, 

description of locality, collectors), 

year of collection

76 30056

77 30061

78 30141

79 30063

80 18011

81 18014

82 18016

83 18017

in banks of the Rio Minho at same 

locality (73) as 30050. N. F. Sohl 

and W. O. Ross, 1971.

Guinea Corn Formation. From a 

0.9-m shale, 81.4 m below the 

Summerfield Formation exposed 

in banks of the Rio Minho at same 

locality (73) as 30050. N. F. Sohl 

and W. O. Ross, 1971.

Guinea Corn Formation. Exposures 

in streambed and banks of the Rio 

Minho 2.3 km N. 30 °W. of Frankfield Post Office, Clarendon Parish. 

Collection from a 2.4-m-thick 

blue-gray shale, about 76 m above 

the base of formation. N. F. Sohl 

and W. O. Ross, 1971.

Guinea Corn Formation. Locality 

same as that for 30061 doc. 77) 

but from a cobbly radiolite-rich 

shale about 73.3 m above base of 

formation. N. F. Sohl and W. O. 

Ross, 1971.

Guinea Corn Formation. Locality 

same as that for 30061 (loc. 77), 

but from a brownish hackly shale 

about 70.73 m above base of formation. N. F. Sohl and W. O. 

Ross, 1971.

Guinea Corn Formation. Near second 

outcrop of rudist limestone, 16.9 

to 17.3 km from Spaldings. Soft 

yellow calcareous marl full of rudists, together with corals, decapod 

crustacean fragments, mollusks, 

etc. Logie Green road, faulted 

against red shales, Middlesex 

County, Clarendon Parish, Jamaica. C. A. Matley, 1939.

Guinea Corn Formation. Rudist 

marls, very fossiliferous Logie 

Green section, 17.7 to 19.3 km 

from Spaldings, Middlesex County, 

Clarendon Parish, Jamaica. C. R. 

Matley, 1939.

Guinea Corn Formation. Logie Green 

section, continuation of 18014 beds 

containing more rudists, Middlesex County, Clarendon Parish, Jamaica. C. A. Matley, 1939.

Guinea Corn Formation. "Bad Rock," 

nodular rudist beds, slipping over 

solid limestone. Logie Green section, 17.7 to 19.3 km from 

Spaldings, at embanked area at a 

bend above Rio Minho, Middlesex 

County, Clarendon Parish. C. A. 

Matley, 1939.
42 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Locality No. 

(fig. 20)

USGS Mesozoic 

colln. No. 

(unless otherwise 

identified)

Geographic locality, stratigraphic assignment, 

description of locality, collectors), 

year of collection

84 18019 

85 18020

86 18023

87 18024

 18029 -

89 18031

90 18033

91 18034

92 29924, 30028 

Guinea Corn Formation. Horizon 

probably high in rudist limestone. 

Logie Green section, beyond tuffs 

at 12-mile post, Middlesex County, 

Clarendon Parish. C. A. Matley, 

1939.

Guinea Corn Formation. Nodular 

marly clays or marl with rudists 

(slipping). Logie Green Section, 

19.3 to 20.9 km from Spaldings, 

Middlesex County, Clarendon Parish. C. A. Matley, 1939.

Guinea Corn Formation. Nodular 

limestone interbedded with marl, 

and shale, containing small rudists. Logie Green Section on road 

to Spaldings, at Grantham, near 

but before the crossing of the 

White Rock River, at near road 

level, Middlesex County, Clarendon Parish. C. A. Matley, 1939.

Guinea Corn Formation. On path 

rising above the preceding locality 

and at a higher stratigraphic 

level, Logie Green section, Middlesex County, Clarendon Parish. 

C. A. Matley, 1939.

Guinea Corn Formation. Top beds of 

rudist limestone. White Rock 

River Road about 300 m before 3- 

mile post, Middlesex County, Clarendon Parish. C. A. Matley, 1939.

Guinea Corn Formation. Gastropod 

bed 7.2 cm thick exposed along the 

Frankfield-Trout Hall Road, at 

28.6 km, Middlesex County, Clarendon Parish, Jamaica. C. A. Matley, 1939.

Guinea Corn Formation. Rudist 

limestone. Content Gully, Frankfield, Middlesex County. C. A. 

Matley, 1939.

Guinea Corn Formation. Shale containing thin calcareous layers below rudist limestone. Above left 

bank of stream, carbonized plant 

remains, small oysters, tiny mollusks, etc. Content Gully, Frankfield, Middlesex County. C. A. 

Matley, 1939.

Marchmont Inlier

"Titanosarcolites limestone." Lower 

1.6 m of section in shaly, cobbly 

limestone exposed at a hairpin 

bend of the Ducketts to Greenwich 

road 0.32 km airline E.20° N. of 

Ducketts Crossroads, Ducketts 

land Settlement, Westmoreland

Locality No. 

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collectors),

year of collection

93 29925, 30027 

94 29926

95 29927 

96 29921 

97 30026

98 30030

99 30462

Parish, 18°17'47"N., 77°54'54"W. 

N. F. Sohl, E. G. Kauffman, A. G. 

Coates, and J. E. Hazel, 1966; 

W. O. Ross, A. G. Coates, and 

N. F. Sohl, 1971.

"Titanosarcolites limestone." Locality same as preceding but from the 

middle of a 0.7-m unit of yellowish shale overlying the shaly limestone of the preceding. N. F. Sohl, 

E. G. Kauffman, A. G. Coates, and 

J. E. Hazel, 1966; N. F. Sohl, 

W. O. Ross, and A. G. Coates, 1971.

"Titanosarcolites limestone." Locality same as 29924 (loc. 92) but 

from 1.3-m radiolite limestone bed 

overlying preceding locality (29925). 

N. F. Sohl, E. G. Kauffman, A. G. 

Coates, and J. E. Hazel, 1966.

"Titanosarcolites limestone." Float 

in fields on hillslope below 29924 

(loc. 92), 0.32 km W.30°N. of 

Ducketts Crossroads, Ducketts 

Land Settlement, Westmoreland 

Parish, 18°17'49"N., 77°54'53"W. 

E. G. Kauffman, A. G. Coates, 

J. E. Hazel, and N. F. Sohl, 1966.

"Titanosarcolites limestone." From a 

5-m-thick, gray cobbly limestone 

32 m above a brownish shale in a 

section measured from edge of banana field and directly up hill 

slope to road level 0.48 km (airline) northeast of Ducketts Crossroads, 18°17'54"N., 77°54'57"W. 

A. G. Coates, J. E. Hazel, N. F. 

Sohl, and E. G. Kauffman, 1966.

"Titanosarcolites limestone." Reddishbrown shale exposed in cuts of the 

Cambridge to Lambs River road at 

Ducketts Crossroads, 2.17 km (airline) south-southeast of the railroad 

station in Cambridge, 18°17'43"N., 

77°54'46"W. N. F. Sohl, W. O. 

Ross, and A. G. Coates, 1971.

"Titanosarcolites limestone." Exposures of calcareous sandstone on 

north bank of east-flowing tributary of Great River west of Lambs 

River road, 0.97 km south of Ducketts Crossroads, Westmoreland 

Parish, 18°17'12"N., 77°54'43"W. 

W. O. Ross, A. G. Coates, and 

N. F. Sohl, 1971.

"Titanosarolites limestone." St. 

Leonard to Seafordtown road, 0.56 

km northeast (airline) of Seafordtown junction, Westmoreland Parish, 18°14'59"N., 77°53'55"W.
LOCALITY REGISTER 43

Locality No. 

(fig. 20)

USGS Mesozoic 

colln. No. 

(unless otherwise 

identified)

Geographic locality, stratigraphic assignment, 

description of locality, collectors), 

year of collection

100 30467

101 30460

102 30456

102a 30457

103 30471

104 30472 

105 30473

105a 30476 

E. G. Kauffman and N. F. Sohl, 

1972.

"Titanosarcolites limestone." From 

upper part of 6.5-m-thick unit of 

gray shale below a 1.5-m-thick 

radiolite limestone exposed in cuts 

of the St. Leonards to Marchmont 

road at the south edge of Marchmont, or 0.16 km (airline) south of 

the road intersection in Marchmont, 18°15'34"N., 77°52'53"W. 

N. F. Sohl and E. G. Kauffman, 

1972.

Shales below "Titanosarcolites limestone." Exposures of reddish-brown 

shale that overlie brownish sand 

in cuts of Lambs River to Seafordtown road, 1.45 km (airline) northwest of Lambs River, Westmoreland 

Parish, 18°15'02"N., 77°54'30"W. 

E. G. Kauffman and N. F. Sohl, 

1972.

"Titanosarcolites limestone." Exposures in cuts of the Lambs River 

to Marchmont road, 1.5 km (airline) north-northeast of Lambs 

River. Collection from a 3-m-thick 

gray silty shale 6 m above the 

base of the section, 18°16'24"N., 

77°54'46"W. N. F. Sohl and E. G. 

Kauffman, 1971.

Locality same as loc. 102 (30456), 

but from a 1.5-m-thick shale 11 m 

above base of section. N. F. Sohl 

and E. G. Kauffman, 1971.

"Titanosarcolites limestone." Seafordtown to Chesterfield Bridge 

road, 0.93 km (airline) south of 

Seafordtown road junction. Collection from 2-m-thick brownishgray shale, 3 m above base of section, 18°14'18"N., 70°53'52"W. 

E. G. Kauffman and N. F. Sohl, 

1972.

"Titanosarcolites limestone." Locality 

same as for 30471, but from a 1.5- 

m-thick brownish clay containing 

abundant corals, 5 m above base 

of section. N. F. Sohl and E. G. 

Kauffman, 1972.

"Titanosarcolites limestone." Locality 

same as that for 30471 (loc. 103), 

but from a 6-m-thick nodular to 

cobbly gray radiolite-coral-rich limestone that caps the section. E. G. 

Kauffman and N. F. Sohl, 1972.

"Titanosarcolites limestone." Bluff 

along right bank of east-flowing 

tributary of Great River immediLocality No. 

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collectors),

year of collection

106 30455 -

107 30480 

108 30421

109 30011,30422 

HO 29465

111 29466 -

112 29467

ately west of bridge of secondary 

road, about 40 m south of Chesterfield Bridge-Seafordtown road, 

18°13'50"N., 77°52'27"W. N. F. 

Sohl and E. G. Kauffman, 1972.

"Titanosarcolites limestone." Cuts of 

Ducketts to Lambs River road 1.85 

km (airline) south of Ducketts Crossroads, 18°16'32"N., 77°54'42"W. 

N. F. Sohl and E. G. Kauffman, 

1971.

"Titanosarcolites limestone." 

Brownish-gray shale exposed in 

stream bluff about 3 m above 

water level on right bank of tributary of the Great River, immediately west of bridge of secondary 

road, about 40 m south of the 

Chesterfield Bridge to Seafordtown 

road, 18°13'59"N., 77°52'27"W. 

N. F. Sohl and E. G. Kauffman, 

1971.

Jerusalem Mountain Inlier

"Titanosarcolites limestone." Cobbly 

limestone exposed in roadcuts 1.13 

km (airline) southeast of Jerusalem 

Mountain crossroads, 18°19'0"N., 

78°13'32"W. E. G. Kauffman and 

N. F. Sohl, 1971.

"Titanosarcolites limestone." Exposed in roadcuts of road to Mount 

Eagle 1.07 km (airline) southeast 

of Jerusalem Mountain road intersection on path to Assemblage of God 

Church, 18°18'59"N., 78°13'36"W. 

W. O. Ross, A. G. Coates, and 

N. F. Sohl, 1971; N. F. Sohl and 

E. G. Kauffman, 1971.

"Titanosarcolites limestone." Roadcuts along Jerusalem Mountain to 

Belle Isle road, northeast of houses 

that sit on "oyster limestone" outcrops, about 330 m (airline) northeast of intersection with the Blairs 

Hill to Jerusalem Mountain road. 

18°19'29"N., 78°13'14"W. N. F. 

Sohl, E. G. Kauffman, and J. E. 

Hazel, 1966.

"Titanosarcolites limestone." Float 

from covered interval in sections 

in roadcuts at the same locality as 

that for 29465 (loc. 110). E. G. 

Kauffman, J. E. Hazel, and N. F. 

Sohl, 1966.

"Titanosarcolites limestone." From 

a 0.6-m-thick rubbly limestone 

exposed about 3 m above road 

level at same locality as that for
44 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Locality No. 

(fig. 20)

USGS Mesozoic 

colln. No. 

(unless otherwise 

identified)

Geographic locality, stratigraphic assignment, 

description of locality, collectors), 

year of collection

113 29473

114 29475

115 29476

116 29953,30023 

117 30019

118 29951

119 29955,30017

30018

121 30007

29465 (loc. 110). J. E. Hazel, E. G. 

Kauffman, and N. F. Sohl, 1966.

"Titanosarcolites limestone." From 

1-m-thick rubbly limestone about 

100 m east of 29465 (loc. 110), 

about 7 m above base of measured 

section. N. F. Sohl, J. E. Hazel 

and E. G. Kauffman, 1966.

"Titanosarcolites limestone." From 

massive 1-m-thick limestone unit 

about 7.5m above base of measured section, 120 m east of 29465 

(loc. 110). E. G. Kauffman, J. E. 

Hazel, and N. F. Sohl, 1966.

"Titanosarcolites limestone." From 

cobbly limestone in shale overlying 29475, about 8 m above base 

of measured section 120 m east of 

29465 (loc. 110). J. E. Hazel, N. F. 

Sohl, and E. G. Kauffman, 1966.

Maldon Inlier

Woodland Shale. From a yellowishbrown silty clay 3 m above base of 

section and about 3.25 m below 

the Maldon Limestone. Point to 

Flamstead Road, 1.93 km (airline) 

north of the crossroads at Flamstead, St. James Parish. J. E. 

Hazel, N. F. Sohl, E. G. Kauffman, 

and A. G. Coates, 1966.

Vaughansfield Limestone. Collection 

from float weathering out on hill 

slopes downslope (north) from 

Flamstead to Maroontown road, 

St. James Parish, 1.77 km (airline) 

southwest of Maroontown. A. G. 

Coates, W. O. Ross, and N. F. Sohl, 

1971.

Maldon Limestone. Exposures 

titanosarcolites-rich limestone in 

the schoolyard of Maldon schoolyard, 0.21 km (airline) N.74°W. of 

Maroontown, St. James Parish. 

J. E. Hazel, N. F. Sohl, E. G., 

Kauffman, and A. G. Coates, 1966.

Maldon Limestone. Yellowish-brown 

shale 1.1 m above base of measured section in roadcuts and adjacent fields near Chatsworth School 

at Shaw Castle, 1.45 km (airline) 

N. 33 °W. of Maroontown, St. James 

Parish. N. F. Sohl, E. G. Kauffman; 

A. G. Coates, and J. E. Hazel, 

1966; W. O. Ross, A. G. Coates, 

and N. F. Sohl, 1971.

Green Island Inlier 

"Barrettia Limestone." Limestone 

from high on west-facing wall of

Locality No. 

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collectors),

year of collection

122 29939, 30014 

123 30065 

124 18283

125 19464,19465 

126 

127 

128 

129 

27169 

(895P Palmer

Collection)

27266 

(2793 Palmer 

Collection)

27277 - 

(2941 Palmer 

Collection)

27278 

(2942 Palmer 

Collection)

north-trending valley 0.97 km (airline) S.40 °W. of Green Island, 

18°22'49"N, 78°16'49"W. N. F. 

Sohl, A. G. Coates, and W. O. Ross, 

1971.

Sunderland Inlier 

Stapleton Formation. Roadcut exposures south of Stapleton on road to 

Kensington and from a 0.4-m-thick 

bed of coarse bioclastic limestone 

2.4 km (airline) south-southeast of 

Sunderland Bridge, St. James Parish, 18°22'47"N., 77°50'07"W. 

N. F. Sohl, E. G. Kauffman, A. G. 

Coates, and J. E. Hazel, 1966; 

N. F. Sohl, W. O. Ross, and A. G. 

Coates, 1971.

St. Ann's Great River Inlier 

Exposures of dark-gray shale in 

bluffs of Saint Ann's Great River, 

upstream and about 100 m higher 

than the shales exposed at the 

ford south-southeast of St. Ann's 

Bay. N. F. Sohl and W. O. Ross, 

1971.

CUBA

Habana Formation? From exposures 

in a small unnamed creek, 1.0 km 

west of the stack of Central Perseverancia, Santa Clara Province, 

Emilio Aleman, date unknown.

Habana Formation? Exposures in 

bed of river east-southeast of 

house of Senor Aparicio 25.3 km 

east of Central Miranda on south 

bank of Rio Sumidero, Oriente, 

Province. G. E. Lewis, 1945.

Habana Formation. 150 m north of 

Grua Esperanza, which is 6 km 

east of Madruga, Habana Province. R. H. Palmer, 1942.

Habana Formation. 500 m S. 20° E. 

of Negrito and 8.5 km southsoutheast of Central Soledad, Santa Clara Province. R. H. Palmer, 

1942.

Habana Formation. El Abra, 20 km 

east and 6 km south of Central 

Hormiguero, Santa Clara Province. R. H. Palmer, 1942.

Habana Formation. South slope of 

Loma Campana about north of 

Grua Campana, Central Hormiguero, Santa Clara Province. R. H. 

Palmer, 1942.
LOCALITY REGISTER 45

Locality No. 

(fig. 20)

USGS Mesozoic

colln. No.

(unless otherwise

identified)

Geographic locality, stratigraphic assignment,

description of locality, collector!s),

year of collection

130 27295 - 

(Original No. 

3144)

131 9568

132 1632

133 1926 -

134 1941 .

135 1946, 18791 

136 11853

137 H854

138 11883

139 H885

Habana Formation. Copper mine, 6 

km south of Fomento and west of 

Fomento-Sipiabo road, Santa Clara 

Province. Cuba. R. H. Palmer, 1944.

MASSACHUSETTS, U.S.A. 

Raritan Formation. Sideritic erratic 

cobble derived from cliffs of Gay 

Head, Martha's Vineyard. J. B. 

Woodworth, 1889.

TEXAS, U.S.A.

"Caprina Limestone" (Edwards 

Limestone). About 9.65 km northeast of Kerrville on the road to 

Fredricksburg, Kerr County. R. T. 

Hill, T. W. Vaughan, and T. W. 

Stanton, 1895.

Buda Limestone(?). 7.24 km a little 

south of west of Kent, Culberson 

County. T. W. Stanton, 1898.

Devils River Limestone (upper part). 

Near base of box canyon about 

91.5 m below level of the track of 

the G. H. and S. A. Railroad at 

the high bridge over the Pecos 

River, 4.8 km east of Shumla, Val 

Verde County. T. W. Stanton, 1898.

Devils River Limestone (upper part). 

Painted Caves on the Rio Grande 

River 1.6 km below the mouth of 

the Pecos River, Val Verde County. 

T. W. Stanton, 1898; W. F. Cummins, 1898(7).

Washita Group. Top of mesa about 

2.4 km south of Texon Oil Company Santa Rita well, 22.4 km 

west of Big Lake, Reagan County. 

T. W. Stanton and J. A. Udden, 

1923.

Upper limestone of the Washita 

Group. Cut and adjacent ravine on 

the Kansas City and Orient Railway, 4 km southeast of Big Lake, 

Reagan County. T. W. Stanton 

and J. A. Udden, 1923.

Devils River Limestone. About 27.4 m 

below top of formation in cuts of 

the highway on east side of the 

Pecos River bridge, Comstock 

Quadrangle. T. W. Stanton, 1923.

Devils River Limestone (top). 1.5 m 

of limestone in bed of Middle Fork 

of Sycamore Creek, 1.2 km southeast of Havilien Oil Company's 

Weatherbee Well No. 10, about 

33.8 km east of Del Rio (Del Rio 

Quadrangle) Kinney County. T. W. 

Stanton and W. H. Ritchey, 1923.

Locality No. 

(fig. 20)

USGS Mesozoic 

colln. No. 

(unless otherwise 

identified)

Geographic locality, stratigraphic assignment, 

description of locality, collectors), 

year of collection

140 13150 Fredericksburg Group. From top of

9.15-m cliff-forming limestone 

that caps summit of Big Spring 

Mountain mesa, Howard County. 

W. B. Lang and T. W. Stanton, 1925.

141 13196 Washita Group (possibly Denton

Clay equivalent). Top of low mesa 

2.4 km north of west of Leon 

Spring Reservoir, west of Fort 

Stockton, Pecos County. W. B. 

Lang and T. W. Stanton, 1925.

142 14185 Washita Group. Actaeonella bed,

7.6 to 9.1 m below big sandstone 

at top of Trinity Group north of 

highway 49.9 km east of Marathon, Brewster County. T. W. 

Stanton and W. P. Popenoe, 1928.

143 14186 Washita Group. On north side of

highway 49.9 km east of Marathon, 

from a loose fragment of limestone 

on slope beneath sandstone of 

upper part of Trinity Group. Probably float from above the Trinity, 

Brewster County. T. W. Stanton 

and W. P. Popenoe, 1928.

144 14236 Washita Group (Buda Limestone?).

Davis Mountain Station junction 

of Texas and Pacific Railroad and 

Bankhead highway, 1.6 km east of 

Kent and 30 m above the base of 

exposure, Culberson County. Collectors: T. W. Stanton and W. P. 

Popenoe, 1928.

145 "Thorp Springs Limestone Member"

of the Glen Rose Formation. Exposures in roadcuts on IM 1715, 0.96 

km northwest of Morgan Mill, 

Erath County.

146 UT1036TX1-6 Edwards Limestone. Locality unTexas Memorial specified, Williamson County. 

Museum

147 1694

148

NEW MEXICO, U.S.A. 

4 km southwest of Doyles Well, between and in line with the corner 

monument and the highest peak of 

Hatchita Mountain (apparently exposures on the south flank of 

Sierra Rico) in sec. 2, T. 20 S., R. 

14 W., Hidalgo County. E. A. 

Mearns, 1892.

MEXICO

Limestone exposures near Ocuilapa, 

2 leagues from Ocozocuautla and 2 

leagues from Chapopote, Municipio 

de Ocozocuautla de Espinsa, State 

of Chiapas. (Maldonado-Koerdell,
46 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Locality No. 

(fig. 20)

USGS Mesozoic 

colln. No. 

(unless otherwise 

identified)

Geographic locality, stratigraphic assignment, 

description of locality, collectors), 

year of collection

149

150

151 H467 

152 24392 

153 24394 

154 26615

155 28187

156 28218

157 28216 

158 28219 

159 28217

1950, p. 189.) J. T. Singewald, Jr., 

1922.

Rancho Neuvo Beds (upper part). 

Ocuilapa Road at 5 km northwest 

of Ocozocuautla, State of Chiapas. 

L. J. Chubb, 1956.

Rancho Nuevo Beds (upper part). 

Escarpment 1.5 km northwest of 

Ocozocuautla, State of Chiapas. 

L. J. Chubb, 1956.

Aquilapa, north of Ocozocuautla, 

State of Chiapas. Raymond Leibensperger, 1922.

Cardenas Formation. 3.2 km south 

of Rayon, State of San Luis Potosi. 

Kenneth Segerstrom and Carl 

Fries, Jr., 1952.

Cardenas Formation. Locality same 

as that for USGS 24392 (loc. 152); 

collections from a few meters 

higher in the section. Kenneth 

Segerstrom and Carl Fries, Jr., 

1952.

100 m east of point 923 on the second limestone ridge west of Cuautala, Zacatepec district, State of 

Morelos. Carl Fries, Jr., 1955.

Cardenas Formation. Calcareous 

shale 28.8 m thick and 502.5 m 

below the top of Bruce Wade's 

measured section (unit 67 of Imlay, 1944, p. 1137), east and southeast of Cardenas, State of San 

Luis Potosi. Bruce Wade, 1923.

Cardenas Formation. Locality same 

as that for 28187, but from unit 69 

of Imlay's (1944, p. 1137) portrayal 

of Wade's Canon Canada section, 

a 5.5-m-thick unit of shale and 

limestone 482 m below top of section. Bruce Wade, 1923.

Cardenas Formation. Locality same 

as that for 28187, but from unit 80 

of Imlay's (1944, p. 1137) portrayal 

of Wade's Canon Canada section, 

a 60-m-thick unit of brown calcareous sandstone and shale 342.7 m 

below top of section. Bruce Wade, 

1923.

Cardenas Formation. Locality same 

as that for 28187, but from uni 98 

of Imlay's (1944, p. 1137) portrayal 

of Wade's Canon Canada section, 

an 8.8-m-thick red and gray shale 

unit, 60 m below the top of the 

section. Bruce Wade, 1923.

Cardenas Formation. About 2 km 

west of Lima and southwest of 

Valles, State of San Luis Potosi. 

Bruce Wade, 1923.

Locality No. 

(fig. 20)

USGS Mesozoic 

colln. No. 

(unless otherwise 

identified)

Geographic locality, stratigraphic assignment, 

description of locality, collectors), 

year of collection

160 28327

161

162

163

164

165

166

167

168

CAS. 27831, -

28103 

CAS. 10664- 

10666, 27831

CAS. 444,10663 - 

CAS. 33721 -

CAS. 53169 - 

CAS. 444A -

CAS. 733- 

CAS. 1293-

169 2672

170 2179

GUATEMALA

Albian Limestone. Gravity station 

(Sohio Company) No. 972. 2 km 

east of Sojio, Robert H. Ray Geophysical Survey, about 12 km 

south of La Libertad, Peten. Barnum Brown, 1948 and 1950.

CALIFORNIA, U.S.A.

[Collectors and date not recorded] 

Stinking Canyon, 2.4 km north of

Fraziers Corners, Shasta County. 

4.8 km north of Bellavista, 3.2 km

above the Cawkins home on Little

Cow Creek, Shasta County. 

Rocky Gulch near Hornbrook, Siskiyou County. 

South Fork Cow Creek, Shasta

County.

Woodman Creek, Shasta County. 

Near Klamath River 2.4 km south

of Hornbrook, Siskiyou County. 

8 km southwest of Corona, Orange

County. 

Bella Vista District, Shasta County.

ALASKA, U.S.A.

Right bank of the Yukon River, 8 km 

above Nohatatietin, Yukon. A. J. 

Collier, 1902.

From above Nulato and below Pickart's coal mine, on the north or 

right bank of the Yukon River, 

Koyukuk River region. F. C. 

Schrader, 1899.

SYSTEMATIC PALEONTOLOGY

In the following descriptions, morphologic terminology generally conforms with that used in "Treatise on 

Invertebrate Paleontology" (Cox, 1960). In the sections 

on measurements, the following abbreviations are 

used:

H, total height of shell, measured in millimeters parallel to axis of coiling.

MD, maximum diameter of shell, measured in millimeters normal to axis of coiling. 

PA, pleural angle, measured in degrees. 

HB, height of body whorl, measured parallel to axis of 

coiling.

Under the section on types, the following abbreviations are used:

USNM, U.S. National Museum 

USGS, U.S. Geological Survey Mesozoic Locality
SYSTEMATIC PALEONTOLOGY 47

UT, University of Texas

CAS, California Academy of Sciences

The numbers listed for areas cited in sections on 

occurrence refer to detailed locality information provided in the locality register (p. 35).

Superfamily ACTEONIDAE 

Family ACTAEONELLIDAE

Genus TROCHACTAEON Meek, 1863

Type by original designation, Actaeonella renauxiana 

d'Orbigny

Diagnosis. Elongate ovate to subcylindrical, moderately thick walled shells. Breadth of body whorl 

greatest above midheight. Spire gently concave to convex in profile and less than half total shell height. 

Aperture elongate, broadening anteriorly, posteriorly 

sinused; inner lip callused, columella normally bearing 

three prominent folds but rarely two or four; outer lip 

smooth or bearing one low ridgelike plait.

Discussion. A detailed review of the older views on 

the genus has been given by Kollmann (1967). Although 

many subdivisions of the genus have been made by 

Meek (1863) and Hacobjan (1963, 1972), we recognize 

only four valid subgenera, as discussed below: Trochactaeon (Trochactaeon) Meek, 1863; T. (Neocylindrites) 

Sayne, 1932; T. (Sevanella) Hacobjan, 1972; T. (Mexicotrochactaeon) Hacobjan, 1972.

Subgenus Trochactaeon Meek, 1863

Type by original designation Actaeonella renauxiana 

d'Orbigny.

Diagnosis. Proportional breadth of shell high for 

genus (height/width ratio 1.4:1 to 2:1) and thick walled. 

Spire height variable, whorl sides well rounded. 

Growth lines arcuate. Aperture expands anteriorly; 

parietal lip smooth; columella with three or rarely four 

plaits, one as a high palatal ridge.

Discussion. Subjective synonyms include: Spiractaeon Meek, 1863; Mesotrochactaeon Hacobjan, 1972; 

Neotrochactaeon Hacobjan, 1972; Purpuractaeon 

Hacobjan, 1963.

In 1863, Meek erected the genus Spiractaeon for 

high-spired species that we here assign to Trochactaeon (Trochactaeon). Recognition of Spiractaeon is 

untenable because the proportional height of spire 

within the individual species Meek assigned is so variable that it encompasses the full spectrum of form 

shown within Trochactaeon (Trochactaeon) and 

Spiractaeon.

Neotrochactaeon was proposed by Hacobjan (1972) to 

accommodate species whose shells have an especially 

low spire and strongly inflated whorls. The included 

species show a considerable variety of form, so far as 

shell obesity and spire height are concerned. In this

sense, many species would fall within the definition of 

Spiractaeon Meek, here considered a synonym of T. 

(Trochactaeon). Other species, such as Actaeonella 

(Trochactaeon) burckhardti Bbse and A. (T.) quadriplicatus Bbse bear a ridge on the inner surface of the outer 

lip and are more properly considered members of T. 

(Mexicotrochactaeon). The taxonomically crosscutting 

nature of Neotrochactaeon makes it of little value, and 

it is thus considered unusable.

We also consider the genus Mesotrochactaeon Hacobjan, 1972, as a synonym of T. (Trochactaeon). It was 

established for a group of Trochactaeon species with 

"elongate-oval to semicylindrical, narrow shells with a 

high or medium spire and elongate last whorl; lateral 

walls of whorls weakly convex; aperture significantly 

widened anteriorly, with the appearance of the ridgelike fold of the outer lip; three plaits on columella" 

(original diagnosis by Hacobjan, 1972, p. 8; translated 

from the Russian). In this group, Hacobjan included 

some nine species from the Cenomanian(?) and Turonian in an area ranging from California through North 

Africa and southern Europe to the Transcaucasus. He 

separated these species from Trochactaeon "by the 

semicylindrical rather than conoid outlines of the 

whorls and of the shell and by presence of a more developed spire" (original diagnosis by Hacobjan, 1972, p. 8; 

translated from the Russian). The group stands in an 

intermediate position between T. (Neocylindrites) and 

Trochactaeon s.s. relative to overall shell form. However, the whorls are broad in their lower part, as in 

T. (Trochactaeon), and because this characteristic is 

probably of more importance than the overall shell form, 

we prefer to leave this group within T. (Trochacteon).

Hacobjan (1972) established the new genus Purpuractaeon for five species of Trochactaeon that have nodes 

on the exterior of the shell. Their relationship will be 

discussed under Trochactaeon (Trochactaeon) woodsi 

Rennie (1930). The diagnosis of Purpuractaeon is as follows: "Shell large conical to oval, spire relatively high; 

whorls convex, sutures with a ramp; sculpture consists 

of spiral nodes at the upper margin of the whorls; aperture narrow posteriorly, anteriorly widening like a 

chamber; outer lip with a rounded ridgelike plait internally; strong columella with three plaits." (Original 

diagnosis by Hacobjan, 1972, p. 12; translated from the 

Russian.)

As diagnosed, Purpuractaeon differs from Trochactaeon (Trochactaeon) only by the presence of nodes at the 

shoulder. The nodes are formed by a periodic extension 

of the mantle. The function of such nodes is presently 

unknown, even in many living node-bearing gastropods. The presence or absence of spines or nodes, however, does not seem to reflect a major change in the 

organization of soft parts. Accordingly, we consider
48 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

these species as belonging to T. (Trochactaeon) and 

place Purpuractaeon in synonomy.

Subgenus Neocylindrites Sayn, 1932 

(Synonym: Paleotrochactaeon Hacobjan, 1972)

Type by original designation Trochactaeon (Neocylindrites) barremicus Sayn, 1932.

Diagnosis. "Trochactaeon with cylindrical or nearly 

cylindrical sides; whorls enclose the preceding ones to 

more than two-thirds of their height; aperture in the 

lower part only slightly wider than above; 2 to 3 columellar plaits." (Original diagnosis by Kollmann, 1967, 

p. 215; translated from the German.)

Discussion. T. (Neocylindrites) differs from T. (Trochactaeon) by its high cylindrical or subcylindrical 

whorls, each of which strongly overlaps the preceding 

one. The whorls are narrow, and the shell is therefore 

proportionally more slender.

Hacobjan (1972) proposed the genus Palaeotrochactaeon on the basis of species that originally were encompassed within the concept of T. (Neocylindrites). He 

included the following species: T. absalonis Fraas, 

T. angustatus Ptselintsev, T. briarti Geinitz, T. cordieroi Choffat, T. crisminensis Choffat (type species), 

and T. cylindraceus Stoliczka.

The diagnosis of Palaeotrochactaeon Hacobjan (1972, 

p. 6) is as follows: "Shell small, conical semicylindrical 

to cylindrical, with short spire and high last whorl, regularly tapering at base; surface of whorls flattened and 

smooth; aperture elongate and narrow, with straight 

thick walls, columella with three plaits." This translation of Hacobjan's diagnosis from the Russian virtually 

parallels that of T. (Neocylindrites).

Sayne's (1932) original description of the type species 

of T. (Neocylindrites) indicates a wide range of interspecific morphologic variability. Choffat (1901-1902) 

figured five specimens of Trochactaeon crisminensis, 

the type species of Palaeotrochactaeon Hacobjan, that 

show a degree of variability similar to that of T. (N.) 

barremicus in whorl convexity, midwhorl depression, 

and variable height of the spire. In both T. (Neocylindrites) and Palaeotrochactaeon the aperture is high and 

narrow. In view of the range of character variability 

shown by the two proposed taxa, plus the overall similarity of the two, we see no reason to distinguish 

Palaeotrochactaeon from T. (Neocylindrites).

Subgenus Sevanella Hacobjan, 1972

Type by original designation Sevanella laticostata 

Hacobjan, 1972.

Diagnosis. "Shell of medium to large size ovi-conical, 

with high spire. Whorls overall convex but irregular in 

profile because of presence of a broad and deep spiral 

depression in medial part; sculpture consists of strong,

transverse, foldlike ribs or of spiral rows of nodes that 

occur both on upper and lower portions of whorls; aperture short, narrow, and about equal width throughout 

its length; plait of outer lip and parietal callus rather 

strong, thus giving aperture a somewhat sinuate character; apertural walls much thickened. Columella massive and bears three plaits." (Original diagnosis of 

Hacobjan, 1972, p. 12; translated from the Russian.)

Trochactaeon (Sevanella) differs from Trochactaeon 

(Trochactaeon) by possessing a strong median depression of the body whorl and in having strong rounded 

transverse ribs above and below this depression. The 

palatal ridge is exceptionally strong. According to 

Hacobjan's (1972, pi. 2) figures, the shells also bear a 

small parietal ridge which was not mentioned in the 

diagnosis.

The subgenus is known only from the type species 

from the upper Coniacian of Armenia (fig. 8).

Subgenus Mexicotrochactaeon Hacobjan, 1972

Type by original designation Actaeonella (Trochactaeon) acutissima Bose = Actaeonella (Trochactaeon) 

coniformis Bose, 1906.

Diagnosis. "Shell narrow, biconical, or with a conical turriculate spire; last whorl low; upper part of 

whorls weakly convex, but narrowing strongly below 

toward point, becoming conical in outline anteriorly; 

aperture narrow, siltlike, short and very slightly broadened anteriorly; ridgelike plait on the outer lip weakly 

developed; columella with 3 plaits." Original diagnosis 

by Hacobjan, 1972, p. 13; translated from the Russian.)

Discussion. Bose (1906) described 10 species of Trochactaeon from the same stratigraphic horizon within 

the Cardenas, Mexico section. These are: Actaeonella 

(T.) coniformis; A. (T.) acutissima; A. (T.) occidentalis; 

A. (T.) gigantea, A. (T.) inconstans; A. (T.) irregularis; 

A. (T.) brevis; A. (T.) planilateralis; A. (T.) potosiana; 

A. (T.) variabilis. With the exception of Actaeonella 

(Trochactaeon) gigantea Sowerby, Myers (1968), in his 

study of the Cardenas section, considers them all as 

belonging to the same species, and, following article 

24(a) of the International Code of Zoological Nomenclature, he chose the name Trochactaeon coniformis for 

Bose's simultaneously published names. Hacobjan 

(1972) established the new genus Mexicotrochactaeon 

and selected Actaeonella (Trochactaeon) acutissima 

Bose, 1906, as type species. Myers (1968), the first reviser of Bose's species, considered T. acutissima as a 

junior synonym of Trochactaeon coniformis.

After examination of a large suite of specimens of the 

type species, we find that T. (Mexicotrochactaeon) can 

only consistently be characterized by the presence of a 

parietal fold or ridge. Minute differences in the whorl 

shape and especially in the palatal ridge, which Hacob-
SYSTEMATIC PALEONTOLOGY 49

jan used as criteria for separating T. (Mexicotrochactaeon) and the other subgenera of Trochactaeon, should 

not be used for establishing new genera or subgenera 

within this group.

Trochactaeon (Trochactaeon) woods! (Rennie)

Plate 4, figures 6-8, 12; plate 5, figures 1-5; plate 6, figures 1, 2, 4, 5; 

plate 7, figures 4, 5

1906. Actaeonella (Trochactaeon) sp. Woods, South African Museum

Annals, v. 4, p. 328, pi. 41, fig. 1. 

1930 Actaeonella (Trochactaeon) woodsi Rennie, South African Museum

Annals, v. 28, p. 232, pi. 28, figs. 17-21.

Diagnosis. A Trochactaeon with 11 to 12 strong 

nodes at whorl shoulder.

Original description. "Shell elongately oval, stout. 

Spire of about 6 whorls, moderately convex, the shape 

obscured by a row of very large, stout tubercles; spiral 

angle of 70°-80°. Last whorl longer than the spire, 

cylindrical in form, tapering gradually towards the 

anterior base tapering more rapidly; posterior part 

with a sloping ledge adjacent to the well marked but 

shallow suture, the upper part of the whorl ornamented 

with 10 to 12 very stout tubercles. Surface smooth, 

except for faint growth lines on the last whorl, slightly 

convex toward the aperture. Columella short, with three 

strong folds, of which the posterior is much stronger 

than the other two, and the anterior is weakest. Outer 

lip thin." (Rennie, 1930, p. 232.)

Additional description. Shells large to very large; 

maximum diameter 53-73 percent of total height. 

Spire variable in height, whorls convex, all before last 

one having 8-10 strong conical tubercles; growth lines 

strongly opisthoclinely sinused over tubercles, and 

curving opisthoclinely at broad collarlike substural 

ramp. Body whorl 68-89 percent of total height, with 

11 to 12 tubercles; regularly tapering below tubercles 

towards base in early stages, but convex sided or occasionally having midwhorl depression in later growth 

stages. Aperture broad, with deep posterior sinus at 

suture; columellar lip only slightly expanded and reflected at edge, which is formed by uppermost plait. 

Plaits decreasing in strength anteriorly. Parietal lip 

thin, decreasing slightly in thickness towards posterior 

part of aperture. Plaits resorbed anteriorly on all whorls 

except for last. Parietal wall of penultimate whorl 

resorbed adjacent to columella and forming a scar divided by a small ridge; on earlier whorls, this zone is 

completely resorbed and smooth.

Measurements (fig. 26). The full range of size among 

some 27 specimens is: H, 78-166 mm; MD, 52-102 mm; 

HB, 61-126 mm; PA, 60°-110°.

Discussion. The following Late Cretaceous species 

of T. (Trochactaeon) that have tubercles or nodes have 

been described:

170

150

130

110

LJJ I

90

70

50

20 30 50 70 90 

MAXIMUM DIAMETER

110

FIGURE 26. Scatter diagram comparing height with width among 

selected specimens of Trochactaeon (Trochactaeon) woodsi 

(Rennie).

T. (Trochactaeon) woodsi (Rennie) from Pondoland, 

South Africa (Santonian-Campanian), and Puerto Rico 

(upper Campanian-lower Maastrichtian).

T. (Trochactaeon) beyrichi (Drescher) from the Upper 

Quader of Giersdorf, the Papanberg near Blankenburg, 

and the sandstone of Kieslingswalde near Habelschwerdt 

(Bystrzyca Kiedzkaq), Silesian Poland (Coniacian).

Purpuractaeon conicus Hacobjan, 1963, from the 

Devi River, Armenian SSR. Because it is considered 

here as a Trochactaeon, the name becomes a junior secondary homonym of Trochactaeon conicus (Muenster) 

(upper Coniacian), but we refrain from substituting a 

new name without being able to study the type material.

T. (Trochactaeon) pcelinsevi (Hacobjan, 1963), also 

described as a Purpuractaeon from the Devi River, 

Armenian SSR. (Upper Coniacian).

Cassidaria cretacea Mueller, 1851, assigned by Holzapfel (1888) to Actaeonella, possesses nodes and a spire 

suggestive of this genus. The species was described 

from steinkerns; therefore, doubts exist about its true 

affinities.

The specimens that Rennie figured are small, and 

the heights of the spires vary considerably. The specimens show a depression on the side of the body whorl 

and have 10 to 12 nodes that lie in the same position on 

the whorl as do nodes on the specimens from Puerto 

Rico upon which the additional description was based.
50 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Although our specimens are generally much larger 

and broader than the African specimens, several have 

the same proportions. Sections (pi. 5, fig. 5) show that 

the earlier growth stages consisting of 4 to 5 whorls are 

usually not as much inflated as the later stages. The 

number and shape of tubercles are also the same in the 

specimens from Puerto Rico and those from Pondoland. 

In view of the lack of any significant differences between the specimens from the two areas, we consider 

them as belonging to one species.

T. (Trochactaeon) beyrichi (Drescher) has the same 

proportions as T. (Trochactaeon) woodsi but possesses 

only 8 very broad and rounded nodes per whorl. As in 

the very large specimens from Puerto Rico, the nodes 

are flatter on the body whorl.

In T. (Trochactaeon) pcelinsevi (Hacobjan), the nodes 

are large and rounded, as in Trochactaeon beyrichi, but 

on the basis of only the single figured specimen, the 

whorls are more inflated than those of T. (Trochactaeon) beyrichi. In total aspect, however, T. (Trochactaeon) pcelinsevi is rather similar to large specimens of 

T. (Trochactaeon) woodsi.

The species is common in certain areas in southwestern Puerto Rico. At locality 39 (USGS 30512) specimens 

occur as a virtually paucispecific bed of individuals. 

The specimen figured on pi. 5, fig. 1, is silicified and 

shows especially well the subspinose flangelike periodic extension of the nodes that is not seen on the 

larger worn specimens. This specimen also shows delicate subsutural features that are obliterated on other 

specimens because of poor preservation. Immediately 

below the suture is a channel with a concave base 

about 1.2 mm broad margined by a raised sharp-edged 

paper-thin rim. The growth lines are crescently sinused 

within this channel, which marks the position of the 

posterior (exhalent) siphon. On the bordering rim, the 

lines are strongly prosoclinely flexed, a direction that 

carries onto the steep slope above the shoulder, where 

the prosocline inclination is at an angle of about 105 ° 

to the axis of coiling. The growth lines bend down to 

only slightly prosocline over the shoulder, save where 

the shoulder nodes are produced into strongly opisthoclinely directed raised flanges (see pi. 5, fig. 1).

Occurrence. Puerto Rico: Barranquitas Quadrangle, 

Pozas Formation, Botijas Limestone Member, at localities 8 and 9 and Reves Member, at localities 5, 6, and 

7; San German Quadrangle, Cotui Limestone, at localities 39, 40, 41, 44, 45, and 46, and Sabana Grande Formation at locality 43. South Africa: Natal (Pondoland), 

Umzamba Formation.

Age. The type occurrence in the Umzamba Formation on the coast of Natal (Pondoland), South Africa, 

has been considered by Kennedy and Klinger (1975) to 

range in age from late Coniacian to early Campanian.

Subsequently, Kennedy (1979) suggested a Santonian 

to Campanian Age for the formation. The exact position within this range from which T. (T.) woodsi was 

collected is not known. In Puerto Rico, the species is 

associated with molluscan faunas considered of possible latest Campanian Age and ranging into the earliest Maaestrichtian.

Types: Holotype, Transvaal Museum; hypotypes, 

Puerto Rico, USNM 306006, 306007, 306011-30618.

Trochactaeon (Trochactaeon) cf. T. (Trochactaeon) woodsi (Rennie)

Plate 4, figures 9-11; plate 6, figure 3; plate 7, figures 1-3.

Description. Shells globular, spire proportionally 

low; about seven whorls; body whorl about 80 percent 

of total height; whorl sides convex, maximum diameter 

in upper third. Body bearing 11 broadly round-topped, 

flattened nodes on subsutural shoulder; growth lines 

weakly sinused over subsutural area, gently prosocline 

over upper third of body below shoulder and then gently 

prosocline over basal slope. Aperture proportionally 

broad, columella thick, columellar lip only slightly 

expanded and bearing three plaits; parietal lip of equal 

thickness throughout its full length; palatal ridge 

broad and strong.

Measurements. The most complete specimen, which 

lacks about 5 mm of its apical tip, measures 75 mm in 

length, 60 mm in maximum diameter, and 63.7 mm in 

body-whorl length.

Discussion. All the available specimens of this form 

are somewhat deformed and eroded or worn. They 

undoubtedly do, however, represent a noded Trochactaeon with undetermined affinities to T. (Trochactaeon) 

woodsi (Rennie). Compared with the Puerto Rican specimens previously assigned to that species, the nodes of 

the better preserved specimens are more rounded but 

less prominent and thus may represent a new form.

Occurrence. Puerto Rico: Orocovis Quadrangle, 

Pozas Formation, Rio Bauta Member, at localities 25, 

26, and 27.

Age. Lower Campanian.

Types.-Hypotypes, USNM 306008-30610, 306019, 

306020.

Trochactaeon (Trochactaeon) frazierensis (Anderson) 

Plate 8, figures 1-5

1958. Acteonella frazierensis Anderson, Geol. Soc. America mem. 71,

p. 161, pi. 29, figs. 7, 7a. 

1958. Acteonella rustica Anderson, Geol. Soc. America mem. 71,

p. 161, pi. 29, fig. 5.

Diagnosis. Trochactaeon of medium size with a proportionally high spire. Whorls with broad subsutural 

ramp, slightly rounded in upper part of whorl and subcylindrical below. Body whorl inflated medially. Columella short.
SYSTEMATIC PALEONTOLOGY 51

Description. "The mature shell is of moderate size, 

biconical in general aspect, spire of moderate height; 

body whorl of equal length sloping downward to columella; aperture nearly straight, narrow above, and 

expanded below; outer lip thin and nearly straight; 

inner lip converging gradually from outer lip above, 

expands suddenly below to anterior canal; columella 

carries three oblique plications, the anterior being 

reduced in size near anterior end of columella. Shoulders of whorl slightly rounded and tabular above, with 

a slight collar rising on the preceding whorl." (Anderson, 1958, p. 161).

Additions to Description. The spire is somewhat 

convex in profile and proportionally high for the genus. 

The whorl sides are slightly rounded in their upper 

posterior portions and primarily cylindrical below. A 

subsutural ramp is a prominent feature and is nearly 

horizontal to the shell axis. No collar such as described 

by Anderson, however, could be observed. On the larger 

specimens (pi. 8, fig. 5), the body whorl is more convex 

than on smaller specimens (pi. 8, fig. 1), bears a slight 

depression, and is 71 to 78 percent as long as the total 

shell length. The aperture is broad and the columella is 

much shorter than indicated in Andersen's (1958, pi. 29, 

figs. 5, 7, 7a) illustrations (compare pi. 8, figs. 1,4, and 

5). The outer margin of the inductura is deviated about 

20 ° away from the shell axis.

Measurements. The incomplete holotype (GAS 

10664) is 43.4 mm high and has a maximum diameter 

of 21.5 mm and a body-whorl length of 27.9 mm. The 

pleural angle is 48 °. Two paratypes measure as follows 

(in mm except as shown):

H

53.5

52.5

MD

28.4

30.8

HB

38.3

40.0

PA

50°

40°

Discussion. The type specimens of (T.) Trochactaeon 

frazierensis (Anderson) come from lower Turonian beds 

near Redding in Shasta County, Calif. In the same publication in which Anderson (1958) defined this species, 

a second species, Actaeonella rustica, was also described 

from other exposures of lower Turonian rocks within 

the same area. In the description of A. rustica, Anderson (1958, p. 161) stated:

Shell similar to Actaeonella frazierensis n. sp., although relatively 

stouter and more biconic in form, and distinct in most of its other 

external features. The holotype ***consists of the body whorl and 

five spiral whorls including apex, which is slightly reduced by weathering of shell substance***. The body whorl is little impressed near 

its middle, thus modifying its conical form, and in some examples 

giving them a slightly barrel shape.

Andersen's figures of both species are rather sketchily hand drawn and in obvious error in several regards. 

Examination of Andersen's specimens shows, for example, that they are all incomplete and that the total

number of whorls cannot be determined save that there 

are more than the stated five whorls. The most nearly 

complete specimens possess a body whorl that isw 45 to 

60 percent of the total shell height.

Although Anderson (1958) separated his species 

Actaeonella rustica and A. frazierensis primarily because the latter has more convex sides, a comparison 

of the photographs of the type specimen on plate 8, figure 2 with the original hand-drawn figures shows exaggeration in this character. We see no significant difference between the two named species in this regard and 

synonymize A. rustica under A. frazerensis by page 

preference.

Some specimens of T. (Trochactaeon) matensis (Fittipaldi) may have an equally high convex spire, and a 

subsutural ramp of equal strength as T. (Trochactaeon) 

frazierensis (see Pchelintsev, 1953; Lupu, 1965) but, 

they differ in always possessing a proportionally long 

and evenly convex body whorl that lacks any medial 

constriction or depression. Trochactaeon (Trochactaeon) lamarcki (Sowerby), a common species in Upper 

Cretaceous deposits of Central Europe and the Middle 

East, is somewhat similar but possesses a proportionally shorter spire and more convex whorls.

Occurrence. California (Siskiyou and Shasta Counties), Hornbrook Formation and Chico Formation, at 

localities 161 to 165.

Age. Turonian.

Types. Holotype (A. frazierensis), GAS 10664; paratype, GAS 10665. Holotype (A. rustica), GAS 10666.

Trochactaeon (Trochactaeon) packardi (Anderson)

Plate 8, figures 15-17

1902. Actaeonella oviformis Gabb. Anderson, California Acad. Sci.

Proc., 3d ser., v. 2, p. 28, 34. 

1916. Actaeonella oviformis Gabb. Packard, California Univ., Dept.

Geol. Sci. Bull., v. 9, no. 2, p. 144, 147. 

1922. Actaeonella oviformis Gabb. Packard, California Univ. Dept.

Geol. Sci. Bull., v. 13, no. 10, p. 458, pi. 36, fig. 4. 

1954. Acteonella oviformis Gabb. Popenoe, California Div. Mines

Bull. 170, table 1, fig. 3, no. 2. 

1958. Acteonella oviformis Gabb. Anderson, Geol. Soc. America Mem.

71, p. 160, pi. 29, fig. 6. 

1958. Actaeonella packardi Anderson. Geol. Soc. America Mem. 71,

p. 160, pi. 29, fig. 4, 4a, 4b.

Diagnosis. Shell of small to medium size, broadly 

subcylindrical. Spire low to immersed. Subsutural 

ramp broad and horizontal. Palatal swelling weak.

Description. Shell of small to medium size, elongate 

oviform, maximum diameter 47 to 58 percent of total 

height. Spire very short to immersed. Shell of 6 to 7 

whorls that have a broad subsutural ramp horizontal 

to shell axis. Height of body whorl 84 to 99 percent of 

total height. Body slightly convex, often with a weak 

inflation above midline. Sides of columella form a very
52 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

acute angle to shell axis. Columellar lip reflected and 

thick outside aperture; posteriormost plait delimits 

upper edge of columella and is only plait visible outside 

aperture. Within shell, posterior plait strongest, medial one weakest; columella below plaits short; parietal 

lip equally thin throughout its length; palatal swelling 

weak and only present in second half of penultimate 

whorl and in first part of body whorl; columella resorbed 

in earliest parts of last whorl and remainder of shell. 

Measurements (in mm except as shown) (see fig. 27). 

Holotype 

Paratypes

H

66.2 (estimated 68) 

21.7 to 75 

(estimated 80)

MD

30.0

11.2 to 41.7

HB

59.5

21.7 to 73.4

PA

110 C

Discussion. The specimens that Anderson (1958) 

assigned to Acteonella oviformis Gabb are silicified like

90

80-

70

60-

I- 50

I

LLJ

I

40

30

20

10

I I

10 20 30 40 

MAXIMUM DIAMETER

50

FIGURE 27. Scatter diagram comparing height with width among 

selected specimens of Trochactaeon (Trochactaeon) packardi 

(Anderson).

the sandstone in which they have been found, but imperfect replacement results in all isolated specimens 

being more or less incomplete. The type of Actaeonella 

oviformis Gabb is similarly preserved, the outer whorls 

being incomplete. Gabb's specimens, however, have 

straight growth lines and are referable to Actaeonella, 

whereas the specimens described by Anderson (1902, 

1958) under the name oviformis belong to the genus 

Trochactaeon. This is also true of Anderson's Acteonella 

packardi. Anderson's holotype of A. packardi, which is 

the same specimen Packard (1922) figured and assigned 

to Actaeonella oviformis Gabb, is about twice as high as 

most of the silicified specimens of his Acteonella oviformis. Among the specimens from localities CAS 252 

and 733, the size range easily encompasses both Acteonella oviformis and A. packardi of Anderson. As there 

are no significant differences in the height-width ratios 

and in general morphology, Anderson's A. oviformis is 

considered to be a synonym of Trochactaeon (Trochactaeon) packardi.

Trochactaeon (Trochactaeon) matensis (Fittipaldi) 

and T. (Trochactaeon) cylindraceus Stoliczka are morphologically closely similar to T. (Trochactaeon) packardi. 

T. (Trochactaeon) matensis was described by Fittipaldi 

from the Turonian of Campobasso, Italy. It also occurs 

in the Cenomanian and lower Turonian of the TransCaucasus area, where it includes T. (Trochactaeon) 

cylindricus Pchelintsev (1928) and T. (Trochactaeon) 

pseudocylindraceus Pchelintsev (1953). T. (Trochactaeon) matensis, recorded from the upper Cenomanian 

of Cherghes, Romania, by Lupu (1965) has a more convex body whorl and a spire proportionally higher and 

more convex in profile than that of T. (Trochactaeon) 

packardi (Anderson). Longitudinal sections show a 

much stronger palatal swelling than in T. (Trochactaeon) packardi, which narrows the interior of the 

whorls considerably.

The body whorl of T. (Trochactaeon) cylindraceus 

Stoliczka from the Arialoor Group of southern India is 

proportionally shorter than that of T. (Trochactaeon) 

packardi and T. (Trochactaeon) matensis. Stoliczka 

(1868) pointed out the obtuse attenuation of the lower 

part of the body whorl, which bends strongly towards 

the axis from the upper more cylindrical part. The 

internal molds of a Trochactaeon Darteville and Brebion 

(1956) described as Actaeonella (Trochactaeon) cylindracea Stoliczka are too poorly preserved to allow any 

assignment to a known species of Trochactaeon.

Occurrence. California: Siskiyou County, at localities 163 and 166; Shasta County, at localities 161,162, 

and 168; Orange County (Baker Canyon Conglomerate 

Member, Ladd Formation), at locality 167.

Age. Turonian.

Types.-Holotype, CAS 10663; hypotype CAS 733.
SYSTEMATIC PALEONTOLOGY 53

Trochactaeon (Neocylindrites) cumminsi Stanton

Plate 4, figures 1-4; plate 9, figures 9 19

1947. Trochactaeon cumminsi Stanton. U.S. Geol. Survey Prof. Paper 211,

p. Ill, pi. 63, figs. 3-6, 13. 

1947. Trochactaeon parvus Stanton. U.S. Geol. Survey Prof. Paper 211,

p. 112, pi. 63, figs. 1, 2. 

1955. Actaeonella parvus (Stanton). Allison, Jour. Paleontology, v. 29,

p. 429, pi. 44, fig. 5.

Diagnosis. Shell small to medium size, subcylindrical and high. Whorls moderately convex, with a very 

broad and strongly inclined subsutural ramp. One to 

two smaller folds occur below main palatal ridge.

Description. "Shell of medium size, slender, subfusiform, consisting of about 8 whorls, of which the last 

constitutes two-thirds to three-fourths of the entire 

length; spire variable, in some specimens mammillated at the summit, the apical angle commonly about 

65 ° but ranging from 60 ° to nearly 80 °; suture slightly 

impressed; whorls of the spire gently convex, but some 

whorls bear a very slight angulation near the middle, 

which marks the position of a fine impressed line that 

forms the anterior border of a broad slit-band or sinus 

comparable with that in Nerinea; last whorl relatively 

very large, subcylindrical, with an oblique flattening 

or shoulder behind occupied by the slit-band, and somewhat produced and contracted in front, the extreme 

anterior end being obliquely subtruncate; surface 

marked only by fine growth lines that pass almost 

straight across the whorl in a broad curve with its convexity directed forward, the curvature increasing suddenly as the lines approach the slit-band, which they 

cross in a short curve turned in the opposite direction; 

inner lip thickened in front and bearing 3 strong folds; 

aperture long and narrow, slightly increasing in width 

toward the front.

"Internal molds differ greatly from the shell in form, 

the whorls being prominently shouldered with the 

maximum breadth behind, and the last whorl showing 

a well-marked spiral groove or constriction a little in 

advance of the middle, with a second and occasionally a 

third groove less distinctly marked in front of it.

"The largest of the types, one of the more slender forms 

with an apical angle of 60 °, has the following measurements: Height 76mm., height of spire 26mm., maximum 

breadth 27mm., breadth of slitband 5mm. Corresponding measurements of a stouter specimen with an apical 

angle of 70° are 60, 15, 25, and 3.8mm." (Stanton, 

1947, p. Ill, 112.)

As we consider Trochactaeon parvus Stanton a synonym of T. (Neocylindrites) cumminsi, we quote a passage from Stanton's description of Trochactaeon parvus 

(1947, p. 112): "Shell small, slender subfusiform, consisting of 5 or 6 whorls, of which the last constitutes 

nearly two-thirds of the entire length; greatest breadth

near the posterior (upper) margin of the last whorl, 

from which the shell tapers regularly in both directions, 

the apical angle of the spire about 30 ° whereas the 

sides of the last whorl converge toward the narrow 

anterior end with an angle of about 20°; exposed portions of whorls of the spire slightly convex, as in Trochactaeon (Neocylindrites) cumminsi Stanton; surface 

apparently smooth, but the state of preservation is 

such that the growth lines are not visible, nor is the 

slit-band on the sloping shoulder of the whorl; aperture 

long and narrow; inner lip with 3 strong spiral folds 

near the front."

Measurements (see fig. 28). The largest of the measured specimens is 74.0 mm, the shortest, 3.3 mm. 

Small specimens have a maximum diameter as much 

as 50 percent of total height, whereas larger specimens 

have a maximum diameter only 33 to 44 percent of 

total height. The body whorl constitutes 73 to 89 percent of total height.

Discussion. To the description cited above, we may 

add that the basal lip is deeply sinuate and evanesces 

out of the aperture onto the basal part of the body whorl. 

The inductura is very thin (pi. 9, figs. 16, 19). The columellar plaits are visible for a full one-half volution on 

the body outside the aperture, but the columellar rim

70

60

50

40

30

20

10

i r i r i r

10 20 30 40

MAXIMUM DIAMETER

FIGURE 28. Scatter diagram comparing height of shell with 

width among selected specimens of Trochactaeon (Neocylindrites) cumminsi Stanton.
54 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

is poorly defined. Internal molds (pi. 9, figs. 12, 18) 

show two or three grooves on the palatal area that represent ridges on the shell. The largest of these is present 

up to the aperture, but the secondary ridges disappear 

before the aperture is reached. Stanton (1947) pointed 

out the similarities in shell morphology of T. (Neocylindrites) cumminsi and T. (Neocylindrites) parvus. As the 

main difference between these two species, he cited 

(p. 112) the "narrow gradually tapering anterior" of 

T. parvus, as his figures (Stanton, 1947, pi. 63, figs. 1, 

2) suggest. Our examination of the figured specimens 

shows that the fragile outer lip is not preserved so that 

the impression of a very narrow aperture is an artifact 

of preservation and is erroneous. With this difference 

eradicated, we conclude that the two are synonymous, 

T. (Neocylindrites) cumminsi having page preference.

Compared with the European species Trochactaeon 

(Neocylindrites) briarti Geinitz, 1874, and its possible 

synonym T. (Neocylindrites) crisminensis Choffat, 

1901, the sides of the last whorl are barely convex and 

taper straight towards the base. In addition, the subsutural ramp is not as prominent as in T. (Neocylindrites) 

cumminsi. Trochactaeon (Neocylindrites) boutillieri 

Cossmann (1895) together with T. (Neocylindrites) barremicus Sayn (1932) are very small and possess a spire 

that is very variable in height. In the juvenile stages, 

few differences exist between these species and Trochactaeon (Neocylindrites) cumminsi, except for a very 

distinct constriction in the upper part of the last whorl 

in T. (Neocylindrites) boutillieri. This constriction also 

persists in the adult specimens of T. (Neocylindrites) 

barremicus var. mumiaeformis Sayn (1932, p. 9, pi. 1, 

fig. 5).

Occurrence. Texas: Devils River Limestone, Val 

Verde County (localities 134, 135, 138); Washita 

Group, Reagan County (localities 136, 137), Brewster 

County (localities 142,143), and Pecos County (locality 

141); Edwards Limestone, Kerr County, "on Colorado 

River above the mouth of Barton's Creek near Austin" 

(Stanton, 1947, p. 112) (locality 132). Baja California: 

Allisitos Formation.

Age. Middle to late Albian.

7>pes.-Lectotype, USNM 306042; syntypes, USNM 

306038-306041, 306043, 306044; hypotypes, USNM 

306001-306004.

Trochactaeon (Neocylindrites) punctatus (Maldonado-Koerdell), 1950

Plate 4, figure 5; plate 9, figures 1-6

1950. Actaeonella cf. A. (Trochactaeon) coniformis Bose. MaldonadoKoerdell, Mexico Escuela Nac. Cienc. Biol. Anales, v. 6, 

p. 214, figs. 44, 45.

1950. Actaeonella (Trochactaeon) acutissima punctata MaldonadoKoerdell, Mexico Escuela Nac. Cienc. Biol. Anales, v. 6, 

p. 215, figs. 46a, b.

1950. Actaeonella cf. A. (Trochactaeon) occidentals Bbse. MaldonadoKoerdell, Mexico Escuela Nac. Cienc. Biol. Anales, v. 6, 

p. 215, fig. 47.

1950. Actaeonella (Trochactaeon) planilateralis Bose. MaldonadoKoerdell, Mexico Escuela Nac. Cienc. Biol. Anales, v. 6, 

p. 217, fig. 52.

Description. Shells of medium size; maximum diameter 37-39 percent of total height; spire high in small 

specimens, proportionally low and with concave sides 

in large ones; whorls with a broad and inclined subsutural ramp terminated distinctly against whorl sides. 

Body whorl 80 to 89 percent of total height, slightly 

inflated in upper third; whorl sides tapering first at a 

low angle and then with slightly increasing steepness 

towards the narrow base; middle of whorl bearing distinct constriction. Aperture high and narrow. Columella proportionally high, thin, and only slightly expanded, outer rim smooth, bending towards base almost vertically; plaits of about equal strength, distance 

between upper two wider than between middle and 

anteriormost one. In large specimens, one additional 

weak plait occurs between upper two plaits. Parietal 

lip thin, edge inclined at about 20 ° to shell axis. Border 

of inner lip inclined 20 ° to shell axis in parietal region, 

bending downwards almost vertically at columella, 

which is little expanded.

Measurements. -The six specimens (USGS 11467) 

available for measurement show a range of: total height, 

21.1-47.5 mm; maximum diameter, 8.0-19.8 mm; 

pleural angle 50 °-70 °. The lower values are mainly on 

small specimens.

Discussion. Maldonado-Koerdell (1950) figured five 

specimens of actaeonellids from the State of Chiapas in 

southern Mexico. He assigned all his specimens to taxa 

previously described by Bbse in 1906 from the Maestrichtian of Cardenas, Mexico, but he erected one new 

subspecies Actaeonella (Trochactaeon) acutissima 

punctata. Maldonado-Koerdell's material appears to us 

to belong to a single species differing from specimen to 

specimen only in the form and height of the spire. These 

characters are among the most variable in samples of 

almost any species of the genus. Elsewhere herein, we 

have accepted all Bose's species and subspecies to 

which Maldonado-Koerdell assigned his Chiapas specimens to be parts of one variable species Trochactaeon 

(Mexicotrochactaeon) coniformis Bbse. We consider the 

Chiapas specimens to differ from that species in having 

proportionally higher and much less convex whorls 

and a smooth parietal area. These features are characteristic of the subgenus Neocylindrites. We therefore 

raise Maldonado-Koerdell's subspecific name punctata 

to species rank and include all the Trochactaeon from 

Chiapas that he discussed within that species and 

assign them to Neocylindrites.
SYSTEMATIC PALEONTOLOGY 55

Occurrence. Mexico; State of Chiapas, at localities 

148, 151.

Age. Maastrichtian. Maldonado-Koerdell (1950, 

p. 189) assigned the type specimens to the upper Campanian. The listing of associated species, however, suggests a Maastrichtian Age.

Types. The holotype is in the collections of Petroleos 

Mexicanos; hypotypes, USNM 306005, 306031-306035.

Trochactaeon (Neocylindrites) nelsoni Sohl and Kollmann, n. sp.

Plate 8, figures 9-14, 18, 19; plate 11, figures 6, 7, 10

Diagnosis. A T. (Neocylindrites) with broadly convex whorls which are interiorly broad; columella thin, 

little expanded.

Description. Shells of small to medium size, subcylindrical, maximum diameter 39 to 54 percent of total 

height; spire low, whorls with medium broad inclined 

subsutural ramp. Pleural angle generally 60 ° to 70 ° in 

small specimens and 70 ° to 90 ° on larger forms. Body 

whorl length 81 to 93 percent of total height, lateral 

parts forming a distinct angle with ramp; body evenly 

convex over sides, tapering over lower part more rapidly toward base than in upper part; small specimens 

tapering evenly towards base throughout. Aperture 

high, rather broad; columella proportionally long and 

thin, with three plaits that decrease in size anteriorly; 

columella only slightly expanded, outer margin smooth; 

palatal ridge small; resorption of columella starts 

within last whorl; sides of earlier whorls in lower half 

nearly completely resorbed, whereas in upper parts, 

original shell thickness is maintained. Measurements 

are shown on figure 29.

Discussion. T. (Neocylindrites) nelsoni shows a 

great difference in average size at various localities. 

Low in its range zone at the type locality in the Rio 

Bauta Member of the Pozas Formation, it is rather 

small. Upward in its stratigraphic range, average size 

increases, and it reaches its largest size in the Flor de 

Alba Limestone Lentil of the Pozas. As seen in figure 29, 

although size may vary between localities, shell proportions remain remarkably constant save for smaller 

specimens being less cylindrical and more tapering in 

outline.

T. (Neocylindrites) nelsoni differs from T. (Neocylindrites) punctatus (Maldonaldo-Koerdell) 1950 by having more cylindrical sides on the body whorl that go 

straight down from the subsutural ramp without strong 

inflation. In addition, T. (Neocylindrites) nelsoni lacks 

a constriction in the middle of the shell, the whorls are 

broader interiorly and lack the fourth columellar plait 

typical of large specimens of T. (Neocylindrites) punctatus. Both T. gradatus Kollmann (1967) and T. gosaviensis Kollmann (1967) differ in possessing a more convex 

body whorl, a horizontal subsutural ramp, and strongly

70

60

50

I- 40

X

LUI

30-

20

10-

0 10 20 30 40 

MAXIMUM DIAMETER

FIGURE 29. Scatter diagram comparing height with width among 

selected specimens of Trochactaeon (Neocylindrites) nelsoni 

Sohl and Kollmann, n. sp.

reflected columella. Trochactaeon (Neocylindrites) 

cumminsi Stanton (1947) has a very broad and strongly 

inclined subsutural ramp, narrow and high whorls, a 

palatal ridge that is very strong, and two additional 

ridges towards the base. This species is named in honor 

of Arthur Nelson of the U.S. Geological Survey, for his 

contributions to the knowledge of the geology of the 

Florida area, Puerto Rico.

Occurrence. Puerto Rico: Pozas Formation, Rio 

Bauta Member (Orocovis Quadrangle), at locality 28; 

and Flor de Alba Limestone Lentil (Florida Quadrangle), at localities 22, 23, 24; Maravillas Formation 

(Coamo Quadrangle), at locality 20; Cotui Limestone 

(San German Quadrangle), at locality 40 (questionable 

assignment).

Age. Campanian.

Types.-Holotype, USNM 306028; paratypes, USNM 

306024-306027, 306029, 306030, 306064-306066.

Trochactaeon (Neocylindrites) sp. A

Plate 8, figures 6, 7, 8; plate 11, fig. 11

Description. Shells of medium size, maximum diameter 30 to 33 percent of total height. Spire very short, 

with flat sides. Apical angle about 80 °. Sutures, due to 

silification, indistinct. Body whorl high, subcylindri-
56 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

cal, with slightly convex sides. Aperture wider in lower 

part than in upper. Columella with three large plaits of 

equal strength broadly reflected at external margin. 

Deep incision above upper plait. Basal lip thickened 

and deeply canaliculate, cut off at an angle of 120 ° to 

axis.

Measurements. Two typical specimens measure as 

follows (in mm except as shown):

H

41.5 

35.8

MD 

14.0 

10.9

PA

76° 

80°

Discussion. The known well-preserved specimens of 

this species are too few to serve as the basis for a new 

species, but this form seems distinct from previously 

described species. The maximum diameter of Trochactaeon (Neocylindrites) cumminsi is larger than that of 

T. (Neocylindrites) sp. A, and the sides are more convex 

in specimens of comparable height. In T. (Neocylindrites) cumminsi, the body whorl diameter is about 

40 percent of total height, whereas in T. (Neocylindrites) 

sp. A it is about 30 percent and the basal lip, which is 

not very often preserved, seems to be less sinuate and 

evanescent.

Occurrence. Puerto Rico: limestone within the El 

Rayo Formation (Sabana Grande Quadrangle), at locality 36.

Age. Middle Maastrichtian.

Types.-Hypotypes, USNM 306023, 306084, 306069.

Trochactaeon (Neocylindrites) sp. B

Plate 9, figures 7, 8

Description. Shell of medium size, spire low with 

slightly convex or flat profile. Whorl sides are flat and 

taper conically towards base. Columella poorly known, 

but bearing three plaits.

Discussion. Only external molds in a silicified limestone are known. The molds are of shells that are less 

convex than that of Trochactaeon cumminsi Stanton 

and that have lower whorls and a less strongly developed subsutural ramp. They seem to be closely related 

to the T. (Neocylindrites) crisminensis-briarti group of 

Europe.

The specimens described above come from a silicified 

unit on the southern flank of Sierra Rica in southwestern New Mexico. Zeller (1965), in his study of the nearby 

Big Hatchet Mountain area, suggested that the silicified limestone of the Sierra Rica was an equivalent of 

his Mojado Formation. Though Zeller did not report 

the presence of Trochactaeon in the Big Hatchet Mountains, Actaeonella is present. In addition to the Sierra 

Rica Trochactaeon, the limestone blocks contain nerineids and bivalves. On the basis of contained fauna, 

the Mojado Formation is of both Fredricksburg and

Washita age. The nerineid and actaeonellid assemblages 

come from the upper (Washita) part of the formation. 

Thus, it seems most likely that the Trochactaeon here 

discussed is of late Albian Age.

Occurrence. New Mexico: Mojado Formation, at 

locality 147.

Age. Late Albian.

Types.-Hypotypes, USNM 306036, 306037.

Trochactaeon (Mexicotrochactaeon) coniformis (Bose)

Plate 10, figures 1-17; plate 11, figures 1-4

1906. Actaeonella (Trochactaeon) coniformis Bose. Mexico Inst, Geol.

Bol. 24, p. 78, pi. 16, figs. 12-21. 

1906. Actaeonella (Trochactaeon) acutissima Bose. Mexico Inst. Geol.

Bol. 24, p. 79, pi. 16, figs. 411. 

1906. Actaeonella (Trochactaeon) occidentalis Bose. Mexico Inst. Geol.

Bol. 24, p. 81, pi. 17, figs. 2-10. 

1906. Actaeonella (Trochactaeon) inconstans Bose. Mexico Inst. Geol.

Bol. 24, p. 83, pi. 17, figs. 11-19. 

1906. Actaeonella (Trochactaeon) irregularis Bose. Mexico Inst. Geol.

Bol. 24, p. 84, pi. 17, figs. 20 27. 

1906. Actaeonella (Trochactaeon) brevis Bose. Mexico Inst. Geol. Bol. 24,

p. 85, pi. 18, figs. 1-7. 

1906. Actaeonella (Trochactaeon) planilateralis Bose. Mexico Inst. Geol.

Bol. 24, p. 87, pi. 18, figs.8-13. 

1906. Actaeonella (Trochactaeon) potosiana Bose. Mexico Inst. Geol.

Bol. 24, p. 88, pi. 18, figs. 14-25. 

1906. Actaeonella (Trochactaeon) variabilis Bose. Mexico Inst. Geol.

Bol. 24, p. 90, pi. 18, figs. 26 34.

1968. Actaeonella coniformis Bose. Myers (part), Mexico Univ. Nac. 

Autonoma, Inst. Geologia, Paleontologia Mexicana, no. 24, 

p. 75, pi. 7, fig. 3, ?5 (not fig. 4). 

1972. Mexicotrochactaeon acutissima Bose. Hacobjan, Paleont. Zhur.

no. 1, p. 13. 

1972. Mexicotrochactaeon inconstans Bose. Hacobjan, Paleont. Zhur,

no. 1, p. 13. 

1972. Mexicotrochactaeon irregularis Bose. Hacobjan, Paleont. Zhur.

no. 1, p. 13. 

1972. Mexicotrochactaeon planilateralis Bose. Hacobjan, Paleont. Zhur.

no. 1, p. 13. 

1972. Mexicotrochactaeon potosianus Bose. Hacobjan, Paleont. Zhur.

no. 1, p. 13. 

1972. Mexicotrochactaeon variabilis Bose. hacobjan, Paleont. Zhur.

no. 1, p. 13. 

Not: 1950. Actaeonella cf. A. (Trochactaeon) coniformis Bose.

Koerdell, Mexico Escuela Nac. Cienc. Biol. Anales, 

v. 6, p. 214, figs. 44, 45.

Not: 1950. Actaeonella (Trochactaeon) acutissima punctata Bose. 

Maldonado-Koerdell, Mexico Escuela Cienc. Biol. 

Anales, v. 6, p. 215, fig. 46a, b. 

Not: 1950. Actaeonella cf. A. (Trochactaeon) occidentalis Bose.

Maldonado-Koerdell, Mexico Escuela Nac. Cienc. Biol. 

Anales, v. 6, p. 215, fig. 1. 

Not: 1950. Actaeonella cf. A. (Trochactaeon) inconstans Bose.

Maldonado-Koerdell, Mexico Escuela Nac. Cienc. Biol. 

Anales, v. 6, p. 216, fig. 50.

Not: 1950. Actaeonella (Trochactaeon) planilateralis Bose. MaldonadoKoerdell, Mexico Escuela Nac. Cienc. Biol. Anales, 

v. 6, p. 215, fig. 47.

Diagnosis. Small shells with highly variable spire; 

whorls with collarlike subsutural ramp. Parietal fold 

prominent.
SYSTEMATIC PALEONTOLOGY 57

Description. Shells small, with more or less high, 

convex spire. Maximum diameter 32 to 52 percent of 

total height. Shell of six to seven whorls on strongly 

evolute specimens. Whorl sides of high-spired forms 

convex, greatest diameter at midheight, slightly compressed posteriorly and convex below. On low-spired 

forms, only upper depressed part of whorls visible. 

Body whorl covered with fine growth lines. Subsutural 

ramp strongly inclined to axis, collarlike, often concave. Body whorl 51 to 91 percent of total height, with 

depressed shoulder in upper fifth of whorl; below, subconical in outline but somewhat compressed medially. 

Columellar lip slightly inclined towards axis, uppermost of three plaits most prominent; distance between 

lowermost plait and basal lip short. Thickening of 

lower part of parietal lip within aperture continuous 

with oblique sharp parietal fold on internal whorl surface. In large specimens, this fold is slightly concave 

below; parietal zone above plait slightly depressed. 

Resorption of plaits starts at junction of last whorl with 

penultimate whorl; about Vz whorl behind this point, 

columella and parietal region of penultimate whorl 

completely smooth. Palatal ridge indistinct because of 

recrystallization but appears more oblique in penultimate whorl than in body whorl.

Measurements (fig. 30). The 87 specimens measured 

ranged in total height from 9.0 to 55.4 mm, and the 

pleural angle ranged from 19° to 105°

II

UJ

I

50

40

30

20-

10

1 1 1 1

0 10 20 30 40 

MAXIMUM DIAMETER

FIGURE 30. Scatter diagram comparing height with width among 

selected specimens of Trochactaeon (Mexicotrochactaeon) coniformis (Bb'se).

Discussion. In addition to the above description, one 

specimen possessed a narrow longitudinal groove on 

the interior inner surface of the body whorl that seemingly represents a muscle impression. The measurements were made on specimens from units 69, 80, and 

98 of Wade's measured section along the Cardenas railroad in Mexico, published by Imlay (1944). These units 

contain all the "species" of Trochactaeon Bb'se described 

from the cut of the railway, 415 to 416 km from Tampico.

One of the most significant features of the forms collected in unit 80 is the extreme variability, which is 

caused primarily by the relative amount of overlap of 

one whorl onto the preceding one. This overlap is not 

only different in different specimens but often changes 

within one specimen (see pi. 10, fig. 9). The varying 

amount of overlap of the whorls is responsible for the 

differences in proportional height and size of the spire. 

The visible parts of the whorls are convex in the less 

overlapped and flat or even concave in the more overlapped whorls (compare figs. 12 and 17 on pi. 10). These 

differences were the criteria Bb'se used to distinguish 

his many species at Cardenas. As there is no other significant difference in Bbse's "species," we agree with 

Myers (1968), who considered all of them, except for 

Bbse's Actaeonella (Trochactaeon) aff. gigantea Sowerby, as belonging to one variable species, Actaeonella 

coniformis Bb'se.

The few specimens we have from Wade's unit 69, 

which is 127.3 m lower in his section than unit 80, do 

not have as strongly shouldered convex whorls. The 

whorls, however, do have shoulders, the subsutural 

ramps are steep, and the parietal ridge is strong; therefore, there is no reason to separate these specimens 

from Trochactaeon (Mexicotrochactaeon) coniformis. 

Although not separable as a distinct species, the specimens of unit 69 are not as variable as those in unit 80 

and may be compared with the large varietal form 

named Actaeonella (Trochactaeon) acutissima Bb'se 

(1906, pi. 16). This observation is consistent with Bbse's 

mention on his p. 17 that A. (Trochactaeon) acutissima 

occurs in the lower beds in the exposures along the railroad line near Cardenas, together with his Actaeonella 

(Trochactaeon) aff. gigantea Sowerby, which belongs to 

the herein described new species T. (Mexicotrochactaeon) rossi.

Several specimens were collected by Wade at the 

base of unit 98 of the Cardenas section, which he assigned a stratigraphic position considerably higher 

than unit 80. These specimens resemble the forms from 

unit 69. According to Myers (1968), the Cardenas section shows repetition of beds because of a synclinal 

structure. Wade had not noted this structural complexity. Unit 98 probably has about the same stratigraphic 

position as unit 69.
58 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

The forms from Chiapas that Maldonado-Koerdell 

(1950) assigned to Actaeonella coniformis Bbse, A. occidentalis Bose, A. inconstans Bbse, A. planilateralis 

Bbse, and his new subspecies Actaeonella (Trochactaeon) acutissima punctata belong to a single species 

that differs from T. (Mexicotrochactaeon) coniformis. 

For that species, the name T. (Neocylindrites) punctatus 

Maldonado-Koerdell is proposed (see p. 54).

Occurrence. Mexico, State of San Luis Potosi, Cardenas Formation. Myers (1968, p. 76) listed occurrences 

through his Durania ojauchalensis, Arctostrea aquilerae, 

and Tampsia floriformis zones. The species is represented in collections of the U.S. Geological Survey from 

localities 152, 153, 156, 157.

Age. Maastrichtian.

Types.-Hypotypes, USNM 306045-306062, 

UT(WSA 15554a.

Trochactaeon (Mexicotrochactaeon) burckhardti (Bose)

Plate 12, figures 7-13; plate 13, figures 7-9

Actaeonella (Trochactaeon) gigantea Sowerby sp. n. var. mexicana

Bose, Mexico Inst. Geol. Bol. 42, p. 200, pi. 14, figs. 3-8. 

Actaeonella (Trochactaeon) Burckhardti Bbse, Mexico Inst. Geol.

Bol. 42, p. 202, pi. 14, figs. 9-14, pi. 15, figs. 1-11. t 

Actaeonella (Trochactaeon) Burckhardti Bose n. var. obliqueplica'ta

Bose, Mexico Inst. Geol. Bol. 42, p. 203, pi. 16, figs. 1-6. 

Actaeonella (Trochactaeon) pruniformis Bbse, Mexico Inst. Geol.

Bol. 42, p. 204, pi. 16, figs. 7-15, pi. 17, figs. 1-5. 

Actaeonella (Trochactaeon) Humboldti Bbse, Mexico Inst. Geol.

Bol. 42, p. 205, pi. 17, figs. 6-11. 

Actaeonella (Trochactaeon) sp. Bbse, Mexico Inst. Geol. Bol. 42,

p. 206, pi. 17, figs. 12-14. 

Actaeonella (Trochactaeon) quadriplicata Bose, Mexico Inst.

Geol. Bol. 42, p. 207, pi. 17, figs. 15-17. 

Actaeonella luciano Maury, Brazil Serv. Geol. Min., Mon. 8,

p. 257, pi. 10, figs. 1, 2, 6. 

Actaeonella pompei Maury, Brazil Serv. Geol. Min., Mon. 8,

p. 259, pi. 10, fig. 7. 

Actaeonella cf. silvai Maury, Brazil Serv. Geol. Min., Mon. 8,

p. 263.

Diagnosis. Body whorl strongly inflated in upper 

quarter, lower half tapering to slender base; columella 

proportionally short, parietal lip with a weak ridge on 

lower part.

Description. Shells large, ovate; maximum diameter 46 to 58 percent of total height; spire variable in 

height, consisting of 6-7 whorls, early ones usually 

adapically concave in profile but later ones convex, profile rarely convex as a whole; whorls flat side and with 

a broad wrinkled outwardly inclined subsutural ramp. 

Body whorl 80 to 92 percent of total height, strongly 

inflated in upper quarter, flattened to slightly depressed 

medially, tapering and thinning anteriorly; surface 

with coarse prosocline growth lines becoming opisthocline over basal slope. Aperture elongate, proportionally broad; columella one-third of aperture height; 

columella with upper plait sharply upturned, lower

1923.

1923.

1923.

1923.

1923.

1923.

1923.

1930.

1930.

1930.

two plaits weaker; columella below plaits as long as 

part with plaits; inductura with strongly prosocline 

outer margin that circumscribes columella; parietal lip 

thick, extends onto body whorl; low ridge above columella, thinning out nearly completely in middle within 

aperture, about two times thicker in the upper part 

than near columella; sections show plaits that gradually diminish in strength from first half of body whorl 

to outer half of penultimate whorl; a low palatal ridge 

occurs within penultimate whorl and in first half of 

body whorl; palatal walls of early whorls nearly completely resorbed.

Measurements (fig. 31). The range of measurements 

of specimens studied is: H, 63-98.2 mm; MD, 46-52 

mm; HB, 56-82 mm; PA 65°-85°.

Discussion. The seven "species" figured by Bose 

(1923) from Zumpango del Rio, Mexico, show very well 

the variation within the one species for which we select 

the name Trochactaeon (Mexicotrochactaeon) burckhardti (Bbse); we place his other species in synonymy. 

As indicated on figure 31, the height-width ratios are 

very variable. Unfortunately, the parietal lip seems 

not to be preserved in Bbse's specimens; at least, it is 

not mentioned in his descriptions and is not recognizable on the figures. The general shape of the shells 

their low spire, convex body whorl, broad subsutural 

ramp, and proportionally slender columella is charac110r

100-

90

I1 BO

70

60

50.

"30 40 50 60

MAXIMUM DIAMETER

FIGURE 31. Scatter diagram comparing height with width among 

selected specimens of , Trochactaeon (Mexitrochactaeon) 

burckhardti (Bose). 0, holotype of Actaeonella lucianoi Maury; 

+, holotype of A. pompei Maury.
SYSTEMATIC PALEONTOLOGY 59

teristic of Trochactaeon (Mexicotrochactaeon) burckhardti and is in accordance with the shape of the Caribbean specimens described here and with that of the 

Brazilian Actaeonella lucianoi Maury (1930). Actaeonella pompei Maury (1930) was described from a 

single fragment, which, like A. lucianoi, also belongs to 

T. (Mexicotrochactaeon) burckhardti.

Among European species T. (Trochactaeon) sanctaecrucis (Futterer, 1892) is morphologically similar. This 

widely distributed species has, like T. (Mexicotrochactaeon) burckhardti, a strong thickening in the upper 

part of the parietal lip but differs in the thin lower part 

of the lip (Kollmann, 1967). There is no other significant difference in shell morphology between the two 

species. T. (Trochactaeon) galloprovincialis Cossmann 

(1897) is more convex in the lower part of the body 

whorl than is T. (Trochactaeon) burckhardti, has a 

thicker columella, and possesses a thin parietal lip, as 

far as it can be recognized from the completely recrystallized specimens from the type locality in the Barremian of southern France (Kollmann, 1967).

Occurrence. Mexico: Bb'se's type material came from 

Tenantitlan near Zumpango de Rio in the State of 

Guerrero. Brazil: Maury's specimens came from Fazenda 

da Estiva near Tamandore, Pernambuco. Jamaica: 

Central inlier, Guinea Corn Formation, at localities 

58-60, 64, 67; Maldon inlier, Maldon Limestone, at 

locality 119.

Age. "Emscherian" (probably Coniacian, according 

to Imlay, 1944, p. 1124) in Mexico; Emscherian in 

Brazil, according to Maury, 1930; Maastrichtian in 

Jamaica.

7>pes.-Hypotypes, USNM 306076-306080, 306088- 

306090.

Trochactaeon (Mexicotrochactaeon) wrighti Sohl and Kollmann, n. sp.

Plate 11, figures 8, 9; plate 12, figures 1-6, 10

Diagnosis. A Mexicotrochactaeon with gently convex body whorl sides and an especially narrow elongate aperture.

Description. Shells small to medium size, maximum 

diameter 42-79 percent of height. Spire with slightly 

concave to weakly convex sides; whorl sides convex, 

with a broad, collarlike steeply inclined ramp. Body 

whorl 74 to 83 percent of total height, shouldered in 

the upper fifth, conical to slightly convex below, with a 

shallow inflation in upper half. Growth lines fine. Aperture narrow and long. Columella one third of aperture 

height, median plait weakest; broad incision above columella, which may be carinated above. Parietal lip 

thick on anterior part, thinning out nearly completely 

near middle, thickening again on posterior part; margin of parietal lip nearly vertical on posterior part, 

becoming flatter on anterior third, circumscribing columella about one quarter of a turn and bending downwards, thus terminating columella. Columella usually 

resorbed internally on all whorls except the last; on 

penultimate whorl, only lower parts of palatal area 

resorbed; on earlier whorls, sides nearly completely 

resorbed.

Measurements (fig. 32). The size range of 13 complete specimens is: H, 33.6-57.0 mm; MD, 14.5-28.8 

mm; HB, 26.5-45.0 mm; PA, 38°78°.

Discussion. Trochactaeon (Mexicotrochactaeon) 

wrighti has features that seem similar to those of both 

T. (Neocylindrites) punctatus (Maldonado-Koerdell) 

and T. (Mexicotrochactaeon) coniformis. As do those of 

T. (Neocylindrites) punctatus, the shells possess both 

an especially elongate and narrow aperture and whorls 

that are not very convex below the shoulder. They are, 

however, similar to those of T. (Mexicotrochactaeon) 

coniformis in having an incision above the columella. 

This incision is especially deep and wide in the proportionally broad Jamaican specimen illustrated on plate 

12, figure 10. The less concave early whorls and flat 

sides of the body, however, distinguish the specimen 

from Bo'se's species. Specimens collected from the lower 

part of the Cardenas, Mexico, section (bed 51 of Imlay's, 

1944, section) are closer to typical T. (Mexicotrochac70

I

UJ

n:

60-

50

40-

30

20

10

0 10 20 30 40 

MAXIMUM DIAMETER

FIGURE 32. Scatter diagram comparing height with width among 

selected specimens of Trochactaeon (Mexicotrochactaeon) wrighti 

Sohl and Kollmann, n. sp.
60 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

taeon) coniformis. They possess slightly convex whorl 

sides and a distinct subsutural ramp. The incision 

above the columella is very deep and carinate, abuting 

the parietal lip (pi. 12, fig. 1).

Trochactaeon (Mexicotrochactaeon) wrighti possesses 

a proportionally shorter spire than do most other species of the subgenus. Although some specimens of both 

T. (Mexicotrochactaeon) granthamensis n. sp. and T. 

(Mexicotrochactaeon) rossi n. sp. may have a low spire, 

they are more obese, possessing a very convex-sided 

body whorl.

Occurrence. Mexico: State of San Luis Potosi, Cardenas Formation, at localities 152, 155. Jamaica: Marchmont inlier, "Titanosarcolites limestone," at localities 

93, 99.

Age. Maastrichtian.

7>/?es.-Holotype, USNM 306073; Paratypes, USNM 

306067, 306068, 306070-306072, 306074, 306075.

Trochactaeon (Mexicotrochactaeon) palmeri Sohl and Kollmann, 

n. sp.

Plate 11, figure 5; plate 14, figures 1-5, 8

Diagnosis. A T. (Mexicotrochactaeon) with a proportionally high spire and a parietal ridge that is reduced 

to a flat thickening at the aperture.

Description. Shell of medium size; maximum diameter 45-55 percent of height, spire high, flat to slightly 

convex in profile; whorl sides generally convex with or 

without slight depression below shoulder; aperture 

broad but proportionally short; columella short, middle 

columellar plait weakest; parietal lip thick with curved 

rim; parietal ridge expressed only as a flat thickening at 

aperture, but stronger and with sharper crest inside first 

half of last whorl; palatal ridge low within last whorl; 

in penultimate whorl and earlier whorls, stronger 

resorption of side walls below palatal ridge than above; 

resorption of columella begins within last whorl, columella in earlier whorls smooth or with small round ridge.

Measurements (fig. 33). The size range of the available specimens is: H, 27.4-56.8 mm; MD, 14.1-25.3 mm; 

HB, 19.0-36.0 mm; PA, 30°-55°.

Discussion. This species superficially resembles some 

specimens of Trochactaeon (Mexicotrochactaeon) coniformis (Bb'se) but differs by its proportionally broader 

whorls and the less prominent parietal ridge (compare 

pi. 10, fig. 6, and pi. 14, fig. 4). The columellar plaits 

are resorbed within the last whorl, whereas the resorption in T. (Mexicotrochactaeon) coniformis starts with 

the beginning of the penultimate whorl.

Trechmann (1929) ascribed a specimen from the Blue 

Mountains, Jamaica, to Actaeonella (Trochactaeon) cf. 

variabilis (Bb'se). We have not examined the specimen, 

but the description suggests affinities to T. (Mexicotrochactaeon) palmeri.

60

50

40

1 30

I

20-

10

0 10 20 30 40 

MAXIMUM DIAMETER

FIGURE 33. Scatter diagram comparing height with width among 

selected specimens of Trochactaeon (Mexicotrochactaeon) palmeri Sohl and Kollmann, n. sp.

The species is named in honor of Robert O. Palmer, 

who made many of the collections from the Cretaceous 

of Cuba that provided fossils studied here.

Occurrence. Jamaica: Jerusalem Mountain inlier, 

"Titanosarcolites limestone," at localities 108, 111; 

Central inlier, Guinea Corn Formation, at locality 65. 

Cuba: Santa Clara Province, Habana Formation, at 

localities 124, 126, 128, 129. Puerto Rico: San German 

Quadrangle, fossils in tuffs of the El Rayo Formation, 

at locality 42.

Age. Middle to late Maastrichtian.

Types.-Holotype, USNM 306098; paratypes, USNM 

306063, 306095-306097, 306099, 306100.

Trochactaeon (Mexicotrochactaeon) teaensis Sohl and Kollmann, n. sp.

Plate 13, figures 13, 6, 14; plate 14, figures 6, 7, 9-16

Diagnosis. A Mexicotrochactaeon with a proportionally high spire and a low rounded parietal ridge.

Description. Shells of medium size, maximum diameter 44 to 54 percent of height; spire variable in height, 

convex in outline, whorls with an inclined, often indistinct collarlike ramp; shoulder variable with all transitions between virtual absence to slightly convex. Body 

whorl 66-83 percent of total shell height; sides below 

shoulder convex or with broad flat depression and 

angulated at junction with low columella. Aperture 

relatively broad and of medium height. Columella with
SYSTEMATIC PALEONTOLOGY 61

middle plait weakest, part below plaits as high as distance between plaits; parietal ridge rounded, strongest 

in first half of last whorl, decreasing in strength at 

aperture; palatal ridge strong in first half of last whorl, 

but area below ridge resorbed on penultimate whorl; in 

earlier whorls, resorption occurs also on upper part of 

palatal walls; columella with strong medial ridge in 

resorbed parts.

Measurements (in mm). H, 18.6-66.0; MD, 8.8-30.2; 

HB, 15.5-51; PA, 30 °-60°.

Discussion. Trochnctaeon (Mexicotrochactaeon) 

teaensis is a very variable species; it possesses characters suggesting that it is intermediate in form between 

T. (Mexicotrochactaeon) rossi and T. (Mexicotrochactaeon) wrighti. It is like T. (Mexicotrochactaeon) rossi in 

having evenly convex whorls and a broad, rounded, low 

parietal ridge. It is similar to T. (Mexicotrochactaeon) 

wrighti in its width of whorl and in the character of the 

shoulder that occurs on most specimens; however, the 

whorls are proportionally longer.

Trochactaeon (Mexicotrochactaeon) teaensis occurs in 

rocks of Maastrichtian Age from Puerto Rico to Mexico. It is very abundant in the Tea syncline near San 

German, Puerto Rico.

Occurrence. Puerto Rico: San German Quadrangle, 

limestone lentils within the El Rayo Formation, at 

locality 48. Jamaica: Jerusalem Mountain inlier, 

"Titanosarcolites limestone," at locality 109. Mexico: 

State of San Luis Potosi, Cardenas Formation, at localities 155, 156, 157.

Age. Middle to late Maastrichtian.

7>pes.-Holotype, USNM 306105; paratypes, USNM 

306081-306085, 306101-306104, 306106.

Trochactaeon (Mexicotrochactaeon) granthamensis 

Sohl and Kollman, n. sp.

Plate 13, figures 10-13; plate 15, figures 111

Diagnosis. Variably high spired shell with a columella that is bordered above, first by a deep constriction and then by a ridge; parietal lip thickened in upper 

part.

Description. Shells of small to medium size; maximum diameter 43 to 57 percent of total height; spire 

usually high but variable, slightly concave to convex in 

profile and consisting of 7 to 8 whorls; whorl sides flat 

in small specimens, convex in large ones; all whorls 

with a broad, inflated, but little inclined subsutural 

ramp. Body whorl 62 to 89 percent of total height, convex in outline with greatest diameter in posterior 

third, convex below and tapering towards very narrow 

base. Opisthocyrt growth lines strongly curve backwards over the ramp. Growth lines occasionally develop a narrow sinus over upper part of whorl. Aperture 

broad with a proportionally long columella; uppermost

columellar plait acute and bent upwards; plaits decreasing in size anteriorly; part of columella below plaits 

about half total height; deep incision above columella 

bordered posteriorly by parietal ridge; this ridge extends out from margin of columellar lip but becomes 

more prominent inside shell and is present throughout 

last whorl; thickening on upper part of parietal lip 

prominent. In early growth stage, deep constriction in 

middle of parietal lip; low palatal ridge in penultimate 

whorl at later stage.

Measurements (fig. 34). The largest available, but 

incomplete, specimen, has an estimated height of nearly 

100 mm and a maximum diameter of 51 mm. Most 

commonly, shells range from 60 to 75 mm in height 

and 27 to 38 mm in diameter.

Discussion. This species varies considerably in shell 

proportion. High-spired forms tend to have a more slender outline whereas those having a low spire possess a

90

80

70

60

50

40

30

20

10-

I I J__L

0 10 20 30 40 50 

MAXIMUM DIAMETER

FIGURE 34. Scatter diagram comparing height with width among 

selected specimens of Trochactaeon (Mexicotrochactaeon) granthamensis Sohl and Kollmann, n. sp.
62 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

more obese and rounded body whorl (pi. 15, fig. 1). In 

most larger specimens, the shell is quite thin near the 

aperture; thus, the body whorl gives a wrinkled appearance in part because of crushing and part because 

of accentuation of the growth lines (pi. 15, fig. 7). Of all 

the species dealt with in this work, this species contains the largest number of individuals showing shellrepair phenomena. Although breaks may be common 

at almost any stage, the specimen figured on plate 15, 

figure 8, shows extreme injury. In this specimen, the 

body whorl has been broken back for a distance of almost half a turn and then reformed at an offset position.

Occurrence. Jamaica: Central inlier, Guinea Corn 

Formation, at localities 61, 62, 64, 68, 68a, 69, 76, and 

79; Marchmont inlier, "Titanosarcolites limestone," at 

locality 106.

Age. Middle Maastrichtian.

Types.-Holotype, USNM 306115; paratypes, USNM 

306091-306094, 306108-306114.

Trochactaeon (Mexicotrochactaeon) rossi Sohl and Kollmann, n. sp.

Plate 13, figures 4, 5; plate 16, figures 1-9

1906. Actaeonella (Trochactaeon) aff. gigantea Sowerby. Bose, Inst. 

Geol. Mexico Bol. 24, p. 82, pi. 17, fig. 1.

1968. Actaeonella coniformis Bose. Myers, (in part) Mexico Univ. Nac. 

Autonoma, Inst. Geol., Paleontologia Mexicana, No. 24, p. 75, 

pi. 7, figs.4, ?5: not fig. 3 (T.(Mexicotrochactaeon) coniformis).

Diagnosis. Spire and body whorl equally convex in 

profile, subsutural ramp collarlike; parietal lip with 

broad thickening in lower part.

Description. Small to large elongate oviform shells 

consisting of 8 to 9 fairly convex high whorls with 

strongly inclined collarlike subsutural ramps that 

become indistinct on larger specimens. Maximum 

diameter 37 to 54 percent of total height and occurs at 

middle of shell, which is nearly symmetrically biconvex in large- and medium-sized specimens; in small 

specimens, spire may be slightly concave. Body whorl 

58 to 84 percent of total height; aperture long, slightly 

broader in lower part; columella proportionally long, 

thin, and distinctly separated from convex whorl sides 

above; columellar plaits weak but equal in adult specimens; parietal ridge begins within aperture as rounded 

but distinct thickening of lower third of parietal lip, 

but parietal ridge absent in early growth stages. Columellar plaits decrease in size within body whorl and 

are lost in earlier formed whorls where columella is 

highly resorbed and shows only a foldlike ridge at midheight; palatal ridge present on penultimate whorl and 

on most of body whorl but does not reach aperture.

Measurements (fig. 35). The size range of the many 

specimens studied is: H, 22-86 mm (a large fragment is 

estimated to represent a shell about 130 mm high); 

MD, 10.7-50.2 mm; BW, 19.7-88 mm; PA 34°-60°.

110 t

I

UJ

X

100

90

80

70

60

50

40

30

20

i i r

10 20 30 40 

MAXIMUM DIAMETER

50

FIGURE 35. Scatter diagram comparing height with width among 

selected specimens of Trochactaeon (Mexicotrochactaeon) rossi 

Sohl and Kollmann, n. sp.

Discussion. Shells of this species vary widely in 

their proportions, even specimens from the same locality. Common to all larger specimens is the equal convexity of both the body-whorl sides and the spire and 

also the thickening of the lower part of the parietal lip. 

Collections containing small specimens show that convex spires predominate but that a few specimens may 

have a concave spire profile. As the parietal lip in 

small specimens shows no thickening at all, there is 

some question whether they should be included in this 

species. However, as most of them show the same external shell features, we have included them. We also 

include the specimen that Bb'se assigned to Actaeonella 

(Trochactaeon) aff. gigantea Sowerby because it has an 

evenly rounded last whorl and a spire with convex 

sides, but the spire is lower than in most other specimens. Myers (1968, pi. 17, fig. 4) figured a large specimen that should be included in this species as it shows 

a thickening of the parietal lip. Another specimen fig-
SYSTEMATIC PALEONTOLOGY 63

ured by Myers (1968, pi. 17, fig. 5) is a higher spired 

form, which may belong to the same species. Myers 

assigned both these specimens to Actaeonella coniformis 

Bose, but he did not include Bbse's Actaeonella (Trochactaeon) aff. gigantea Sowerby in his synonymy. This 

suggests that he considered that form to be different 

from all the other "species" of Bose.

Trochactaeon (Mexicotrochactaeon) rossi may be distinguished by the strong convexity of the body whorl 

and by the thick swelling in the upper part of the parietal lip. Other species having a similar high spire may 

be distinguished by their lack of the very broad and 

less inclined subsutural ramp. The shell of T. (Mexicotrochactaeon) granthamensis is not as broad, the 

whorls are narrow, and the parietal ridge is sharper.

The species is named in honor of William O. Ross of 

the U.S. Geological Survey, who assisted in collecting 

and preparation of much of the material upon which 

this study is based.

Occurrence. Mexico: State of San Luis Potosi, Cardenas Formation (upper part), according to Myers (1968). 

Jamaica: Marchmont inlier, "Titanosarcolites Limestone," at localities 99, 102a, 105a; Central inlier, 

Guinea Corn Formation, at localities 73, 74, 78, 82, 84, 

87, 89. Cuba: Questionable occurrence in the Habana 

Formation, Habana Province, at locality 126.

Age. Middle Maastrichtian.

Types. Holotype, USNM 306120; paratypes, USNM 

306086, 306087, 306116-306119.

Trochactaeon (Mexicotrochactaeon) melonensis 

Sohl and Kollman, n. sp.

Plate 16, figures 10-13; plate 17, figures 6-8

Diagnosis. A Mexicotrochactaeon with a high spire 

and an exceptionally developed parietal ridge.

Description. Oviform shells of medium size, maximum diameter about half total height; spire variable, 

mostly high, convex in profile; whorl sides slightly convex, subsutural ramp narrow and inclined. Body whorl 

convex, occasionally indistinctly shouldered; base well 

rounded. Aperture incompletely known, broad, and 

comparatively low; middle of three columellar plaits 

weakest; part of columella below plaits as high as distance between plaits; parietal lip with a broadly expanded edge inclined 35 ° to shell axis; parietal lip thin 

except for a strong round-topped ridge that ends within 

aperture; ridge situated higher on parietal lip in earlier whorls than on last whorl. Sectioned shells show 

that wider shells possess a stronger palatal ridge; side 

walls thick and not resorbed in earlier whorls, but columella is resorbed.

Measurements. Most available specimens are incomplete, but large specimens approach 80 mm in 

height, with a maximum diameter near 40 mm. One

small specimen measures 35 mm in height and 18 mm 

in diameter. The pleural angle of small stages is near 

80° but decreases in latest growth stages to 30°-35°.

Discussion. This species is moderately variable in 

the proportional height of the spire and in whorl outline. In most specimens, the spire height approaches or 

even occasionally exceeds half the total shell length. 

The two specimens figured on plate 16, figs. 11 and 13, 

adequately express the range of body-whorl convexity, 

one being well rounded, whereas the other approaches 

a cylindrical upper whorl profile. This species resembles 

T. (Mexicotrochactaeon) rossi in its general appearance, 

but its body whorl is more evenly rounded, broader, 

and proportionally not as high. T. (Mexicotrochactaeon) 

melonensis has an extraordinarily strong parietal 

ridge, which distinguishes it from other species that 

possess broad whorls. T. (Trochactaeon) obtusus 

(Zekeli), which has been recorded from several localities 

in Austria and Yugoslavia (Kollmann, 1967), is similar 

in its overall shell form but lacks a parietal ridge and 

therefore belongs to the subgenus T. (Trochactaeon).

Occurrence. Puerto Rico: Melones Limestone, Cabo 

Rojo Quadrangle, at locality 13, and Puerto Real Quadrangle, at localities 31 and 33.

Age. Early Maastrichtian.

Types.-Holotype, USNM 306123; paratype, USNM 

306121, 306122, 306129, 306130.

Trochactaeon (Mexicotrochactaeon) spp.

Discussion. Specifically indeterminate specimens 

occur at many localities. Most probably belong to species already described here, but their state of preservation precludes precise placement. They are included 

here, as many are the only record of the genus in certain Maastrichtian rock units.

Occurrence. Puerto Rico: Melones Limestone, Cabo 

Rojo Quadrangle, at locality 14; El Rayo Formation, 

San German Quadrangle, at locality 50; Cotui Limestone, San German Quadrangle, at locality 46.

Jamaica: Guinea Corn Formation, Central inlier, at 

localities 54-56, 62-64, 66, 68, 72, 77, 81, 82, 85, 88, 

90, and 91; Titanosarco/ites-bearing limestone, Marchmont inlier, at localities 93, 96, 97, and 106, Jerusalem 

Mountain inlier, at localities 110, 111, 114, and 115; 

Maldon Limestone, Maldon inlier, at localities 118,119; 

Vaughansfield Limestone, Maldon inlier, at locality 117.

Genus Actaeonella d'Orbigny, 1842 

(Synonym Volvulina Stoliczka, 1865)

Type by subsequent designation (Hermannsen, 1846) 

Volvaria laevis Sowerby.

Diagnosis. With the addition that rarely only two 

columellar plications may be present, Meek's (1863, 

p. 89) characterization of the genus remains pertinent:
64 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Shell ovate-volvuliform, rather thick, involute, more or less attenuate above, widest below the middle entirely without any traces of a 

spire. Surface nearly smooth. Aperture very narrow, arcuate, and 

equalling the greatest length of the shell. Outer lip smooth, generally rather obtuse. Inner lip thickened near the base of the aperture, 

and twisted outwards so as to form on the columella three prominent 

revolving folds; also usually a little thickened at the summit of the 

aperture.

Discussion. D'Orbigny (1842) in describing the 

genus Actaeonella failed to designate a type species. 

The first designation of a type species was that of Hermannsen, who in 1846 selected Volvaria laevis J. de C. 

Sowerby from the Santonian to Campanian of the eastern Alps. Meek (1863) subsequently cited the same 

species as type but failed to cite Hermannsen as first 

subsequent designator; however, he did give a full reference to Sowerby's work. From this point onward, 

various errors in citation of the type species entered 

into the synonymic history. The first major departure 

was Cossmann's (1895) designation of Actaeonella laevis 

d'Orbigny (1842), not Sowerby, as type. D'Orbigny's 

species, based upon specimens from the Turonian of 

Uchaux, France, are distinct from Sowerby's species. 

Evidently, Cossmann was ignorant of the existence of 

Sowerby's figures of V. laevis, as he neither cited or discussed them but proposed the name A. terebrellum for 

the Alpine species. In the following year, Cossmann 

(1896) cited A. laevis (Sowerby) as a valid name and 

placed his A. terebrellum in synonymy. At the same 

time, he recognized the separation of d'Orbigny's A. 

laevis from Sowerby's species by erecting a new species, A. uchauxensis. In a separate paper, read in April 

1896 but not published until 1896, he used d'Orbigny's 

A. laevis as the basis for description of yet another species, A. grossouvrei. Thus, to this point, the synonymy 

of these two species is as follows:

Sowerby's laevis 

1832 Volvaria laevis Sowerby 

1846 Actaeonella laevis Hermannsen 

1863 Actaeonella laevis Meek

1895 Acteonella terebrellum Cossman

1896 Acteonella laevis Cossmann

d'Orbigny's laevis 

1850 Acteonella laevis (Sowerby) d'Orbigny

1895 Acteonella uchauxensis Cossmann

1896 Acteonella grossouvrei Cossmann.

Cossmann and several subsequent authors ignored 

Hermannsen's and Meek's earlier designations and 

continued to cite A. uchauxensis as the type of the 

genus. This stand has been supported by Nuttall (1969) 

and Nuttall and Leong (1972). In his request to the 

International Commission on Zoological Nomenclature, Nuttall (1969) argued that compilations such as 

that of Zilch (1959) and Pchelintsev and Korobkov 

(1960) cited Acteonella uchauxensis Cossmann as the

type species. Subsequently, Nuttall and Leong (1972) 

cited Article 70 of the Code, which deals with misidentified type species, as support for the original Nuttall 

proposal.

In contrast to Nuttall's opinion, we see no reason to 

reject Hermannsen's (1846) explicit designation of 

Actaeonella laevis (Sowerby) as the type species. That 

Hermannsen did not cite the plate and figures provided 

by Sowerby is no reason to believe that he meant Actaeonella laevis d'Orbigny (= A. uchauxensis) to be considered the type species, and certainly no proof, as Nuttall 

believes, that Hermannsen misidentified d'Orbigny's 

species. Rather, it seems clear that if misidentification 

is the charge, it is Cossmann who must bear the stigma. 

In 1895, when he designated d'Orbigny's species, 

which later he named A. uchauxensis, Cossmann was 

certainly not aware that A. laevis (Sowerby) existed 

and had been previously designated as the type species 

of Actaeonella. The fact that such subsequent compilers 

as Zilch and Pchelintsev and Korobkov have in error 

followed Cossmann's designation does not seem a compelling reason to ignore the primary literature and perpetuate misinterpretation of the record. Obviously, 

subsequent compilers have followed Cossmann's 1895 

designation without independently searching the literature. In other type designations, such as those for 

Euchelus Phillipi, Calliostoma Swainson, and Rimella 

Agassiz, they have accepted Hermannsen as a source. 

Thus, if Hermannsen cannot be ignored as a source of 

type designation for these genera, he cannot be ignored 

for designation of the type species of Actaeonella.

Attempts have been made to subdivide Actaeonella. 

The most diverse subdivision is that proposed by Djalilov (1972), who suggested the following taxa: 

Subgenus Sogdianella (Type species: Actaeonella supernata Pchelintsev, 1953; Turonian; Transcaucasus,

USSR). 

Subgenus Pchelincevella (Type species: Actaeonella

caucasica Zekeli, 1852; Coniacian, Transcaucasus,

USSR). 

Genus Ovactaeonella (Type species: Ovactaeonella ferganica Djalilov, 1972, Coniacian; Fergana, Kuraminskiy Range, USSR).

In addition to the above and in the same year, Hacobjan (1972) proposed the genus Omphaloactaeonella 

(type species Actaeonella ovata Pchelintsev, 1953; Turonian, Armenia).

Kollmann and Sohl (1980) have presented evidence 

for the inclusion of Sogdianella in the family Itieriidae 

of the Nerineacea, rather than the Actaeonellidae. Of 

the remander, Ovactaeonella and Omphaloactaeonella, 

both proposed in 1972, seem to represent either the 

same or an overlapping generic concept. Djalilov and 

Hacobjan included virtually the same species in these
SYSTEMATIC PALEONTOLOGY 65

two genera, but in subsequent publications have maintained the independent generic names. In both genera, 

however, the included species possess the hollow columella and central pillar of the Itieriidae rather than 

the Actaeonellidae. Thus, we are not considering them 

further, and they are herewith transferred out of the 

Actaeonellidae in the accompanying species catalogue.

The only remaining taxon is Pchelinseuella Djalilov 

(1972, p. 22) which is "distinguished from the nominate subgenus by the amygdaloid, squatter shell" 

(translation). The distinction of this subgenus seems 

vague to us, of no practical value, and is not accepted 

here.

The evolution, distribution, and life habits of Actaeonella are discussed elsewhere in this paper. Some features of shell morphology, not elsewhere touched upon, 

need special mention. The first is external sculpture. 

The vast majority of actaeonellids possess a smooth 

surface marked only by fine growth lines. A few species 

such as Actaeonella decorata Hacobjan, A. ornata 

Pchelintsev, and A. khalilovi Aliev from the Cenomanian of the Armenian SSR, show spiral sculpture consisting either of incised lines or raised coarse spiral 

cords interrupted by transverse swellings that lend a 

spirally elongate noded appearance to the cords.

Shells of Actaeonella are seldom bored. One of the 

few exceptions found in our material is figured on plate 

21, figure 15. Likewise, lack of epibiont encrustation 

and lack of evidence of shell breakage and repair support the interpretation of an infaunal habit.

Internally, Actaeonella lacks evidence of shell-wall 

thinning and rarely shows secondary shell deposition 

in parts of the shell that is so common in Trochactaeon 

shells. One especially unusual character shown by two 

species described here (A. robinsoni and A. cubensis) is 

the presence of only two columellar plications. We 

know of no other species possessing this feature.

Actaeonella pecosensis Stanton

Plate 18, figures 9-14; plate 20, figure 10

1947. Actaeonella pecosensis Stanton, U.S. Geol. Survey Paper 211, 

p. 110, pi. 67, figs. 2, 5, 6, 8, 9.

Diagnosis. A high slender Actaeonella with evenly 

convex sides and acute posterior tip; columella with an 

additional weak fourth plait below three primary plaits.

Description. "Shell of moderate size, slender fusiform, with the spire entirely concealed and the aperture extending the full length of the shell; greatest 

breadth of shell a little in advance of the middle, from 

which the shell gradually tapers to the acute posterior 

end which has an apical angle of about 30 °; anterior end 

broader, more abruptly narrowed, and obliquely truncate or slightly emarginate in front; surface smooth, 

not even showing growth lines on the best preserved

specimens studied; inner lip thickened toward the 

front and bearing three strong spiral folds of which the 

posterior fold is most prominent. The elevation on 

which the folds are borne is continued as a narrower 

rounded ridge on the back of the last whorl where it 

borders the anterior end of the shell. The aperture is 

very narrow, its widest part being near the columellar 

folds" (Stanton, 1947, p. 110).

Additional description. Maximum diameter 29-37 

percent of total height. Columella proportionally long, 

in late growth stages bears three plaits of about equal 

strength, with a fourth very weak plait very close but 

anterior to the third plait; columella below three upper 

plaits slightly more inclined towards shell axis than 

above; lateral edge of columella thickened, inclined 

about 40 ° toward axis, upper two plaits extend towards 

this edge, but lower ones end well before edge. Shell 

base unknown.

Measurements (fig. 36).-The holotype (USNM 77560) 

is 43 mm high and has a maximum diameter of 14 mm. 

Other specimens reach a maximum height of 60.5 mm 

and have a maximum diameter of 20.5 mm. The pleural 

angle of all specimens ranges from 32 ° to 45 °.

Discussion. Cut sections (pi. 20, fig. 10; pi. 18, fig. 14) 

show that the proportional maximum shell diameter is

70

g

LLJ

X

60-

50

40

30

20 h

10

0 10 20 30 40 

MAXIMUM DIAMETER

FIGURE 36. Scatter diagram comparing height with width among 

selected specimens of Actaeonella pecosensis Stanton.
66 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

greater in small growth stages (H/D=2.5 to 3). In smaller 

specimens, the columellar length is also proportionally 

greater than in larger specimens. No development of 

the weaker fourth columellar plait seems to have taken 

place in these early growth stages. Thus, in early stages, 

individuals can hardly be distinguished from Actaeonella delgadoi Choffat. Larger specimens of A. delgadoi 

have a higher maximum diameter than large specimens of A. pecosensis and also a more obtuse posterior 

tip.

The species is abundant in the upper part of the Devils 

River Limestone of the margins of the Maverick Basin 

in Texas. Stanton (1947, p. 110) noted that it is a common associate of Nerinea riograndensis.

Occurrence. Texas:Devils River Limestone (upper 

part), at localities 134,139, and Edwards Limestone, at 

localities 132, 146.

Age. Middle to late Albian.

Types. Holotype, USNM, 77560; paratypes, USNM 

77561a-b, 103173; hypotypes, USNM 306138, 306139, 

UT1036 TX6.

Actaeonella delgadoi Choffat, 1901

Plate 18, figures 17 20

1901. Actaeonella delgadoi Choffat, Faune cretacique du Portugal, v. 1,

p. 109, pi. 1, figs. 1-6. 

1916. Actaeonella gracilis Douville, Acad. Sci. Inst. France Mem.,

ser. 2, v. 54, no. 1, p. 147, pi. 18, fig. 34. 

1955. Acteonella gracilis Douville. Mahmoud, Inst. Desert Egypte Pub. 8,

p. 155, pi. 17, figs. 29-31. 

1963. Actaeonella gracilis Douville. Abbass, [Cairo] Geol. Mus., Palaeont.

Ser., Mon. 2, p. 105. 

1972. Acteonella grossouvrei Cossmann. Collignon, Morocco Serv. Geol.

Notes et Me'm. 228, p. 18, pi. 3, figs. 12-14. 

1972. Actaeonella (Pchelincevella) delgadoi Choffat. Djalilov, Paleont.

Zhur. no. 1, p. 14. 

1977. Acteonella (Pchelincevella) delgadoi Choffat. Djalilov, Akad.

Nauk Tadzhakstan SSR, p. 116.

Description. Shells small, maximum diameter 38 to 

41 percent of total height; sides subcylindrical, slightly 

convex with a pleural angle of 75° to 90°; basal part 

rounded; aperture narrow; columella proportionally 

long, about 20 percent of total height, very thick and 

separated from parietal region by a deep incision; three 

plaits of columella decrease in strength anteriorly, 

upper and middle plaits concave on their upper surface; occasionally, a faint fourth plait appears below 

anteriormost one, which is only visible between half 

and three-quarters of a whorl behind aperture. Laterally, edge of columella bends strongly downward; edge 

either reflected or elevated; upper two plaits end at 

this rim, lower plait ends somewhat before edge.

Measurements (fig. 37). Available specimens range 

from 21.6 to 29.2 mm in height and 8.2 to 14.2 mm in 

maximum diameter.

Discussion. Actaeonella sublaevis d'Orbigny has a

60

50

40

30

20

10

0 10 20 30 40 

MAXIMUM DIAMETER

FIGURE 37. Scatter diagram comparing height with width among 

selected specimens of Actaeonella delgadoi Choffat.

columellar form similar to that of A. delgadoi, but the 

profile of the upper part of the shell is strongly concave. 

In addition, small shells of A. sublaevis are inflated 

anteriorly and drawn out to an acute posterior tip (see 

Briart and Cornet, 1865), but large specimens have a 

more cylindrical shape. A. pecosensis Stanton differs in 

possessing a more acute apical tip, a shell that is constantly convex throughout with only a small medial 

constriction, and a proportionally shorter columella.

Occurrence. Texas: Glen Rose Formation, "Thorp 

Springs Limestone Member," at locality 145, and from 

an unspecified locality in the Walnut Formation (specimens from the USNM Renfro Collection). Also, Portugal, Egypt (Massif de Moghara), and Madagascar.

Age. Early and middle Albian in Texas, but Albian 

to Cenomanian elsewhere.

Types.-Hypotypes, USNM 306140-306143.

Actaeonella cf. A. sublaevis Briart and Cornet

1865.

Plate 18, figures 21, 22 

Actaeonella sublaevis Briart and Cornet, Acad. Roy. Sci., Lettres,

et Beaux Arts Belgique Mem., v. 34, p. 42, pi. 3, figs. 15-16. 

1939. Actaeonella (Volvaria) sublaevis Briart and Cornet. Marliere,

Roy. Hist. Nat. Belgique Mem. 89, p. 147, pi. 8, fig. 10a, b. 

1970. Actaeonella delgadoi Choffat. Wolff, Univ. Munich, Disiert.,

p. 142, pi. 7, figs. 6-7. 

1976. Actaeonella sublaevis Briart and Cornet. Kollmann and Yochelson,

Naturhist. Mus., Wien Annal., v. 80, p. 186, pi. 6, figs. 55-57.
SYSTEMATIC PALEONTOLOGY 67

Description. Shell large, lower part with weakly 

convex sides. At about two-thirds of total shell height 

as measured from base, sides turn abruptly inward 

towards axis; because inclination toward axis lessens 

higher on shell, whorl sides below posterior tip are 

strongly concave. Columella is proportionally high, 

upper two of three plaits are sharp and bent upwards, 

lowermost plait is rounded on top.

Discussion. Only fragments and steinkerns with 

partly preserved shells of this large Actaeonella are 

present in our collections. Because of the poor state of 

preservation of all available material, assignment to 

Actaeonella sublaevis is tentative and affords no basis 

for further speculation. The subcylindrical lower part 

of the shell and the strongly concave sides of the upper 

part (pi. 18, fig. 21) are features found in large specimens of Actaeonella sublaevis, especially the specimen 

illustrated by Marliere (1939) and Kollmann (1976). 

This specimen has a total height of 38 mm, which is 

only one-third that of our largest specimen. Actaeonella delgadoi differs in having evenly convex whorls 

but is closely related to Actaeonella sublaevis.

Occurrence. Belgium: Meule de Bracquegnies. Austria: the species has been found at several localities in 

the eastern Alps (Wolff, 1970; Kollmann, 1967, 1976). 

Texas: Buda Limestone, at localities 133 and 144; Edwards Limestone, at locality 146.

Age. middle Albian to early Cenomanian.

Types.-Hypotypes, USNM 306144, UT1036 TX5.

Actaeonella browni Sohl and Kollmann, n. sp.

Plate 18, figures 1-8

Diagnosis. A slender Actaeonella whose greatest 

whorl diameter lies at about two-thirds the total height.

Description. Shells of medium to small size. Shell 

profile convex over lower surfaces, apical tip acute 

with posterior whorl surface flat to gently convex; maximum diameter at about upper two-thirds point, and 

equals about one-third of total height. Aperture elongate narrow, incision lacking between parietal lip and 

very thick columella. Columella bears three plaits of 

equal strength, area anterior to plaits low, with narrow basal excavation.

Measurements (fig. 38). Available specimens reach a 

maximum length of at least 35 mm, but one incomplete 

specimen (pi. 18, fig. 5) indicates that some individuals 

may have been as long as 50 mm and almost 20 mm wide.

Discussion. This species is represented by abundant 

silicified specimens from the type locality. None of 

them, however, possesses a complete aperture. The 

available specimens show a minimum variation in 

shape and, similarly, only a very weak change in proportions during growth. Early growth stages show a

60 r

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MAXIMUM DIAMETER

FIGURE 38. Scatter diagram comparing height with width among 

selected specimens of Actaeonella browni Sohl and Kollmann, 

n. sp.

slight tendency toward greater obesity than do later 

stages, but the amount is hardly noticeable.

Actaeonella browni is morphologically very similar 

to both A. sublaevis Briart and Cornet and A. delgadoi 

(pi. 18). All possess a noticeably proportionally thick 

and well-developed columella. A. browni differs from 

A. delgadoi in its proportionally more slender shell 

that has a less convex whorl profile and a maximum 

diameter that is higher on the whorl. Compared with 

A. sublaevis from the Albian to Cenomanian of Belgium and Austria, A. browni differs in having a zone of 

maximum diameter higher on the whorl and in having 

flat to slightly convex whorl sides above this zone to 

the apex rather than having a concave profile in the 

early stages of growth as in that species.

The species is named in honor of the collector of the 

material, Barnum Brown.

Occurrence. Guatemala: Peten area, at locality 160.

Age. Late Albian (or, less likely, early Cenomanian).

Types. Holotype, USNM 306131; paratypes, USNM 

306132-306137.

Actaeonella borneensis Nuttall and Leong

Plate 19, figures 16-21; plate 20, figure 11

1959. Actaeonella cf. A. laevis d'Orbigny, Chubb. Am. Assoc. Petroleum

Geologists Bull, v. 43, p. 752. 

1972. Acteonella borneensis Nuttall and Leong, Malaysia Geol.

Survey Geol. Papers, v. 1, p. 3, pi. 1, figs. A-E.
68 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Diagnosis. An Actaeonella of accentuated elongate 

droplike outline.

Description. Medium-sized shells of very elongate 

droplike outline, maximum diameter at about onethird total shell height. Body whorl convex in profile to 

about midwhorl; posterior slope straight to slightly 

convex, terminating in acute apical tip. Whorls are 

very narrow internally. Columella with three plaits, 

posteriormost one most prominent.

Measurements (fig. 39). The type specimens reach a 

maximum length of about 110 mm and a maximum 

diameter of about 40 mm.

Discussion. In. most regards, Actaeonella borneensis 

is similar to A. teaensis Sohl and Kollmann; however, 

in section, the maximum diameter is at the height of 

the protoconch or midwhorl rather than below. In addition, at all growth stages of A. borneensis, the whorls 

are slightly convex and nearly cylindrical over the 

anterior half of the whorl. As in the early growth stage 

of A. teaensis, the anterior half of the whorl is less cylindrical, expanding rapidly in width from the base to 

point of greatest diameter. In later growth stages, A. 

teaensis also possesses cylindrical whorls, but the

110

100

90

80

70

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10 20 30 40 

MAXIMUM DIAMETER

50

FIGURE 39. Scatter diagram comparing height with width among 

selected specimens of Actaeonella borneensis Nuttall and 

Leong. A, Borneo; +, Cuba; , Mexico.

upper whorl slopes are concave rather than generally 

convex as in A. borneensis.

Occurrence. Borneo: Sungai Malua, Sabah. Cuba: 

Santa Clara Province, at locality 130. Mexico: State of 

Chiapas, upper Rancho Nuevo Beds, at localities 149, 

150.

Age. The type material from Borneo is from an uncertain Cenomanian to Campanian level, according to 

Nuttall and Leong. In Mexico, Chubb (1959) suggested 

a Campanian Age, but the associated faunas would 

probably now be considered early Maastrichtian. In 

Cuba, the specimens are in an imprecisely dated sequence of Upper Cretaceous rocks. In summary, the 

best dating suggests a Campanian to Maastrichtian 

Age for the species.

Types.-Hypotypes, USNM 306156-306161.

Actaeonella oviformis Gabb

Plate 18, figures 15, 16

1869. Actaeonella oviformis Gabb, California Geol. Survey,

p. 173, pi. 28, fig. 58.

1926. Acteonella oviformis Gabb. Stewart, Acad. Nat. Sci. Philadelphia, Proc., v. 78, p. 432, pi. 21, fig. 13. 

Not: 1877. Acteonella oviformis Gabb, Acad. Nat. Sci. Philadelphia Jour., 2d ser., v. 8, p. 281, pi. 40, fig. 6 (= T. 

(Trochactaeon) or T. (Mexicotrochactaeon)). 

Not: 1954. Acteonella oviformis Gabb. Popenoe, California

Div. Mines Bull. 170, p. 18, fig. 2 (= T. (Trochactaeon)). 

Not: 1958. Acteonella oviformis Gabb. Anderson, Geol. Soc.

America Mem. 71, p. 160, pi. 29, fig. 6 (= T. (Trochactaeon) packardi).

Discussion. Only the holotype of this species is 

known, and it is so poorly preserved that it has been 

variously interpreted. The specimen is a fragment of a 

medium-sized Actaeonella with broadly convex sides; it 

measures 42.5 mm in height and 21.6 mm in diameter. 

The whorls bear distinct orthocline growth lines, and 

the lower part of the whorls is considerably expanded. 

There are three columellar plaits, the lower one being 

very flat. The presence of orthocline growth lines ally 

the shell to Actaeonella, whereas, as discussed elsewhere herein, the forms described by Anderson (1902, 

1958) and Packard (1916,1922) as Actaeonella oviformis 

belong to the genus Trochactaeon.

The incomplete specimen that Gabb (1871) described 

from Peru as Actaeonella oviformis is either a T. (Trochactaeon) or a T. (Mexicotrochactaeon).

Occurrence. Cotton wood Creek, Shasta County, 

Calif.

Age. Chico Formation (Ceriomanian-Turonian). The 

stratigraphic position given by Anderson (1958) is not 

valid, as it refers to another species in the middle 

Turonian.

Type.-Holotype, ANSP 4323.
SYSTEMATIC PALEONTOLOGY 69

Actaeonella duckettsensis Sohl and Kollman, n. sp.

Plate 19, figures 1-8; plate 20, figures 3, 6

Diagnosis. An Actaeonella of accentuated slender 

cylindrical form.

Description. Small to medium-size subcylindrical 

shells, maximum diameter 27 to 36 percent of total 

height; tip obtuse, whorl sides convex; growth lines 

faint, slightly prosocyrt in lower half of shell, but orthocline above. Aperture long and narrow; columella 

thin, inductura meets columella outside of aperture at 

an acute angle, exterior edge of columella elevated or 

slightly reflected; columella prolonged, and excavated 

over its full width, grading below into a snoutlike basal 

lip.

Measurements (fig. 40). Complete specimens are as 

much as 47 mm high and 13 mm in diameter. The measurements of several incomplete specimens suggest 

that maximum heights may be slightly greater than 

50 mm.

Discussion. This species may be distinguished by its 

high subcylindrical shape, its obtuse apical tip, thin 

columella, and the snoutlike extension of the base 

(pi. 19, fig. 1). Cut and polished sections (pi. 20, figs. 3 

and 6) show that the protoconch is located well above 

the midpoint of the body whorl. In this regard, Actaeonella duckettsensis is somewhat similar to A. robinsoni. The latter species, however, has a more rounded

60

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MAXIMUM DIAMETER

FIGURE 40. Scatter diagram comparing height with width among 

selected specimens of Actaeonella duckettsensis Sohl and 

Kollmann, n. sp.

anterior whorl profile, proportionally broader shell, 

and concave posterior whorl slope. No other species 

considered here approaches the relative protoconch 

position of A. duckettsensis.

A rock sample from the Rosario Mine, Comayagua, 

Honduras, in the collections of the Naturhistorisches 

Museum, Basel, Switzerland, contains an Actaeonella 

that probably belongs to this species.

Occurrence. Jamaica: Green Island inlier, "Barrettia Limestone," at locality 121; Central inlier, Guinea 

Corn Formation, at localities 61, 62, 64, 75, and Slippery Rock Formation, at locality 53; Marchmont inlier, 

"Titanosarcolites Limestone," at localities 97, 106, 

107.

Age. Possibly latest Campanian, definitely early 

and middle Maastrichtian.

7>pe.-Holotype, USNM 306150; paratypes, USNM 

306145-306149, 306165.

Actaeonella jicarensis Sohl and Kollmann, n. sp.

Plate 19, figures 9 15; plate 20, figure 7

Diagnosis. Actaeonella with flattened whorl sides 

between point of maximum diameter and apical tip.

Descriptions. Small shells; greatest thickness 

slightly below midwhorl, maximum diameter 31-41 

percent of total height, whorl sides flat between area of 

maximum width and acute apical tip; whorl subcylindrical below point of maximum diameter, slightly convex and rounding down to base. Aperture narrow, but 

expanding to a trumpetlike exhalant siphon posteriorly and slightly widened anteriorly; columella very 

short with two weak plaits at aperture that strengthen 

interiorly; area below plaits short and inclined about 

40 ° to shell axis within aperture; plaits vanish outside 

aperture; columella passes into a basal lip that is 

slightly varicose externally, broadly excavated, and 

forms an angle of 115 ° with shell axis.

Measurements (fig. 41). Specimens range from 12 to 

near 30 mm in height and from 4 to about 11 mm in 

diameter; pleural angle ranges from 30 ° to 40 °.

Discussion. Large suites of specimens from the type 

locality in the Sabana Grande Quadrangle of Puerto 

Rico show the species to possess only minor variation 

in form. In all specimens, the columellar plications are 

very weakly developed at the aperture. Occasionally 

these are so weak that the anteriormost is virtually 

obliterated, save for a faint swelling. The normally flat 

profile of the posterior whorl surface occasionally may 

become faintly concave.

The combination of the presence of only two columellar plications, maximum diameter below midwhorl, and a flat posterior whorl profile serve to 

distinguish this species from all others in the Western 

Hemisphere.
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CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

40 

0 10 20 

MAXIMUM DIAMETER

FIGURE 41. Scatter diagram comparing height with width among 

selected specimens of Actaeonella jicarensis Sohl and 

Kollmann, n. sp.

Occurrence Puerto Rico: Sabana Grande Quadrangle, limestones of the El Rayo Formation, at locality 36.

Age. Middle Maastrichtian.

Types.-Holotype, USNM 306154; paratypes USNM 

306151-306153, 305155.

Actaeonella coquiensis Sohl and Kollman n. sp.

Plate 20, figures 1, 2, 5, 8; plate 21, figures 7-12, 15

Description. Small to very small shells, maximum 

diameter at the midshell and constitutes 27-36 percent 

of total height. Sides of body whorl strongly concave 

above midheight, ending in an acute apial tip; on early 

whorls, diameter diminishes rapidly below middle, giving shell a biconical appearance; in late stages of 

growth, shell nearly cylindrical below middle but 

rounding down near base. Aperture narrow, columella 

with three plaits, lowermost of which is weakest 

within aperture on early whorls but becomes of equal 

strength near aperture; columella tapers as it grades 

into basal lip; upper columellar plait gradually 

becomes an external thickening of basal lip; lower 

plaits end at columellar margin.

Measurements (fig. 42). Specimens reach a maximum height of more than 50 mm, but most commonly 

are nearer 30 mm. In diameter, the largest specimen

30

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10

10 20

MAXIMUM DIAMETER

FIGURE 42. Scatter diagram comparing height with width among 

selected specimens of Actaeonella coquiensis Sohl and 

Kollmann, n. sp.

measures 22 mm, but most specimens seldom exceed 

12 mm. The pleural angle ranges from 25 °to 35 °.

Discussion. Actaeonella elongata Kollmann (1965) 

from the Gosau area, Austria, has a similar form and 

ontogeny. It differs in having stronger columellar 

plaits and a maximum diameter that lies well below 

the middle of the shell. Actaeonella laevis (Sowerby) is 

similar in possessing weak columellar plaits but differs 

in its regular whorl convexity throughout the total 

shell length.

Actaeonella coquiensis is abundant at the type locality in shales of the Coamo Formation (of Berryhill and 

Glover, 1960) at Coqui Hill near Salinas in southcentral Puerto Rico. Here, specimens in a shale unit 

low in the section are small (20-30 mm long) but show 

an average size increase higher in the same section, 

where some large incomplete specimens are judged to 

have attained a length of about 60 mm. We tentatively 

assign to this species, abundant specimens occurring 

with a diverse molluscan assemblage in shales of the 

Maldon Limestone at Shaw Castle, Maldon inlier, 

Jamaica. The small size and variable preservation of
SYSTEMATIC PALEONTOLOGY 71

these specimens leaves some question as to their 

assignment, but morphologically they seem closest to 

Actaeonella coquiensis.

Both of the above localities are from approximately 

the same Maastrichtian stratigraphic level. The specimens herein assigned to this species from immediately 

above the Barrettia Limestone in the Stapleton Formation of the Sunderland inlier, Jamaica, are from a lower 

stratigraphic position associated with Torreyites and 

Barrettia gigas. This association most often suggests an 

early Maastrichtian Age, but a latest Campanian Age 

cannot be discounted.

Occurrence. Puerto Rico: Coamo Formation, Central Aguirre Quadrangle, at localities 16, 18. Jamaica: 

Maldon inlier, Maldon Limestone, at locality 119; Sunderland inlier, Stapleton Formation, at locality 122; 

Marchmont inlier, "Titanosarcolites Limestone," at 

locality 101.

Age. Latest Campanian(?)-early to middle Maastrichtian.

7>pes.-Holotype, USNM 306188; paratypes, USNM 

306162-306164, 306186, 306187, 306189-306192, 

306204.

Actaeonella cf. A. coquiensis Sohl and Kollman

Plate 2, figure 6; plate 20, figure 9; plate 21, figures 13, 19-23

Discussion. Many medium-sized specimens of 

Actaeonella from a few localities in both Cuba and 

Puerto Rico cannot be placed with surety but do show 

similarity to A. coquiensis. In the early stages of 

growth, the lower part of the whorls increases in width 

from the base toward the midheight, which lies below 

the level of the protoconch. In later growth stages the 

lower part of the shell becomes increasingly cylindrical. The upper or posterior part of the shell has flat or 

slightly concave sides that slope gradually to the apical 

tip. The aperture is narrow and elongate with a proportionally long columella that bears three plaits, of 

which the upper one is the strongest. Below the plaits, 

the columella is strongly concave. The available 

specimens have a maximum height of 66 mm a maximum diameter of 30 mm. The pleural angle ranges 

from 30° to 45°

All the larger free specimens are imcomplete, lacking the apical parts of the shell. Thus, it seems prudent 

to await collection of better preserved material before 

final assignment is made.

Specimens of Actaeonella coquiensis are similar in 

showing a parallel ontogenetic change in whorl shape 

with increased size.

Occurrence. Puerto Rico: Melones Limestone, Puerto Real Quadrangle, at localities 31, 32. Cuba, Santa 

Clara Province, at locality 127.

Age. Middle Maastrichtian in Puerto Rico and of

uncertain but presumably the same age in Cuba.

Types. Hypotypes, USNM 305600, 306167, 

306193-306198.

Actaeonella marchmontensis Sohl and Kollmann, n. sp.

Plate 21, figures 1-6

Description. Shell generally small but rarely 

attaining medium size; whorl sides evenly but only 

moderately convex, apex obtuse; maximum diameter 

34-52 percent of total height. Aperture very narrow, 

with a short columella of medium thickness; uppermost columellar plait stronger than two lower ones; 

middle plait closer to lower than to upper one; uppermost plait borders columella at junction with parietal 

region and disappears when edge bends sharply down 

towards base; middle plait grades into basal lip; area 

below middle plait becomes progressively more inclined relative to axis within aperture; lowermost plait 

diminishing in strength, disappearing before columella grades into excavated basal lip and not visible at 

aperture.

Measurements (fig. 43). Our specimens range from 

3.2 to 38 mm in height and have a maximum diameter 

of 1.6 to 4.3 mm. The pleural angle ranges from 40° to 64°.

Discussion. This species differs from Actaeonella 

briggsi by its flattened area below the middle coluLU

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MAXIMUM DIAMETER

FIGURE 43. Scatter diagram comparing height with width among 

selected specimens of Actaeonella marchmontensis Sohl and 

Kollmann, n. sp.
72 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

mellar plait, the less rounded whorl sides, and more 

slender form.

Actaeonella laevis (Sowerby) has an obtuse tip like 

Actaeonella marchmontensis, but it attains a larger 

size. The maximum diameter of A. laevis is less than 30 

percent of the total height. The columella is short, like 

that of A. marchmontensis, but the area below the middle plait is not flattened within the aperture.

Actaeonella marchmontensis is the most common 

species in the Titanosarcolites-bearing limestones and 

shales of Jamaica. It occures within the same sections 

as A. duckettsensis but may be differentiated by its less 

cylindrical form.

Occurrence. Jamaica: Marchmont inlier, from 

Titanoscarcolites-bearing shales and limestone, at 

localities 92, 93, 97, 98, 100, 103, 106; Maldon inlier; 

Maldon Limestone, at locality 119; and Woodland 

Shale, at locality 116. Central inlier, Guinea Corn Formation, at localities 57, 60, 64, 77, 79, 80.

Age. Middle Maastrichtian.

7>/?es.-Holotype, ISNM 130671; paratypes, 130668- 

130670, 130672, 130673.

Actaeonella briggsi Sohl and Kollmann n. sp.

Plate 20, figure 4; plate 21, figures 14, 16 18.

Description. Shell small, with strongly but evenly 

curving convex sides, maximum diameter at middle of 

shell and constituting 43 to 49 percent of total height; 

apex broadly obtuse. Aperture proportionally broad. 

Columella long and thick, an inverted cone in form; 

columellar plaits decrease in strength anteriorly; 

margin of columella with parietal lip strongly downturned towards basal lip. Basal lip inclined, not 

thickened, and narrowly excavated.

Measurements (fig. 44). The available specimens 

range from 14.5 to 31.0 mm in height and have a maximum diameter of 6.8 to 13.4 mm. The plural angle 

ranges from 43° to 50°.

Discussion. Actaeonella briggsi differs from Actaeonella marchmontensis by its proportionally greater 

maximum diameter and by the shape of the columella, 

which is a broadly inverted cone. In contrast, the columella of A. marchmontensis is strongly inclined toward 

the axis below the middle columellar plait. Compared 

with other proportionally obese species of Actaeonella 

of the region, A. briggsi lacks the extreme concave upper whorl face of A. robinsoni and A. cubensis and has 

a proportionally longer columella.

The species is named for Reginald P. Briggs, in recognition of his many contributions to the understanding 

of the geology of Puerto Rico.

Occurrence. Puerto Rico: Barranquitas Quadrangle, 

Pozas Formation, Reves Member, at localities 4, 5, 6; 

San German Quadrangle, El Rayo Formation, at locali50

40

30

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MAXIMUM DIAMETER

FIGURE 44. Scatter diagram comparing height with width among 

selected specimens of Actaeonella briggsi Sohl and Kollmann, 

n. sp.

ties 48, 49a, 51, 51a, and Cotui Limestone (top), at 

locality 47.

Age. Early to middle Maastrichtian.

Types.-Holotype, USNM 306201; paratypes, USNM 

306166, 306199, 306200.

Actaeonella cubensis Sohl and Kollmann n. sp.

Plate 17, figure 2; plate 23, figures 2-5, 7, 8

1955. Actaeonella (Volvulina) aff. laevis (Sowerby). Lewis and 

Straczek, U.S. Geol. Survey Bull. 975-D, p. 194..

Diagnosis. An Actaeonella with strongly concave 

upper whorl profile, inflated whorls below midheight, 

and two columellar plications.

Description. Shell of medium size, strongly inflated 

on anterior half with strongly concave sides above midheight. Aperture broad, lacks a constriction between 

columella and parietal region. Columella short, bears 

two sharp plaits; columella bends strongly downwards 

out-side of aperture; lower part of columella and adaxial part of basal lip slightly reflected. In longitudinal 

section, first four whorls form a high spiral shell that is 

gradually concealed by fourth whorl; succeeding 

whorls are completely involute.

Measurements (fig. 45). The pleural angle of available specimens ranges from 33° to 40°.

Discussion. The presence of only two columellar 

plications distinguishes this species from all others in 

the Western Hemisphere except Actaeonella robinsoni. 

A. cubensis differs from that species especially in its 

more concave upper-whorl profile and in the great in-
SYSTEMATIC PALEONTOLOGY 73

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MAXIMUM DIAMETER

FIGURE 45. Scatter diagram comparing height with width among 

selected specimens of Actaeonella cubensis Sohl and Kollmann, 

n. sp.

flation of the whorls below midheight.

Occurrence. Cuba: Oriente Province, Habana (?) 

Formation (Lewis and Straczek 1955, p. 194), at locality 125.

Age. Judging by the associated fauna, Lewis and 

Straczek's assignment of the species to the late Senonian seems feasible. The specimens probably come 

from a late Campanian or Maastrichtian level.

Types. -Holotype, USNM 306183; paratypes: 

306127, 306182, 306184-306186.

Actaeonella robinsoni Sohl and Kollmann, n. sp.

Plate 17, figures 1, 4, 5; plate 22, figures 1, 2, 5

P1906. Actaeonella (S.S.) aff. grossouvrei Cossmann. Bose, Mexico 

Inst. Geol. Bol. 24, p. 91..

Diagnosis. An Actaeonella with a convex upper 

whorl profile and columella bearing only two columellar plications.

Description. Shells of medium size; maximum 

diameter at middle of shell and constitutes 40 percent 

of total height, whorls broadly convex over anterior 

part but sides concave in profile on upper part of shell, 

ending in acute tip. Aperture moderately wide. Columella proportionally long and bears strong columellar 

plaits; upper plait vanishes at aperture, lower plait 

passes into varicose basal lip, which is only partly 

known; labral edge of columella indistinct.

Measurements. The holotype is 58.6 mm high, has a 

maximum diameter of 24.8 mm, and a pleural angle of 

38 °. Paratypes range from 30 to 66 mm in height, have 

a maximum diameter of 13 to 32 mm, and a pleural 

angle ranging from 30° to 38°.

Discussion. This species is rare but possesses sufficiently distinctive characters to merit formal naming. 

Described species of Actaeonella that have an accentuated concave upper whorl profile are considerably less 

common than those having convex profiles. Among 

these, A. robinsoni is especially distinguished by having only two columellar plications rather than three. 

In this latter character, A. robinsoni is similar to A. 

cubensis Kollman and Sohl. A. cubensis differs notably 

in its midwhorl obesity and especially in the accentuated concavity of the upper whorl.

The species is named for Dr. E. Robinson of the University of the West Indies, Kingston, Jamaica, in recognition of his contributions to understanding of the 

geology of Jamaica.

Occurrence. Jamaica: Central inlier, Guinea Corn 

Formation, at localities 69, 70; Mexico, San Luis 

Potosi, Cardenas Formation, at locality 158.

Age. Maastrichtian.

Types.-Holotype, USNM 306176; paratypes, USNM 

306124-306126, 306174, 306175.

Actaeonella n. sp. A.

Plate 17, figure 3; plate 22, figs. 3, 4; plate 23, figures 1, 6

Discussion. The specimen figured on plate 22, 

figure 4, probably represents the largest known 

Actaeonella. All available specimens are incompletely 

preserved, but we estimate that some must have exceeded 180 mm in length and about 95 mm in maximum diameter. In cut section the shells are seen to be 

strongly inflated, the maximum diameter being below 

midheight. The upper whorl profile is slightly concave 

to flat, and the basal profile is rounded. The aperture is 

widest in the anterior part; the columella is thick and 

bears three plaits that decrease in strength anteriorly. 

The lower quarter of the whorl bears a strong palatal 

ridge. In this feature, this species differs from all 

Actaeonella known from the Western Hemisphere.

This actaeonellid shows several features comparable 

with those of described species from other regions. For 

example, the strong palatal ridge in the lower part of 

the whorl is similar to that of A. crassa (Dujardin, 

1835), which is of wide occurrence in the Upper Cretaceous rocks of Europe and eastern Asia (Kollmann, 

1965; Hacobjan, 1976). The Puerto Rican form here 

discussed, however, is proportionally much more obese. 

A. inflata Petho (1906), described from the upper 

Campanian-Maastrichtian of Fruska Gora in northern 

Serbia, possesses a similarly obese shell, but the profile
74 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

of the upper part of the shell is convex rather than concave to flat.

The few available specimens and their poor state of 

preservation precludes formal naming of this species.

Occurrence. Puerto Rico: Orocovis Quadrangle, 

Pozas Formation, Rio Bauta Member, at locality 26; 

Coamo Quadrangle, Maravillas Formation, at locality 

19.

Age. Campanian-early Maastrichtian.

Types.-Hypotypes, USNM 306128, 306177, 306180, 

306181.

Actaeonella sp. B.

Plate 22, figure 7

Discussion. Several incomplete specimens of an indeterminate Actaeonella have been found in collections 

preserved in erratic cobbles of sideritic ironstone concretions in glacial till from Martha's Vineyard, Mass. 

Although these specimens cannot be assigned to a 

given species, they are of special importance, as they 

represent the northernmost unquestioned occurrence 

of the genus in eastern North America (fig. 11).

The shells have a blunted apex and slightly convex 

whorl sides. The small but best preserved specimen (pi. 

22, fig. 7) measures 9 mm in height and has a maximum diameter of 3.8 mm. Other apertural features are 

obscured because of state of preservation.

Occurrence. Massachusetts: Martha's Vineyard, at 

locality 131.

Age. Late Cenomanian.

Types.-Hypotype, USNM 360179.

Actaeonella sp. C.

Plate 2, figure 5

Discussion. The collections of the U.S. Geological 

Survey contain two collections of fossils from along the 

Yukon River in Alaska that have yielded three specimens of an Actaeonella. According to D.L. Jones (oral 

commun. 1981), the assemblages are from the lower 

part of the Upper Cretaceous (Cenomanian-Turonian). 

Both collections are small and were made during the 

early exploratory phase of geologic work in Alaska 

(1899 and 1902). The many subsequent collections 

made from the Cretaceous of Alaska have failed to 

recover additional specimens.

These two occurrences are remarkable because they 

are the northernmost records of the family in the 

world. During Cenomanian time, Actaeonella had its 

greatest northward expansion in range (see fig. 7); It 

has been found in both eastern Siberia and Massachusetts, but both of these occurrences are many degrees of 

paleolatitude south of Alaska. This anomalously farnorth occurrence may be associated with a "suspect 

terrane" as viewed by Coney and others (1980). That is,

these specimens may be part of an autocthonous province transported to its present position from a more 

southerly source and subsequently accreted to the Cordilleran margin.

Occurrence. Alaska: Yukon River, at localities 169, 

170.

Types.-Hypotype, USNM 306203.

Actaeonella spp.

Actaeonella also occurs at other Cretaceous localities 

in Jamaica and Puerto Rico. These specimens, however, are either poorly preserved, fragmental, or 

known only in cut sections; proper specific placement 

cannot be made. The specimens are in rocks ranging in 

age from Albian to Maastrichtian and no doubt represent several species, some of which may be new.

Species of Albian Age occur in calcareous silty mudstones in both the Barrancas Limestone Member of the 

Magueyes Formation (locality 12, pi. 22, fig. 6) and the 

Rio Maton Limestone Member of the Robles Formation 

in central Puerto Rico. In both these units. Actaeonella 

is rare but is found in association with assemblages 

dominated by rudist bivalves and nerineid gastropods.

Undetermined Actaeonella is widespread in Campanian to Maastrichtian-Age rocks. In Puerto Rico, these 

fossils have been collected in the Botijas Limestone 

Member of the Pozas Formation in the Barranquitas 

Quadrangle (localities 10 and 11); the Sabana Hoyos 

Limestone Member of the Maravillas Formation of the 

Orocovis Quadrangle (localities 29, 30); the Maravillas 

Formation of the Rio Descalabrado (localities 34, 35) 

and Coamo (locality 21) Quadrangles; and the El Rayo 

Formation (locality 49) and Cotui Limestone (locality 

38) of the San German Quadrangle. In Jamaica, undetermined Actaeonella occurs in the Inoceramus 

Shales of the Anns Great River inlier (locality 123) in 

the Guinea Corn Formation of the Central inlier (localities 62, 64 and 76) and in the Titanosarcolites bearing 

limestones of the Jerusalem Mountain inlier (localities 110, 112-115).

Types.-Hypotype, USNM 360178.

CATALOGUE OF ACTAEONELLID SPECIES

The following listing of species assigned to the Family Actaeonellidae provides the basis for the construction of the distribution maps (figs. 5-13) and diversity 

graphs (figs. 3, 4). The listing reflects views of the 

authors on overall generic and subgeneric assignments. We have not investigated all species in equal 

detail because of lack of adequate comparative material. Thus, the listing should not be viewed as a final 

synonymy. For example, no decision has been reached 

on the validity of the exceptionally large number of
CATALOGUE OF ACTAEONELLID SPECIES 75

species that Pchelintsev and other authors have proposed from the Cenomanian, Turonian, and Coniacian 

of the Transcaucasus. In this regard, the catalogue is 

more a guide to the existing literature and a status 

report than a final expression of our views on validity 

of all species-level taxa.

A separate listing following this section includes 

those genera and species that Kollmann and Sohl 

(1980) and other authors have removed from the 

Actaeonellidae; most of these taxa are now assigned to 

the nerineacean family Itieriidae.

Trochactaeon (Neocylindrites) abbreviatus (Conrad)

1852. Nerinea abbreviata Conrad

1867. N. abbreviata Fraas

1891. N. abbreviata Whitfield

1900. Trochactaeon abbreviatus Bbhm

1927. Actaeonella (Trochactaeon) abbreviata Blanckenhorn

1939. Actaeonella abbreviata Delpey (part)

Age. Aptian.

Occurrence. Lebanon; Syria; Israel.

Trochactaeon (Neocylindrites) absalonis (Fraas)

1867. Phasianella absalonis Fraas

1867. P. absalonis Fraas

1878. Actaeonella absalonis Fraas

1890. A. absalonis Blanckenhorn

1900. Trochactaeon absalonis Bbhm

1976. Palaeotrochactaeon absalonis Hacobjan

Age. Cenomanian-Turonian.

Occurrence. Lebanon; Syria; Israel; USSR (Armenian SSR).

Trochactaeon (Trochactaeon) acuminatus (Fric) 

1897. Actaeonella acuminata Fric

Age. Coniacian. 

Occurrence. Czechoslovakia.

The following species has been assigned to T. (T.) 

acuminata but is here assigned elsewhere:

1934. Actaeonella acuminata Andert (see T. (T.) beyrichi Drescher)

Actaeonella (Trochactaeon) acutissima Bose, 1906

(see Trochactaeon (Mexicotrochactaeon) coniformis Bose)

Trochactaeon agbulakhensis G. A. Aliev, 1959, 1963

(see Trochactaeon (T.) intermedius Choffat)

Actaeonella agricolai Vidal

1921. Actaeonella agricolai Vidal 

1949. A. agricolai Bataller

Age. Maastrichtian (Garumnien). 

Occurrence. Spain (Province of Lerida).

Trochactaeon (Trochactaeon) alibeglyensis Pchelintsev

1954. Trochactaeon alibeglyensis Pchelintsev

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Trochactaeon (Neocylindrites) angustatus Pchelintsev

1953. Trochactaeon angustatus Pchelintsev

1967. Actaeonella (Trochactaeon) angustata Rahman

1974. Palaeotrochactaeon angustatus Hacobjan

1976. P. angustatus Hacobjan

Age. Cenomanian

Occurrence. USSR (Armenian SSR); Austria (Tyrol).

Trochactaeon (Trochactaeon) armenicus Pchelintsev

1954. Trochactaeon armenicus Pchelintsev

Age. Late Turonian.

Occurrence. USSR (Armenian SSR).

Trochactaeon (Trochactaeon) arnaudi Cossmann

1896. Trochactaeon arnaudi Cossmann 

1972. Mesotrochactaeon arnaudi Hacobjan 

1974. M. arnaudi Hacobjan 

1976. M. arnaudi Hacobjan

Age. Turonian.

Occurrence. France (Charente); USSR (Armenian 

SSR).

Trochactaeon (Trochactaeon) artsachensis (Hacobjan)

1976. Mesotrochactaeon artsachensis Hacobjan

Age. Turonian.

Occurrence. USSR (Armenian SSR).

Trochactaeon (Trochactaeon) atschadjurensis Pchelintsev, 1954

1954. Trochactaeon atschadjurensis Pchelintsev

Age. Coniacian.

Occurrence. USSR (Transcaucasus).

Trochactaeon (Trochactaeon) asiaticus Pchelintsev, 1953

(see Trochactaeon (Trochactaeon) schirabadensis (Djalilov))

Trochactaeon (Trochactaeon) babkovi Djalilov

1972. Trochactaeon babkovi Djalilov 

1977. T. babkovi Djalilov

Age. Campanian.

Occurrence. USSR (Tadzahkistan depression).

Actaeonella baconica Benko-Czabalay

1962. Actaeonella baconica Benko-Czabalay

Age. Albian.

Occurrence. Hungary (Bakony Mountains).

Trochactaeon (Neocylindrites) haconicus Benko-Czabalay

1962. Trochactaeon baconicus Benko-Czabalay 

1976. Paleotrochactaeon baconicus Hacobjan

Age. Albian.

Occurrence. Hungary (Bakony Mountains).

Trochactaeon (Neocylindrites) barremicus Sayn

1932. Trochactaeon barremicus Sayn

1932. T. barremicus var. cylindrica Sayn

1932. T. barremicus var. mumiaeformis Sayn
76 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Age. Barremian. 

Occurrence. France (Drome).

Trochactaeon (Neocylindrites) baylei Leymerie of Daguin

1927. Actaeonella (Trochactaeon) baylei Leymerie of Daguin

Age. Senonian. 

Occurrence. Morocco.

Trochactaeon (Neocylindrites) belgicus Cossmann

1865. Actaeonella conica Briart and Cornet

1896. Trochactaeon belgica Cossmann (name substituted for

Actaeonella conica Briart and Cornet, 1865) (not Zekeli, 1852,

and part of Actaeonella briarti Geinitz, 1874) 

1939. Actaeonella (Trochactaeon) conica Marliere

Age. Late Albian. 

Occurrence. Belgium.

Trochactaeon (Trochactaeon) beyrichi Drescher

1863. Actaeonella beyrichi Drescher

1910. A. beyrichi Weinzettl

1934. A. beyrichi Andert

1934. A. acuminata Andert

1963. Purpuractaeon beyrichi Hacobjan

Age. Coniacian

Occurrence. Czechoslovakia; German Democratic 

Republic; Poland.

Trochactaeon (Trochactaeon) biconicus Pchelintsev

1954. Trochactaeon biconicus Pchelintsev

Age. Early Turonian. 

Occurrence. USSR (Armenian SSR).

Actaeonella borneensis Nuttall and Leong

1959. Actaeonella cf. laevis d'Orbigny, Chubb 

1972. A. borneensis Nuttall and Leong

Age. Late Campanian-Maastrichtian. 

Occurrences. Borneo', Cuba (Santa Clara Province); 

Mexico (Chiapas).

Trochactaeon (Neocylindrites) boutillieri Cossmann

1896. Trochactaeon boutillieri Cossmann

1900. T. boutillieri Cossmann

1916. T. boutillieri Cossmann

1932. T. boutillieri Sayn

Age. Barremian.

Occurrence. France (Bouches du Rhone, Gard, 

Drome).

Actaeonella (Trochactaeon) brevis Bose, 1906

(see Trochactaeon (Mexicotrochactaeon) coniformis (Bose)

Trochactaeon (Neocylindrites) briarti Geinitz

1874. Trochactaeon briarti Geinitz 

1905. Actaeonella conica Deninger 

1910. A. briarti Weinzettl 

1974. P. briarti Hacobjan 

1976. P. briarti Hacobjan

Age. Late Cenomanian.

Occurrence. Belgium; Czechoslovakia; Saxony; 

USSR (Armenian SSR).

Actaeonella browni Sohl and Kollmann

Age. Late Albian or Cenomanian. 

Occurrence. Guatemala.

Trochactaeon (Mexicotrochactaeon) burckhardti Bose

1923. Actaeonella (Trochactaeon) burckhardti Bose

1923. A. (T.) burckhardti var. obliquecostata Bose

1923. A. (T.) gigantea Sowerby, sp. n. var. mexicana Bose

1923. A. (T.) humboldti Bose

1923. A. (T.) quadriplicata Bose

1930. Actaeonella luciano Maury

1930. A. pompei Maury

1930. A. cf. silvai Maury

1976. Neotrochactaeon burckhardti Hacobjan (not pi. 15,

figs. 3-5, 7-10, and pi. 14, figs. 12, 13) 

1976. Trochactaeon obliqueplicatus Hacobjan (included is T. impressus

Pchelintsev, 1954, as a synonym

Age. Coniacian, Maastrichtian.

Occurrence. Mexico (State of Guerrero); Brazil; 

Jamaica.

The following species has been assigned to T. (M.) 

burckhardti but should be placed elsewhere:

1950. Actaeonella (Trochactaeon) burckhardti Bose? Maldonado-Koerdell. 

Actaeonella briggsi Sohl and Kollmann

Age. Maastrichtian. 

Occurrence. Puerto Rico.

Actaeonella caucasica caucasica Zekeli

1852. Actaeonella caucasica Zekeli

\928a.A. caucasica Pchelintsev

1953. A. caucasica Pchelintsev

? 1958. A. caucasica Aliev

1965. A. caucasica caucasica Kollmann

1972. A. (Pchelincevella) caucasica Djalilov

1974. Actaeonella caucasica Hacobjan

1976. A. caucasica Hacobjan

1977. A. (Pchelincevella) caucasica Djalilov

Age. Cenomanian.

Occurrence. USSR (Transcaucasus and questionably Elenstop Mountains, Shankhorsk region).

Actaeonella caucasica grossouvrei Cossmann

1896. Actaeonella grossouvrei Cossmann 

1901. A. grossouvrei Choffat 

1920. A. grossouvrei Roman and Mazeran 

1965. A. grossouvrei Roman and Mazeran 

1972. A. (Pchelincevella) grossouvrei'Djalilov 

1977. A. (P.) grossouvrei Djalilov

[Assigned to Actaeonella uchauxensis Cossmann by Hacobjan, 1974 and 

1976]

Age. Turonian.

Occurrence. France (Uchaux); Portugal.

The following species have been assigned to Actaeo-
CATALOGUE OF ACTAEONELLID SPECIES 77

nella caucasica grossouvrei but should be placed elsewhere:

1906. Actaeonella aff. A. grossouvrei Bose 

1972. A. grossouvrei Collignon (see A. delgadoi)

Actaeonella caucasica styriaca Kollmann

1892. Actaeonella (Volvulina) laevis Futterer

1895. A. (V.) laevis Bohm

1964. A. (Actaeonella) caucasica Kollmann

1962. Actaeonella caucasica styriaca Benkb-Czabalay

P1964. Actaeonella caucasica styriaca Benkb-Czabalay

1965. A. caucasica styriaca Kollmann

? 1966.^4. caucasica Lupu

1972. A. (Pchelincevella) styriaca Djalilov

1977. A. (Pchelincevella) styriaca Djalilov

[Assigned to Actaeonella crassa by Hacobjan, 1974 and 1976]

Age. Late Coniacian-Santonian. 

Occurrence. Austria; Italy (Venetian Alps); Hungary; Romania.

Actaeonella caucasica schiosensis Bohm

1895. Actaeonella (Volvulina) schiosensis Bohm

1900. A. (V.) schiosensis Fittipaldi

1965. Actaeonella caucasica schiosensis Kollmann

1972. A. (Pchelincevella) schiosensis Djalilov

1974. Actaeonella schiosensis Hacobjan

1976. A. schiosensis Hacobjan

1977. A. (Pchelincevella) schiosensis Djalilov

Age. Late Cenomanian-early Turonian. 

Occurrence. Italy (Venetian Alps; Campobasso).

Actaeonella caucasica zouparriensis Choffat

1901. Actaeonella zouparriensis Choffat 

1965. A. caucasica zouparriensis Kollmann 

1972. A. (Pchelincevella) zouparriensis Djalilov 

1977. A. (P.) zouparriensis Djalilov

Age. Turonian. 

Occurence. Portugal.

Trochactaeon (Trochactaeon) caucasicus Pchlintsev

1953. Trochactaeon caucasicus Pchelintsev

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Trochactaeon (Trochactaeon) choffati Pchelintsev

1954. Trochactaeon choffati Pchelintsev

Age. Coniacian.

Occurrence. USSR (Transcaucasus).

Trochactaeon (Trochactaeon) collignoni (Darteville and Brehion)

1956. Actaeonella (Trochactaeon) collignoni Darteville and Brebion 

1974. Trochactaeon collignoni Hacobjan

Age. Early Cenomanian-late Turonian. 

Occurrence. Gabon; USSR (Armenian SSR).

Trochactaeon (Trochactaeon) communis Pchelintsev

1954. Trochactaeon communis Pchelintsev

Age. Coniacian.

Occurrence. USSR (Armenian SSR).

Trochactaeon (Trochactaeon) conicus Miinster

1844. Tornatella conica Miinster in Goldfuss

1850. Actaeonella conica d'Orbigny

1852. A. conica Zekeli

1959. A. (Trochactaeon) conica Pokorny

1962. Trochactaeon (Trochactaeon) conicus Benko-Czabalay

1966. Actaeonella (Trochactaeon) conica Marincas

1967. Trochactaeon (Trochactaeon) conicus Kollman

1974. Spiractaeon conicus Hacobjan

1974. Spiractaeon conica Hacobjan

1976. S. conica Hacobjan

Age. Santonian.

Occurrence. Austria; Romania; Hungary; USSR 

(Transcaucasus, Central Asia).

The following species have been assigned to T. (T.) 

conicus (Miinster) but should be placed elsewhere:

1865. Actaeonella conica Stoliczka (see T. (T.) goldfussi (d'Orbigny))

1865. A. Conica Briart and Cornet (see T. (T.) belgicus Cossmann)

1881. Tornata conica Quenstedt (see T. (T.) goldfussi)

1892. Actaeonella conica Futterer (see T. (Neocylindrites) sp.)

1939. (Trochactaeon) conica Marliere (see T. (T.) belgicus Cossmann)

1957. A. conica Pejovic

1965. A. (Trochactaeon) conica Lupu

Trochactaeon (Trochactaeon) conicus Hacobjan, not Miinster

1963. Purpuractaeon conicus Hacobjan 

1972. P. conicus Hacobjan

Age. Coniacian.

Occurrence. USSR (Devi River, Armenian SSR).

Trochactaeon (Mexicotrochactaeon) coniformis coniformis (Bbse)

1906. Actaeonella (Trochactaeon) coniformis Bose

1906. A. (T.) acutissima Bbse

1906. A. (T.) occidentalis Bbse

1906. A. (T.) inconstans Bbse

1906. A. (T.) irregularis Bbse

1906. A. (T.) brevis Bose

1906. A. (T.) planilateralis B5se

1906. A. (T.) potosiana Bose

1906. A. (T.) variabilis Bbse

1968. Actaeonella coniformis Myers

1972. Mexicotrochactaeon acutissima Hacobjan

1972. M. irregularis Hacobjan

1972. M. planilateralis Hacobjan

1972. M. potosiana Hacobjan

1972. M. variablilis Hacobjan

1976. Spiractaeon brevis Hacobjan

Age. Maastrichtian.

Occurrence. Mexico (Cardenasa area, State of San 

Luis Potosi).

All following assigned herein to T. (Neocylindrites) 

punctatus Maldonado-Koerdell, 1950:

1950. A. (Trochactaeon) coniformis Maldonado-Koerdell

1950. A. (Trochactaeon) acutissima punctata Maldonado-Koerdell

1950. A. cf. A. (T.) occidentalis Maldonado-Koerdell (p. 215, fig. 17.)
78 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

1950. A. cf. (T.) inconstans Maldonado-Koerdell 

1950. A. (T.) planilateralis Maldonado-Koerdell

Trochactaeon (Trochactaeon) conoideus (Matheron)

1842. Mitra conoidea Matheron

1902. Actaeonella (Trochactaeon) conoidea Cossmann

Age. Santonian.

Occurrence. France (Bouches-du-Rhone).

Actaeonella coquiensis Sohl and Kollman

Age. Maastrichtian 

Occurrence. Puerto Rico; Jamaica

Trochactaeon (Neocylindrites) cordeiroi (Choffat)

1888. Cylindrites delgadoi Choffat, in Choffat and Loriol

1888. C. delgadoi Choffat

1956. Actaeonella (Trochactaeon) cordeiroi Darteville and Brebion

1976. Paleotrochactaeon cordeiroi Hacobjan

Age. Albian-early Cenomanian. 

Occurrence. Angola; Bas-Congo; USSR (Armenian 

SSR).

Trochactaeon (Trochactaeon) cossmanni Choffat

(see T. (T.) giganteus subglobosus (Miinster))

Trochactaeon (Trochactaeon) cossmanni var. obesus Hojnos

(see T. (T.) giganteus subglobosus Miinster)

Actaeonella crassa (Dujardin)

1835. Volvaria crassa Dujardin 

1842. Actaeonella crassa d'Orbigny 

1863. A. crassa Schafhautl 

1865. Volvulina crassa Stoliczka

1953. Actaeonella crassa Pchelintsev

1954. A. gracilis Pchelintsev

1954. A. crassa Pchelintsev

P1956. A. cf.A. crassa Darteville and Brebion

P1964. A. crassa Djalilev (assigned to A. longa by Hacobjan, 1976)

1965. A. aff. A. crassa Kollmann

1972. A. (Pchelincevella) crassa in Djalilov

1972. A. (P.) gracilis Djalilov

1974. Acteonella crassa Hacobjan

1976. A. crassa Hacobjan

1977. A. (P.) crassa Djalilov 

1977. A. (P.) gracilis Djalilov

Age. Cenomanian-Coniacian.

Occurrence. France; Austria; USSR (Transcaucasus, Darvaza, Turkmen, Sikhote-Alin), questionably, 

Gabon, west Africa.

The species listed below has been assigned to A. 

crassa but should be placed elsewhere:

1916. Actaeonella crassa Douville (Campanian; Tibet)

Trochactaeon? cretaceus (Miiller)

1851.

1859.

1888.

1974.

Cassidaria cretacea Miiller 

Globiconcha maxima Miiller 

Actaeonella cretacea Holzapfel (pi. 7,

to Sevenella by Hacobjan, 1974) 

Purpuractaeon cretaceus Hacobjan

figs. 11, 15, 16, assigned

Age. Coniacian or Santonian. 

Occurrence. West Germany (Aachen).

Trochactaeon (Neocylindrites) crisminensis Choffat

1901. Trochactaeon crisminensis Choffat

1953. T. subangustatus Pchelintsev

1954. T. subangustatus Pchelintsev

1966. Actaeonella (T.) subangustatus Lupu

1972. Paleotrochactaeon crisminensis Hacobjan

1974. P. crisminensis Hacobjan

1976. P. crisminensis Hacobjan

(Hacobjan, 1976, also included Trochactaeon cylindraceus Stoliczka)

Age. Late Albian-Cenomanian. 

Occurrence. Portugal; USSR (Armenian SSR, Central Asia).

Actaeonella cubensis Sohl and Kollman

Age. Maastrichtian. 

Occurrence. Cuba.

Trochactaeon (Neocylindrites) cumminsi (Stanton)

1947. Trochactaeon cumminsi Stanton

1947. T. parvus Stanton

1955. Actaeonella (Trochactaeon) parvus Allison

Age. Albian.

Occurrence. Texas; Mexico (Baja California).

Trochactaeon (Trochactaeon) cylindraceus (Stoliczka)

1868. Actaeonella cylindracea Stoliczka

1976. Paleotrochactaeon cylindraceus Hacobjan (excludes pi. 14, fig. 10 

[= P. crisiminensis Choffat of Hacobjan])

Age. Otatoor Group Cenomanian(?)-Turonian.

Occurrence. Southern India; Gabon; Angola?; USSR 

(Armenian SSR).

The species listed below have been assigned to T. (T.) 

cylindraceus (Stoliczka) but should be placed elsewhere:

1884. Trochactaeon cylindraceus Whiteaves (affinities unknown) 

1956. Actaeonella (Trochactaeon) cylindracea Darteville and Brebion

Trochactaeon (Trochactaeon) cylindricus Pchelintsev

1928. Trochactaeon cylindricus Pchelintsev

1953. T. cylindricus Pchelintsev

1954.. T. cylindricus Pchelintsev

1976. Mesotrochactaeon cylindricus Hacobjan

Age. Cenomanian.

Occurrence. USSR (Armenian SSR).

Trochactaeon (Trochactaeon) cycloideus Pchelintsev

1954. Trochactaeon cycloideus Pchelintsev

Age. Late Senonian. 

Occurrence. USSR (Armenian SSR).

Actaeonella decorata Hacobjan

1976. Acteonella decorata Hacobjan

Age. Late Cenomanian. 

Occurrence. USSR (Armenian SSR).
CATALOGUE OF ACTAEONELLID SPECIES 79

Trochactaeon? damesi Futterer

1892. Cylindrites damesi (Futterer)

Age. Early Senonian. 

Occurrence. Italy (Venetian Alps).

Trochactaeon (Trochactaeon) darwasensis Djalilov

1964. Trochactaeon darwasensis Djalilov 

1977. Spiractaeon darwasensis Djalilov

(Hacobjan 1976, assigned pi. 24, fig. 3, of Djalilov, 1964 to Spiractaeon 

obtusus)

Age. Late Coniacian.

Occurrence. USSR (Darvaza, Turkmen SSR).

Trochactaeon? delgadoi (Choffat)

(see Trochactaeon (Neocylindrites) cordeiroi (Choffat))

1888. Cylindrites delgadoi Choffat, in Choffat and Loriol, 1888 

1956. Cylindrites(?) delgadoi Darteville and Brebion

Age. Albian. 

Occurrence. Angola.

Actaeonella delgadoi Choffat

1901. Actaeonella delgadoi Choffat

1916. A. gracilis Douville

1955. A. gracilis Mahmoud

1963. A. gracilis var. Maqharensis Abbass

1972. A. grossouvrei Collignon

1972. A. (Pchelincevella) delgadoi Djalilov

1977. A. (P.) delgadoi Djalilov

Age. Albian-Cenomanian.

Occurrence. Portugal; Egypt (Sinai); Madagascar; 

USA (Texas).

Trochactaeon dilatatus Pchelintsev, 1954

(see T. (T.) giganteus subglobosus (Minister))

Trochactaeon (Trochactaeon) dolioliformis Pchelintsev

1953. Trochactaeon dolioliformis Pechelintsev

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Trochactaeon (Trochactaeon) dubertreti (Delpey)

1939. Actaeonella dubertreti Delpey

Age. Late Cenomanian. 

Occurrence. Lebanon.

Actaeonella duckettsensis Sohl and Kollmann

Age. Maastrichtian. 

Occurrence. Jamaica.

Trochactaeon (Trochactaeon) ellipsoides (Fittipaldi)

(see Trochactaeon (T.) matensis (Fittipaldi))

1900. Actaeonella ellipsoides Fittipaldi

1953. Trochactaeon subotusus Pchelintsev

1953. T. matensis Pchelintsev (part) (pi. 48, fig. 16 = T. (T.) tovusensis

Hacobjan)) 

1953. T. regularis Pchelintsev (part) (fide, Hacobjan, 1974, not. pi. 50,

figs. 8, 10)

Age. Cenomanian-Turonian.

Occurrence. Italy (Venetian Alps); USSR (Armenian 

SSR).

Trochactaeon (Trochactaeon) ellipticus Zekeli

(see Trochactaeon conicus Miinster)

1852. Actaeonella elliptica Zekeli

1954. Trochactaeon ellipticus Pchelintsev

1976. Spiractaeon ellipticus Hacobjan

Actaeonella elongata Kollman

1852. Actaeonella laevis Zekeli (part) 

1965. A. elongata Kollmann

Age. Late Santonian. 

Occurrence. Austria.

Trochactaeon (Trochactaeon) excelsus Pchelintsev

1954. Trochactaeon excelsus Pchelintsev 

1974. Neotrochactaeon excelsus Hacobjan

Age. Coniacian.

Occurrence. USSR (Armenian SSR).

Actaeonella? faba (Kner)

1849. Volvariafaba Kner

1972. Actaeonella (Pchelincevella) faba Djalilov

1977. A. (Pchelincevella) faba Djalilov (possibly belongs to Ellipsoscapha)

Age. Maastrichtian. 

Occurrence. USSR (Lvov).

Trochactaeon (Trochactaeon) frazierensis (Anderson)

1958. Actaeonella frazierensis Anderson

1958. A. rustica Anderson

1979 (1976). A. frazierensis Jones, Sliter, and Popenoe

Age. Turonian.

Occurrence. USA (northern California).

Actaeonella fusiformis Coquand

1865. Actaeonella fusiformis Coquand 

1887. A. fusiformis Mallada 

1939. A. fusiformis Delpey 

1949. A. fusiformis Bataller 

? 1955. A. fusiformis Allison

Age. Aptian.

Occurrence. Spain (Teruel); Lebanon.

Trochactaeon (Trochactaeon) galloprovincialis (Cossmann)

1842. Actaeonella gigantea d'Orbigny

1850. A. gigantea d'Orbigny

1895. A. (Trochactaeon) gigantea Cossmann

1897. Trochactaeon giganteus var. galloprovincialis Cossmann

1901. T. giganteus Choffat

1901. T. giganteus var. glandiformis Choffat

1967. T. (Trochactaeon) aff. galloprovincialis Kollmann (assigned to 

Neotrochactaeon giganteus (Sowerby) by Hacobjan, 1976)

Age. Turonian(?)-Santonian.

Occurrence. Austria; Portugal; France (LeBeausset).
80 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Trochactaeon (Trochactaeon) giganteus giganteus (Sowerby)

1832. Tornatella gigantea Sowerby

1957. Actaeonella renauxiana Pejovic

1959. A. (Trochactaeon) gigantea gigantea Pokorny

1959. A. (T.) gigantea subglobosa Pokorny

1966. A. (T.) gigantea Todirita-Mihailescu

1966. A. (T.) gigantea ventricosa Todirita-Mihailescu

1966. A. (T.) gigantea ventricosa Marincas

1967. Trochactaeon (T.) giganteus Kollmann 

1967. T. (T.) giganteus giganteus Kollmann 

1972. Neotrochactaeon gigantea Hacobjan 

1974. N. giganteus Hacobjan 

1976. N. giganteus Hacobjan

Age. Santonian-early Campanian. 

Occurrence. Austria; Yugoslavia; Hungary; Portugal; 

Romania; Greece; ?USSR (Transcaucasus-Cenomanian).

The species listed below have been assigned to T. (T.) 

giganteus giganteus (Sowerby) but should be placed 

elsewhere:

1842. Actaeonella gigantea d'Orbigny (= Trochactaeon (T.) galloprovincialis Cossmann)

1844. A. gigantea Miinster (= Trochactaeon (T.) kuehni Kollmann 

1967, to Neotrochactaeon ventricosus (Hojnos) by Hacobjan, 

1976)

1844. Tornatella subglobosa (Miinster) (= Trochactaeon (T.) giganteus 

subglobosus)

1850. Actaeonella subglobosa d'Orbigny

1851. Actaeon giganteus Miiller

1852. Actaeonella gigantea (Sowerby) Zekeli (= Trochactaeon subglobosus (Miinster) fide Kollmann, 1967)

1852. A. lamarcki Zekeli (pi. 6, fig. 2 = T. (T.) giganteus subglobosus 

(Miinster), fide Kollmann, 1967)

1884. Tornatella giganteus (Sowerby) Quenstedt (fig. 128 = Trochactaeon goldfussi (d'Orbigny))

1888. Actaeonella gigantea Holzapfel

1896. Trochactaeon giganteus Cossmann

1897. T. giganteus var. galloprovincialis Cossmann (elevated to specific 

rank by Kollmann 1967; included in T. (T.) giganteus by 

Hacobjan, 1976)

1901. T. cossmanni Choffat (= T. (T.) giganteus subglobosus (Miinster))

1901. T. giganteus Choffat (= Trochactaeon (T.) galloprovincialis 

Cossmann)

1901. Trochactaeon giganteus var. intermedia Choffat (= Trochactaeon 

(Trochactaeon) intermedius (Choffat))

1901. T. giganteus var. glandiformis Choffat (= T. (T.) galloprovincialis 

Cossmann, fide Kollmann, 1967)

1906. Actaeonella (Trochactaeon) aff. gigantea Rose (= Neotrochactaeon fide Hacobjan, 1972; see Trochcactaeon (Mexicotrochactaeon) burckhardti)

1910. Actaeonella gigantea Weinzettl

1911. A. gigantea Fric (= Neocylindrites sp.) 

1921. A. (Trochactaeon) gigantea ventricosa Hojnos 

1921. Trochactaeon transylvanicus Hojnos (= T. (T.) giganteus subglobosus Miinster)

1923. Actaeonella (Trochactaeon) gigantea var. mexicana Bose 

1929. Actaeonella gigantea Steinmann

1950. A. (Trochactaeon) gigantea globosa Maldonado-Koerdell

1951. Actaeonella gigantea Pasic (= Mesotrochactaeon subglobosa fide 

Hacobjan, 1976)

1957. A. giganteus var. glandiformis Pejovic (= Trochactaeon (Trochactaeon) lamarcki lamarcki (Sowerby))

1959. Trochactaeon giganteus (Sowerby), Mitzopoulos (= Neotrochactaeon ventricosa (Hojnos) fide Hacobjan, 1976)

1959. Actaeonella (Trochactaeon) gigantea ventricosa Pokorny (= Trochactaeon (Trochactaeon) giganteus subglobosa (Miinster))

Trochactaeon (Trochactaeon) giganteus subglobosus (Miinster)

1844. Tornatella subglobosa Miinster 

1850. Actaeonella subglobosa d'Orbigny 

1852. A. gigantea Zekeli

1852. A. lamarcki Zekeli (part)

1853. A. gigantea Reuss (part)

1896. Trochactaeon cossmanni Choffat

1901. T. cossmanni Choffat

1921. T. giganteus var. ventricosus Hojnos

1921. T. cossmanni'var. obesus Hojnos

1921. T., typus II Hojnos

1921. T. transyivanicus Hojnos

1951. Actaeonella gigantea Pasic

1954. Trochactaeon cossmanni Pchelintsev

1954. T. dilatatus Pchelintsev

1959. Actaeonella (Trochactaeon) gigantea subglobosa Pokorny

1962. Trochactaeon (Trochactaeon) giganteus subglobosus Benko-Czabalay

1967. T. (T.) giganteus subglobosus Kollmann

1976. Neotrochactaeon ventricosa (Hojnos) Hacobjan

Age. Santonian-early Campanian(?). 

Occurrence. Austria; Yugoslavia; Hungary; 

Romania; Portugal; USSR (Transcaucasus).

Trochactaeon (Trochactaeon) ghazirensis (Delpey)

1939. Actaeonella ghazirensis Delpey 

1952. A. ghazirensis Termier and Termier

Age. Cenomanian-Turonian. 

Occurrence. Lebanon.

Trochactaeon (Trochactaeon) glandiformis (Zekeli)

(see T. (T.) sanctaecrucis)

1852. Actaeonella glandiformis Zekeli (figs. 9a-9b = Trochactaeon collignoni Darteville and Brebion, fide Hacobjan, 1974)

1901. Trochactaeon glandiformis Choffat (pi. 1, fig. 20, not pi. 1,

fig. 21 = Neotrochactaeon choffati (Pechelintsev) fide Hacobjan, 1976)

1954. Trochactaeon sub glandiformis Pchelintsev

1954. T. subovatus Pchelintsev

1974. Neotrochactaeon glandiformis Hacobjan

1976. N. glandiformis Hacobjan

Age. Late Turonian-Santonian.

Occurrence. Portugal; Austria; USSR (Transcaucasus).

Trochactaeon (Trochactaeon) goldfussi (d'Orbigny)

1844. Tornatella lamarckii Munster 

1850. Actaeonella goldfussi d'Orbigny 

1850. A. toucasiana d'Orbigny

1852. A. lamarcki Zekeli (pi. 6, fig. 4, 5, not fig. 2, 3 = Trochactaeon 

giganteus fide Kollmann, 1967)

1853. A. goldfussi Reuss

1884. Tornata conica Quenstedt

1884. T. gigantea Quenstedt

1884. T. lamarcki Quenstedt

1959. Actaeonella (Trochactaeon) goldfussi Pokorny

1962. Trochactaeon (Trochactaeon) goldfussi Benko-Czabalay

1966. Actaeonella (Trochactaeon) goldfussi Marincas
CATALOGUE OF ACTAEONELLID SPECIES 81

1966. A. (T.) goldfussi Todirita-Mihailescu

1967. Trochactaeon (Trochactaeon) goldfussi Kollmann

Age. Late Turonian-Santonian.

Occurrence. France; Austria; Romania; Hungary.

Trochactaeon (Neocylindrites) gosaviensis Kollman

1967. Trochactaeon (Neocylindrites) gosaviensis Kollmann

Age.. Coniacian-Santonian. 

Occurrence. Austria.

Actaeonella gracilis Douville, 1916

(see A delgadoi Choffat)

Actaeonella gracilis Pchelintsev, 1954

(see/4. crassa Dujardin)

Actaeonella grossouvrei Cossmann, 1896

(see A. caucasica grossouvrei Cossmann)

Trochactaeon (Trochactaeon) gracilis Pchelintsev

1954. Trochactaeon gracilis Pchelintsev 

1976. Spiractaeon obtusus Hacobjan

Age. Late Coniacian-Santonian. 

Occurrence. USSR (Transcaucasus).

Trochactaeon (Neocylindrites) gradatus Kollman

1967. Trochactaeon (Neocylindrites) gradatus Kollmann

Age. Late Coniacian-Santonian. 

Occurrence. Austria.

Trochactaeon (Mexicotrochactaeon) granthamensis Sohl and Kollmann

Age. Middle Maastrichtian. 

Occurrence. Jamaica.

Trochactaeon (Trochactaeon) gulistanensis Pchelintsev

1954. Trochactaeon gulistanensis Pchelinstev

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Actaeonella (Trochactaeon) humboldti Bdse, 1923

(see Trochactaeon (Mexicotrochactaeon) burckhardti (Bose))

Trochactaeon (Trochactaeon) humei (Greco)

1916. Actaeonella (Trochactaeon) humei Greco

Age. Turanian. 

Occurrence. Egypt.

Trochactaeon (Trochactaeon) idjevanensis (Hacobjan)

1976. Paleotrochactaeon idjevanensis Hacobjan

Age. Late Cenomanian. 

Occurrence. USSR (Armenian SSR).

Trochactaeon impressus Pchelintsev, 1954

(see T. obliqueplicatus Bose of Hacobjan, 1976)

Actaeonella (Trochactaeon) inconstans Bo'se, 1906

(see Trochactaeon (Mexicotrochactaeon) coniformis (Bose))

Actaeonella inflata Petho

1906. Actaeonella (Volvulina) inflata Petho

1982. Ovactaeonella dolium Konovalov and Poyarkova

Age. Maastrichtian.

Occurrence. Yugoslavia (Fruska-Gora).

Actaeonella (Trochactaeon) irregularis Bose, 1906

(see Trochactaeon (Mexicotrochactaeon) coniformis (Bose))

Trochactaeon insignis Pchelintsev, 1954

(see T. pruniformis Hacobjan, 1974)

Actaeonella involuta Coquand

1862. Actaeonella involuta Coquand 

1896. A. involuta Cossmann

Age. Late Santonian.

Occurrence. France (Lot, Charente); Algeria? (on 

the basis of an internal mold of questionable affinities).

Actaeonella jicarensis Sohl and Kollmann

Age. Maastrichtian. 

Occurrence. Puerto Rico.

Trochactaeon (Trochactaeon) kuehni Kollman

1967. Trochactaeon (Trochactaeon) kuehni Kollmann

Age. Santonian. 

Occurrence. Austria.

Actaeonella laevis laevis (Sowerby)

1832.

1845.

1852.

1853.

1860.

1865.

1888.

1895.

1896.

1959

1965

1976

Volvaria laevis Sowerby 

Actaeonella laevis Reuss 

A. laevis Zekeli (part) 

A. laevis Reuss 

A. obliquecostata Stoliczka 

Volvulina laevis Stoliczka 

Actaeonella laevis Holzapfel 

A. terebellum Cossmann 

A. laevis Cossmann 

. A. laevis Choffat 

A. laevis Pokorny 

A. laevis laevis Kollmann

A. laevis Hacobjan

The species listed below have been assigned to 

Actaeonella laevis laevis but should be placed elsewhere:

1842. Actaeonella laevis d'Orbigny (assigned to A. caucasica grossouvrei

Cossmann by Kollmann, 1965, to A. uchauxensis Cossmann by

Hacobjan, 1976)

1852. A. laevis Bronn (assigned to A. caucasica grossouvrei by Kollmann, 1965, to A. uchauxensis by Hacobjan, 1976)

1890. A. laevis Blanckenhorn (assigned to A. styriaca by Kollmann, 1965) 

1892. A. (Volvulina) laevis Futterer 

1895. A. (Volvulina) laevis Bohm (assigned to A. caucasica styriaca by

Kollmann, 1965) 

1895. Actaeonella laevis Cossmann (assigned to A. uchauxensis by

Hacobjan, 1976 

1899. A. (Volvulina) laevis d'Alesandri

1910. Actaeonella laevis Weinzettl

1911. A. laevis Fric

1939. A. laevis Delpey

1956. A. laevis Darteville and Brebion

Age. Coniacian(?)-Santonian.

Occurrence. Austria (Gosau); ?Portugal; West Germany (Aachen region); USSR (Armenian SSR).
82 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Trochactaeon (Trochactaeon) lamarcki lamarcki (Sowerby)

1832. Tornatella lamarcki Sowerby 

1850. Actaeonella lamarcki d'Orbigny 

1850. A. valuta d'Orbigny

1852. A. valuta Zekeli

1853. A. lamarcki Reuss

1867. A. syriaca Fraas

1867. A. salomonis Fraas

1884. A. valuta Quenstedt

1903. A. salomonis Dacque'

1921. A. salomonis Dacque

1921. A. valuta Dacque'

1957. A. giganteus var. glandiformis Pejovic

1959. A. (Trochactaeon) lamarcki Pokorny

1963. A. (T.) salomonis Abbass

1964. A. (T.) lamarcki Kollmann 

1966. A. (T.) lamarcki Marincas

1966. A. (T.) lamarcki Todirita-Mihailescu

1967. Trochactaeon (Trochactaeon) lamarcki lamarcki Kollmann 

1974. Spiractaeon lamarcki Hacobjan 

1976. S. lamarcki Hacobjan

Age. Turonian-Santonian.

Occurrence. Austria; Romania; Yugoslavia; Greece; 

Syria; Egypt; USSR (Armenian SSR).

The species listed below have been assigned to Trochactaeon (Trochactaeon) lamarcki lamarckibut should 

be placed elsewhere:

1844. Tornatella lamarckii Miinster (see Trochactaeon (Trochactaeon)

goldfussi) 

1852. Actaeonella lamarcki Zekeli (pi. 6, figs. 2-5, see Trochactaeon

(Trochactaeon) giganteus subglobosus) 

1851. Actaeonella lamarcki Pasic (see Trochactaeon (Trochactaeon)

giganteus subglobosus)

Trochactaeon (Trochactaeon) lamarcki brandenbergensis Kollmann

1967. Trochactaeon (Trochactaeon) lamarcki brandenbergensis Kollmann

Age. Late Coniacian. 

Occurrence. Austria.

Trochactaeon (Sevanella) laticostata (Hacobjan)

1972. Sevanella laticostata Hacobjan

1974. S. laticostata Hacobjan

1976. S. laticostata Hacobjan

Age. Late Coniacian. 

Occurrence. USSR (Armenian SSR).

Trochactaeon (Trochactaeon) latus Pchelintsev

1954. Trochactaeon latus Pchelintsev

Age. Early Cenomanian. 

Occurrence.-USSR (Armenian SSR).

Actaeonella longa Hacobjan

1954. Actaeonella crassa Pchelintsev 

1976. A. longa Hacobjan

Age. Late Coniacian.

Occurrence. USSR (Armenian SSR; Turkmen SSR. 

(SW. Darvaz)).

Trochactaeon longisulcus Hacobjan

1912. Actaeonella (Trochactaeon) salomonis Pervinquiere

1965. Actaeonella? salomonis Homaoui and Raab

1976. Mesotrochactaeon longisulcus Hacobjan

Age. Turonian.

Occurrence. USSR (Armenian SSR).

Actaeonella lucianoi Maury, 1930

(see Trochactaeon (Mexicotrochactaeon) burckhardti (Bose))

Trochactaeon (Trochactaeon) mammilatus (Hacobjan)

1976. Purpuractaeon mammilatus Hacobjan

Age. Late Coniacian. 

Occurrence. USSR (Armenian SSR).

Actaeonella marchmontensis Kollmann and Sohl

Age. Maastrichtian. 

Occurrence. Jamaica.

Trochactaeon (Trochactaeon) matensis (Fittipaldi)

1900. Actaeonella matensis Fittipaldi

1900. A. ellipsoides Fittipaldi

1953. Trochactaeon matensis Pchelintsev

1966. Actaeonella (Trochactaeon) matensis Lupu (Hacobjan, 1974

removed pi. 5, figs. 9, 11, and pi. 51, figs. 9, 10, 12 of Trochactaeon matensis Pchelintsev to Mesotrochactaeon ellipsoides 

(Fittipaldi))

Age. Cenomanian-Turonian.

Occurrence. Italy (Campobasso); USSR (Transcaucasus); Romania.

Trochactaeon (Mexicotrochactaeon) melonensis Sohl and Kollmann

Age. Maastrichtian. 

Occurrence. Puerto Rico.

Trochactaeon (Neocylindrites) minutus Stoliczka

1868. Trochactaeon minutus Stoliczka

Age. Campanian-Maastrichtian. 

Occurrence. Southern India.

Trochactaeon (Trochactaeon) mirabilis Pchelintsev

1954. Trochactaeon mirabilis Pchelintsev

Age. Early Senonian. 

Occurrence. USSR (Armenian SSR).

Trochactaeon (Mexicotrochactaeon) nelsoni Sohl and Kollmann

Age. Middle to late Maastrichtian. 

Occurrence. Puerto Rico.

Trochactaeon (Trochactaeon) obesus Pchelintsev

1954. Trochactaeon obesus Pchelintsev

Age. Coniacian.

Occurrence. USSR (Transcaucasus).

Actaeonella obliquestriata Stoliczka, 1859

(see Actaeonella laevis laevis (Sowerby))
CATALOGUE OF ACTAEONELLID SPECIES 83

Trochactaeon (Trochactaeon) obtusa (Zekeli)

1852. Actaeonella obtusa Zekeli

1856. A, obtusa Bronn

?1899.^. obtusa d'Alessandri

1939. A. obtusa Delpey

1951. A. obtusa Pasic

1965. A. obtusa Homaoui and Raab

1976. Spiractaeon obtusa Hacobjan (Hacobjan included Trochactaeon

gracilis Pchelintsev, 1954; T. darwasensis Djalilov, 1964 (part);

and T. (Trochactaeon) lamarcki brandenbergensis Kollmann,

1967 as synonyms)

Age. Cenomanian-Santonian.

Occurrence. Austria; Italy; Yugoslavia; Lebanon; 

Israel; USSR (?Tadzhinsk SSR).

Actaeonella (Trochactaeon) occidentalis Bdse, 1906

(see Trochactaeon (Mexicotrochactaeon) coniformis (Bose) 1906)

Actaeonella cf. A. (Trochactaeon) occidentalis Maldonado-Koerdell, 1950

(see Trochactaeon (Neocylindrites) punctatus Maldonado-Koerdell)

Actaeonella olivaeformis Cossmann

1896. Actaeonella olivaeformis Cossmann 

1902. A. olivaeformis Cossmann

Age. Garumnien (Maastrichtian). 

Occurrence. France (Ariege).

Trochactaeon (Neocylindrites) oliviformis (Coquand)

1865. Actaeonella oliviformis Coquand 

1887. A. oliviformis Mallada 

1909. Trochactaeon oliviformis Cossmann 

1949. Actaeonella oliviformis Battaler

Age. Aptian. 

Occurrence. Spain (Tereul).

Trochactaeon (Trochactaeon) ornatus Pchelintsev

1953. Trochactaeon ornatus Pchelintsev 

1976. Paleotrochactaeon ornatus Hacobjan

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Actaeonella orientale Pchelintsev

1960. Actaeonella orientale Pchelintsev

Age. Early Senonian.

Occurrence. USSR (eastern Siberia, Sikhote-Alin).

Actaeonella ornata Pchelintsev

1953. Actaeonella ornata Pchelintsev 

1972. A. (Pchelinsevella) ornata Djalilov 

1974. Actaeonella ornata Hacobjan

1976. A. ornata Hacobjan

1977. A. (Pchelinsevella) ornata Djalilov

Age. Cenomanian.

Occurrence. USSR (Transcaucasus, Armenian SSR).

Actaeonella oviformis Gabb

1869. Actaeonella oviformis Gabb 

1926. A. oviformis Stewart

Age. Turonian. 

Occurrence. California.

The following have been assigned to Actaeonella oviformis but should be placed elsewhere:

1954. Actaeonella oviformis Popenoe (a Trochactaeon) 

1958. A. oviformis Anderson (see Trochacateon (Trochaetaeon) packardl) 

1979. A. oviformis Jones, Sliter, and Popenoe (see Trochactaeon (Trochactaeon) packardi Anderson)

Trochactaeon (Trochactaeon) packardi (Anderson)

1902. Actaeonella oviformis Anderson

1916. A. oviformis Packard

1922. A. oviformis Packard

1958. A. oviformis Anderson

1958. A. packardi Anderson

1979. A. oviformis Jones, Sliter, and Popenoe

Age. Turonian. 

Occurrence. California.

Trochactaeon (Mexicotrochactaeon) palmeri Sohl and Kollmann

Age. Middle to late Maastrichtian. 

Occurrence. Jamaica; Cuba; Puerto Rico.

Trochactaeon (Trochactaeon) ouremensis Choffat

1901. Trochactaeon giganteus var. ouremensis Choffat

1974. Mesotrochactaeon ouremensis Hacobjan

1976. M. ouremensis Hacobjan

Age. Turonian.

Occurrence. Portugal; USSR (Armenian SSR).

Trochactaeon (Trochactaeon) ovalis Pchelintsev

1953. Trochactaeon ovalis Pchelintsev

1974. Mesotrochactaeon ovalis (Pchelintsev) part, Hacobjan, 1974

(pi. 49, fig. 5, pi. 50, figs. 1, 3, 4, to Mesotrochactaeon ellipsoides (Fittipaldi))

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Actaeonella oviformis Gabb of Packard 1916, Anderson 1938

(see Trochactaeon (Trochactaeon) packardi (Anderson))

Trochactaeon parvus Stanton, 1947

(see Trochactaeon (Neocylindrites) cumminsi Stanton)

Trochactaeon (Trochactaeon) parva Blanckenhorn

1890. Actaeonella parva Blanckenhorn 

1927. A. (Trochactaeon) parva Blanckenhorn

Age. Cretaceous. 

Occurrence. Syria; Israel.

Trochactaeon? patagonica (Feruglio)

1936. Actaeonella patagonica Feruglio

Age. Senonian.

Occurrence. Argentina (Patagonia).

Trochactaeon pamiricum Pchelintsev

1960. Trochactaeon pamiricum Pchelintsev

Age.-?
84 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Occurrence. USSR (Pamir Mountains).

Actaeonella pchelincevi Djalilov

1964. Actaeonella pchelincevi Djalilov

1972. A. (Pchelincevella) pchelincevi Djalilov

1977. A. (P.) pchelincevi Djalilov

Age. Late Cenomanian.

Occurrence. USSR (Darvaza, Turkmen SSR).

Trochactaeon (Trochactaeon) pcelinsevi (Hacobjan)

1963. Purpuractaeon pcelinsevi Hacobjan

1972. P. pcelinsevi Hacobjan

1974. P. pcelinsevi Hacobjan

Age. Late Coniacian. 

Occurrence. USSR (Armenian SSR).

Actaeonella pecosensis Stanton

1947. Actaeonella pecosensis Stanton

Age. Albian. 

Occurrence. Texas.

Actaeonella praecaucasica Aliev

1958. Actaeonella praecaucasica Aliev

1959. A. praecaucasica Aliev

1963. A. praecaucasica Aliev

1972. A. (Pchelinsevella) praecaucasica Djalilov

1977. A. (Pchelinsevella) praecaucasica Djalilov

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Actaeonella (Trochactaeon) planilateralis Bose, 1906

(see Trochactaeon (Mexicotrochactaeon) coniformis (Bose))

Actaeonella (Trochactaeon) planilateralis Bose, Maldonado-Koerdell, 1950

(see Trochactaeon (Neocylindrites) punctatus (Maldonado-Koerdell))

Actaeonella pompei Maury 1930

(see Trochactaeon (Mexicotrochactaeon) burckhardti (Bose))

Actaeonella (Trochactaeon) potosiana Bose, 1906

(see Trochactaeon (Mexicotrochactaeon) coniformis (Bose), 1906

Actaeonella praesulcata Aliev, 1959

(see Sogdianella kurdistanica( Aliev)

Actaeonella (Trochactaeon) prisca (Deshayes) Blanckenhorn, 1927 

(affinities unknown)

Trochactaeon (Trochactaeon) projectiliformis Kollmann

1967. Trochactaeon (Trochactaeon) projectiliformis Kollmann

Age. Coniacian-early Santonian. 

Occurrence. Austria.

Actaeonella (Trochactaeon) pruniformis Bose, 1923

(see Trochactaeon (Mexicotrochactaeon) burckhardti (Bose), 1923;

excludes Trochactaeon pruniformis Hacobjan, 1976,

from the Cenomanian of Armenian SSR)

Trochactaeon (Trochactaeon) pseudocylindraceus Pchelintsev

1953. Trochactaeon pseudocylindraceus Pchelintsev

1954. T. pseudocylindraceus Pchelintsev

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Trochactaeon (Neocylindrites) punctatus (Maldonado-Koerdell)

1950. Actaeonella cf. A. (Trochactaeon) coniformis MaldonadoKoerdell

1950. A. (Trochactaeon) acutissima punctata Maldonado-Koerdell 

1950. A. cf. A. (Trochactaeon) occidentalis Maldonado-Koerdell 

1950. A. (Trochactaeon) planilateralis Maldonado-Koerdell

Age. M aastr ichtian. 

Occurrence. Mexico (Chiapas).

Trochactaeon quadratus Pchelintsev, 1954

(see Trochactaeon (Neotrochactaeon) ventricosa Hacobjan, 1976)

Actaeonella (Trochactaeon) quadriplicata Bose, 1923

(see Trochactaeon (Mexicotrochactaeon) burckhardti (Bose))

Trochactaeon (Trochactaeon) regularis Pchelintsev

1953. Trochactaeon regularis Pchelintsev

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Trochactaeon (Trochactaeon) renauxianus (d'Orbigny)

1842. Actaeonella renauxiana d'Orbigny

Age. Turonian.

Occurrence. France; Yugoslavia; Lebanon; Romania.

The species listed below have been assigned to Trochactaeon (Trochactaeon) renauxiana but should be placed 

elsewhere:

1852. Actaeonella renauxiana Zekeli (see Trochactaeon (Trochactaeon)

sanctaecrucis Futterer, fide Kollmann, 1967) 

1884. A. renauxiana Quenstedt (see T. (T.) sanctaecrucis Futterer,

fide Kollmann, 1967) 

1896. Trochactaeon renauxianus Cossmann (see T. (T.) sanctaecrucis

Futterer, fide Kollmann, 1967) 

71906. Actaeonella cf. A. renauxiana Oppenheim 

1939. A. renauxiana Delpey 

1957. Trochactaeon renauxianus Pejovic'(see T. (T.) giganteus

(Sowerby), fide Kollmann, 1967) 

1959. Actaeonella (Trochactaeon) renauxiana Pokorny (see T. (T.)

sanctaecrucis Futterer, fide Kollman, 1967)

1966. A. (Trochactaeon) aff. A. (T.) renauxiana Todirit|-Mihailescu

1967. A. (Trochactaeon) renauxiana Rahman 

1972. Trochactaeon renauxiana Hacobjan

Actaeonella roashensis Abbass

1963. Actaeonella (Actaeonella) roashensis Abbass

Age. Santonian. 

Occurrence. Egypt.

Actaeonella robinsoni Sohl and Kollmann

Age. Maastrichtian. 

Occurrence. Jamaica', Mexico.

Trochactaeon robustus Pchelintsev, 1954

(see Trochactaeon pruniformis Bose, fide Hacobjan, 1974)
CATALOGUE OF ACTAEONELLID SPECIES 85

Trochactaeon (Mexicotrochactaeon) rossi Sohl and Kollman

1906. Actaeonella (Trochactaeon) aff. gigantea Sowerby, Bose 

1968. A. coniformis Bose, Myers (part)

Age. Middle Maastrichtian. 

Occurrence. Mexico; Jamaica; Cuba.

Actaeonella rotundata Zekeli, 1852

(to Vernedia of the Itieriidae)

Trochacteaon rugosus G. Aliev, 1959

(to Eotrochactaeon fide Hacobjan 1972, 

to the Itieriidae Kollrnann and Sohl, 1980)

Actaeonella rustica Anderson, 1958

(see Trochactaeon (Trochactaeon) frazierensis Anderson)

Trochactaeon (Trochactaeon) salomonis Fraas, 1867

1867. Actaeonella salomonis Fraas (= Trochactaeon (Trochactaeon)

lamarcki lamarcki) 

1903. A. salomonis Dacque (= T. (T.) lamarcki lamarcki, fide Kollmann, 1967; Hacobjan, 1974, removed fig. 6 to Mesotrochactaeon tovusensis Hacobjan) 

1903. Trochactaeon salomonis Fortau 

1912. Actaeonella (Trochactaeon) salomonis Pervinquiere (= Mesotrochactaeon longisulcus Hacobjan, 1974) 

1927. A. (T.) salomonis Blanckenhorn 

1965. Actaeonella? salomonis Homaoui and Raab (see Trochactaeon

(Trochactaeon) longisulcus Hacobjan)

Age. Late Turonian.

Occurrence. Israel; Lebanon; Egypt (Abu Roash).

Trochactaeon (Trochactaeon) sanctaecrucis (Futterer)

1852. Actaeonella Renauxiana Zekeli

1852. A. glandiformis Zekeli

1853. A. Renauxiana Reuss

1865. A. gigantea Stoliczka

1884. A. renauxiana Quenstedt

1892. A. Sanctae-Crucis Futterer

1895. A. Sanctae-Crucis Bohm

1896. Trochactaeon Renauxianus Cossmann

1959. Actaeonella (Trochactaeon) renauxiana Pokorny

Age. Santonian.

Occurrence. Italy (Venetian Alps); Austria.

Trochactaeon (Trochactaeon) scalatus (Schnarrenberger)

1901. Valuta scalata Schnarrenberger

Age. Cenomanian.

Occurrence. Italy (Venetian Alps).

Actaeonella (Volvulina) schiosensis Bbhm, 1895

(see Actaeonella caucasica schiosensis)

[assigned to A. (Pchelincevella) schiosensis by Djalilov, 1972, 1977, 

and to Acteonella schiosensis by Hacobjan, 1974, 1976])

Trochactaeon? schiosensis (Bbhm)

1895. Conus schiosensis Bohm

Age. Late Cenomanian-early Turonian. 

Occurrence. Italy (Venetian Alps).

1953.

Trochactaeon (Trochactaeon) schirabadensis Pchelintsev

Trochactaeon schirabadensis Pchelintsev

1953. T. asiaticus Pchelintsev

1977. Spiractaeon schirabadensis Djalilov

Age. Santonian.

Occurrence. USSR (Tadzhinski depression).

Actaeonella silvae Maury, 1925

(see Trochactaeon (Mexicotrochactaeon) burckhardti (Bose))

Trochactaeon similis Pchelintsev

1954. Trochactaeon similis Pchelintsev

Age. Early Cenomanian. 

Occurrence. USSR (Armenian SSR).

Actaeonella sinaiensis Abbass

1963. Actaeonella sinaiensis Abbass

Age. Santonian. 

Occurrence. Egypt (Sinai).

Trochactaeon (Trochactaeon) solidus Pchelintsev

1954. Trochactaeon solidus Pchelintsev

Age. Late Senonian.

Occurrence.-USSR (Armenian SSR).

Trochactaeon (Trochactaeon) speciosus Pchelintsev

1954. Trochactaeon speciosus Pchelintsev

Age. Coniacian.

Occurrence. USSR (Transcaucasus).

Trochactaeon stabilis Dvali, 1966

(to Eotrochactaeon fide Hacobjan, 1972, 1976, 

to the Itieriidae Kollmann and Sohl, 1980)

Trochactaeon subangustatus Pchelintsev 1953

(see Paleotrochactaeon crisminensis (Choffat)

Trochactaeon subburckhardti Pchelintsev 1954

(part to Trochactaeon pruniformis Hacobjan not Bose, 

part to Trochactaeon excelsus Pchelintsev, fide Hacobjan, 1976)

Trochactaeon subglandiformis Pchelintsev, 1954

(see Neotrochactaeon glandiformis Hacobjan, 1974)

Actaeonella sublaevis Briart and Cornet

1865. Actaeonella sublaevis Briart and Cornet 

1939. A. (Volvaria) sublaevis Marliere 

1972. A. (Pchelintsevella) sublaevis Djalilov

1976. Actaeonella sublaevis Kollmann

1977. Acteonella (Pchelintsevella) sublaevis Djalilov

Age. Late Albian.

Occurrence. Belgium; Austria; Texas (herein).

Actaeonella cf. A. sublaevis Briart and Cornet, Sohl, and Kollmann

Age. Albian to Cenomanian. 

Occurrence. New Mexico.

Trochactaeon subotusus Pchelintsev, 1953

(fig. 16 to Mesotrochactaeon tovusensis 

and remainder to Mesotrochactaeon ellipsoides by Hacobjan, 1976)

Trochactaeon subovatus Pchelintsev 1954

(see Neotrochactaeon glandiformis Zekeli, fide Hacobjan, 1976)
86 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Trochactaeon (Trochactaeon) subpruniformis Pchelintsev

1954. Trochactaeon subpruniformis Pchelintsev 

1974. T. pruniformis Bose, Hacobjan

Age. Coniacian.

Occurrence. USSR (Armenian SSR).

Trochactaeon (Trochactaeon) subrenauxi Pchelintsev

1953. Trochactaeon subrenauxi Pchelintsev

1954. T. subrenauxi Pchelintsev

Age. Cenomanian.

Occurrence. USSR (Armenian SSR).

Actaeonella syriaca Conrad, Fraas, 1867

(see Trochactaeon (Trochactaeon) lamarcki lamarcki Sowerby)

Trochactaeon (Trochactaeon) szontaghi (Todiriti-Mihailescu)

1966. Actaeonella szontaghi Todirit|-Mihailescu

Age. Late Santonian-early Campanian. 

Occurrence. Romania.

Actaeonella tamandarensis Maury

1930. Actaeonella tamandarensis Maury

Age. Late Turonian-Santonian. 

Occurrence. Brazil (Pernambuco).

Actaeonella terebellum Cossmann, 1896

(see Actaeonella laevis laevis (Sowerby))

Trochactaeon (Mexicotrochactaeon) teaensis Sohl and Kollmann

Age. Middle to late Maastrichtian. 

Occurrence. Jamaica; Puerto Rico; Mexico.

Actaeonella terterensis Aliev

1961. Actaeonella terterensis Aliev 

1963. A. terterensis Aliev 

1972. A. (Pchelincevella) terterensis Djalilov 

1977. Acteonella (Pchelincevella) terterensis Djalilov [assigned to 

Actaeonella uchauxensis by Hacobjan, 1976]

Age. Coniacian.

Occurrence. USSR (Transcaucasus).

Actaeonella torrubiai Vidal

1921. Actaeonella torrubiai Vidal 

1949. A. torrubiai Dataller

Age. Santonian. 

Occurrence. Spain (Gerona).

Actaeonella toucasiana d'Orbigny, 1950

(see Trochactaeon (Trochactaeon) goldfussi)

Trochactaeon (Trochactaeon) tovusensis (Hacobjan)

1953. Trochactaeon subobtusus Pchelintsev (part) 

1976. Mesotrochactaeon tovusensis Hacobjan

Age. Turonian.

Occurrence. USSR (Armenian SSR).

Trochactaeon transylvanicus Hojnos, 1921

(see Trochactaeon (Trochactaeon) giganteus subglobosus (Miinster))

Actaeonella truncata Stoliczka

1868. Actaeonella truncata Stoliczka

Age. C ampanian-Maastrichtian. 

Occurrence. Southern India.

Trochactaeon (Trochactaeon) truncatus Stoliczka

1868. Trochactaeon truncatus Stoliczka 

1902. T. truncatum Rehbinder

Age. Arialoor Group; Campanian-Maastrichtian. 

Occurrence. Southern India; USSR (Astrakhan).

Trochactaeon (Sevanella) tuberculatus Hacobjan

1974. Sevanella tuberculata Hacobjan 

1976. S. tuberculata Hacobjan

Age. Coniacian.

Occurrence. USSR (Armenian SSR).

Actaeonella tumida Makarenko

1961. Actaeonella tumida Makarenko

Age. Paleocene(?) (reworked). 

Occurrence. USSR (Crimea).

Trochactaeon (Trochactaeon) tumidus Pchelintsev

1953. Trochactaeon tumidus Pchelintsev

1954. T. tumidus Pchelintsev

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Actaeonella uchauxensis Cossmann

(see Actaeonella caucasica grossouvrei Cossmann)

1896. Actaeonella uchauxensis Cossmann

1974. Acteonella uchauxensis Hacobjan

1976. Actaeonella uchauxensis Hacobjan

Age. Turonian.

Occurrence. France (Drome); USSR (Armenian SSR).

Actaeonella (Trochactaeon) variabilis Bose, 1906

(see Trochactaeon (Mexicotrochactaeon) coniformis (Bose):

Actaeonella (Trochactaeon) variabilis Daguin, 1927,

and A. (Trochactaeon) cf. variabilis Trechmann

= questionable placement)

Trochactaeon (Trochactaeon) variospiralis (Hacobjan)

1976. Spiractaeon variospiralis Hacobjan

Age. Coniacian.

Occurrence. USSR (Armenian SSR).

Trochactaeon (Trochactaeon) vasmikuchensis Djalilev

1964. Trochactaeon vasmikuchensis Djalilev 

1977. Mesotrochactaeon vasmikuchensis Djalilev

Age. Cenomanian.

Occurrence. USSR (Darvaza, Turkmen SSR).

Trochactaeon (Trochactaeon) vediensis (Hacobjan)

1963. Purpuractaeon vediensis Hacobjan

1974. P. vediensis Hacobjan

1976. P. vediensis Hacobjan
GENERA AND SPECIES EXCLUDED FROM THE ACTAEONELLIDAE 87

Age. Coniacian.

Occurrence. USSR (Armenian SSR).

Neotrochactaeon ventricosa (Hojnos) Hacobjan, 1976

(see Trochactaeon (Trochactaeon) gtganteus subglobosus (Minister))

Actaeonella voluta d'Orbigny, 1850

(see Trochactaeon (Trochactaeon) lamarcki lamarcki (Sowerby))

Trochactaeon (Trochactaeon) woods! (Rennie)

1906. Actaeonella (Trochactaeon) sp. Woods 

1930. A. (T.) woodsi Rennie

Age. Santonian-Campanian (Africa); early Maastrichtian (Puerto Rico).

Occurrence. South Africa (Pondoland); Puerto Rico 

(San German area).

Trochactaeon (Mexicotrochactaeon) wrighti Sohl and Kollmann

Age. Maastrichtian. 

Occurrence. Jamaica.

Trochactaeon (Trochactaeon) cf. T. (T.) woodsi (Rennie) 

Sohl and Kollmann

Age. Early Campanian. 

Occurrence. Puerto Rico.

Trochactaeon (Trochactaeon) zekelii Pchelintsev

1954. Trochactaeon zekelii Pchelintsev

Age. Early Senonian. 

Occurrence. USSR (Armenian SSR).

Actaeonella zouparriensis Choffat, 1901

(see Actaeonella caucasica zouparriensis}

Many specimens have been cited in the literature and 

assigned to either Actaeonella or Trochactaeon but have 

not merited assignment to a described species. Some 

specimens do not belong in either genus; others only repeat 

occurrences based upon well-founded species. In both instances, these specimens contribute nothing to the 

geographic or stratigraphic data presented here. Listed 

below, however, are several that bear special mention, as 

they extend the geographic range or stratigraphic occurrence of the genus and are included on the accompanying 

maps:

Actaeonella sp. Delpey 

1939. Actaeonella sp. Delpey

Age. Albian. 

Occurrence. Lebanon.

Actaeonella sp. Duboul

1947. Actaeonella sp. Doboul

Age. Early Campanian. 

Occurrence. France (Provence).

Neocylindrites(?) sp. (Dietrich)

1914. Actaeonella (Trochactaeon) Dietrich

Age. Aptian.

Occurrence. East Africa (Tenadaguru).

Actaeonella n. sp. Darteville and Brebion

1932. Actaeonella? sp. Riedel

1956. Actaeonella n. sp. Darteville and Brebion

Age. Santonian. 

Occurrence. Cameroons.

Trochactaeon sp. Spengler

1913. Trochactaeon sp. Spengler

Age. Campanian.

Occurrence. India (Province of Assam).

Actaeonella sp. Sohl and Kollmann

Age. Middle Albian. 

Occurrence. Puerto Rico.

Trochactaeon (Neocylindrites) sp. A, Sohl and Kollmann

Age. Maastrichtian. 

Occurrence. Puerto Rico.

Trochactaeon (Neocylindrites) sp. B, Sohl and Kollmann

Age. Late Albian. 

Occurrence. New Mexico.

Actaeonella sp. Yii

1982. Actaeonella sp. Yii

Age. Aptian?.

Occurrence. Qinqhai Province, Tibet.

GENERA AND SPECIES EXCLUDED 

FROM THE ACTAEONELLIDAE

White (1888, p. 201-202) diagnosed the genus Cylindritella as follows:

Shell subcylindrical; spire short or moderately extended; aperture 

narrow, more or less distinctly channeled in front but not beaked, and 

probably always notched narrowly behind; outer lip simple or everted 

but anteriorly the columella bears three more or less distinct folds or 

ridges, the anterior one of which is very oblique, and forms the right 

side of the anterior channel; the obliquity of each succeeding fold 

decreases posteriorly so that the posterior one is sometimes nearly 

transverse. The surface is probably always smooth; the volutions of the 

spire are flat or slightly convex; the suture inconspicuous; and, in some 

cases at least, there is more or less callus deposited upon the spire 

about the posterior end of the aperture as it is sometimes in Oliva.

White (1888) and, subsequently, Zilch (1959) considered Cylindritella as an actaeonellid gastropod. Investigation of White's types, which are kept at the U.S. 

National Museum of Natural History, Washington, B.C., 

demonstrate that Cylindritella does not belong to the 

Opistobranchia but to the Neogastropoda, as was sug-
88 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

gested earlier by Kollmann (1967). This conclusion is 

based on the following observations:

1. The whorls of Cylindritella are posteriorly very narrow and acute in cut section. They show no subsutural ramp or any other signs of an exhalant siphon.

2. The internal molds of the whorls are anteriorly prolonged and narrow, showing that the shells had a 

very short siphonal canal.

3. The strength of the columellar plaits decreases from 

the posterior to the anterior.

Following Kollmann (1967), we suggest that Cylindritella should be transferred to the Marginellidae. We 

cannot decide here whether it is a distinct genus within 

this family or whether it belongs to one of the established genera.

Kollmann and Sohl (1980) have provided evidence that 

the genera Sogdianella Djalilov, 1972, Ovactaeonella 

Djalilov, 1972 (including Omphaloactaeonella Hacobjan, 

1972), Peruviella Olsson 1934, and Eotrochactaeon Hacobjan, 1972, all belong to the nerineacean family Itieriidae 

rather than to the Actaeonellidae where the original proposers placed them. The following listings reflect these 

reassignments:

Ovactaeonella agdjakendensis (Aliev)

1959. Actaeonella agdjakendensis Aliev

1963. A. agdjakendensis Aliev

1972. Ovactaeonella agdjakendensis Djalilov

1976. Omphaloacteonella agjakendensis Hacobjan

1977. Ovactaeonella agjakendensis Djalilov

Age. Cenomanian.

Occurrence. USSR (Azerbaijan SSR).

Ovactaeonella anchieti (Choffat), 1888

(see Peruviella do/z'um (Roemer))

Ovactaeonella armenica (Pchelintsev)

1953. Actaeonella armenica Pchelintsev

1972. Ovactaeonella armenica Djalilov

1977. O. armenica Djalilov (assigned to Omphaloacteonella ovata

(Pchelintsev) by Hacobjan, 1976) 

1982. Ovactaeonella dolium Konovalov and Poyarkova

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Eotrochactaeon azerbaidjanensis (Aliev)

1961. Trochactaeon azerbaidjanensis Aliev

1963. T. azerbaidjanensis Aliev

1972. Eotrochactaeon azerbaidjanensis Hacobjan

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Ovactaeonella azerbaidjanica (Aliev)

1959. Actaeonella azerbaidjanica Aliev

1972. Ovactaeonella azerbaidjanica Djalilov

1976. Omphaloacteonella azerbaidjanica Hacobjan

1977. Ovactaeonella azerbaidjanica Djalilov

Age. Early Coniacian.

Occurrence. USSR (Transcaucasus).

Ovactaeonella devii (Dvali)

1966. Actaeonella devii Dvali

1967. Acteonella georgica Gambashidze 

1972. Ovactaeonella devii Djalilov 

1972. Omphaloacteonella devii Hacobjan

1976. Ovactaeonella devii Djalilov

1977. Omphaloacteonella devii Hacobjan

Age. Cenomanian or Turonian. 

Occurrence.-USSE (Georgian SSR).

Peruviella dolium (Roemer)

1849. Actaeonella dolium Roemer

1852. A. dolium Roemer

1888. A. anchietai Choffat

1928. Volvulina texana Adkins

1947. Actaeonella dolium Stanton

1956. A. anchietai Darteville and Brebion

1976. Omphaloacteonella anchietai Hacobjan

1979. Peruviella dolium Kollmann and Sohl

1982. Ovactaeonella dolium Konovalov and Poyarkova

Age. Albian.

Occurrence. Texas; Mexico (State of Chihuahua); 

Angola (Dombe-Grande).

Eotrochactaeon dubertreti Hacobjan

1972. Eotrochactaeon dubertreti Hacobjan

Age. Cenomanian.

Occurrence. USSR (Transcaucasus).

Ovactaeonella ferjanica Djalilov

1972. Ovactaeonella ferjanica Djalilov

1977. O. ferjanica Djalilov

1982. Ovactaeonella dolium Konovalov and Poyarkova

Age. Coniacian.

Occurrence.-USSR (Tadzhik SSR-Fergana, Kuraminski Ronge).

Sogdianella gagii (Dvali)

1966. Actaeonella gagii Dvali 

1972. A. (Sogdianella) gagii Djalilov 

1974. A. gagii Hacobjan

1976. Omphaloacteonella gagii Hacobjan

1977. Actaeonella (Sogdianella) gagii Djalilov

1982. Ovactaeonella dolium Konovalov and Poyarkova

Age. Turonian.

Occurrence. USSR (Armenian SSR).

Actaeonella georgica Gambashidze, 1967

(see Ovactaeonella devii (Dvali)

Peruviella gerthi Olsson

1928. Actaeonella cf. A. gigantea Gerth

1934. Peruvia gerthi Olsson

1936. Trochactaeon sergipensis Maury

1944. Peruviella gerthi Olsson

1979. P. gerthi Kollmann and Sohl
GENERA AND SPECIES EXCLUDED FROM THE ACTAEONELLIDAE 89

1982. Ovactaeonella dolium Konovalov and Poyarkova

Age. Middle and late Albian.

Occurrence. Peru (Amotape region); Brazil (Sergipe 

Province).

Actaeonella cf. gigantea Gerth 1928

(see Peruviella gertht Olsson)

Eotrochactaeon gorindjukensis (Dvali)

1966. Trochactaeon gorindjukensis Dvali 

1972. Eotrochactaeon gorindjukensis Hacobjan

Age. C enomanian.

Occurrence. USSR (Transcaucasus).

Sogdianella grandis (Pchelintsev)

1928. Actaeonella grandis Pchelintsev 

1953. A. grandis Pchelintsev 

1972. A. (Sogdianella) grandis Djalilov

1977. Acteonella (Sogdianella) grandis Djalilov (assigned to Actaeonella crassa Hacobjan, 1976)

Age. C enomanian.

Occurrence. USSR (Transcaucasus).

Sogdinella khalilovi Aliev

1959. Actaeonella khalilovi Aliev

1963. A. khalilovi Aliev

1972. A. (Sogdianella) khalilovi Djalilov

1976. Acteonella khalilovi Hacobjan

1977. A. (Sogdianella) khalilovi Djalilov

Age. Cenomanian (fide Hacobjan, 1976) Coniacian 

(fide Djalilov, 1977). 

Occurrence. USSR (Transcaucasus).

Ovactaeonella konstantini (Dvali)

1966. Actaeonella konstantini Dvali

1972. Ovactaeonella konstontini Djalilov

1982. Ovactaeonella dolium Konovalov and Poyarkova

Age. Cenomanian.

Occurrence. USSR (Azerbaijan SSR).

Sogdianella kurdistanica (Aliev)

1959. Actaeonella kurdistanica Aliev 

1963. A kurdistanica Aliev 

1972. A. (Sogdianella) kurdistanica Djalilov 

1977. A. (S.) kurdistanica Djalilov

Age. Cenomanian.

Occurrence. USSR (Azerbaijan SSR, Transcaucasus).

Ovactaeonella mardakertensis (Aliev)

1961. Actaeonella mardakertensis Aliev

1963. A. mardakertensis Aliev

1972. Ovactaeonella mardakertensis Djalilov

1976. O. mardakertensis Hacobjan

1977. O. mardakertensis Djalilov

Age. Turonian.

Occurrence. USSR (Transcaucasus).

Ovactaeonella ovata (Pchelintsev)

1953. Actaeonella ovata Pchelintsev

1972. Ovactaeonella ovata Djalilov

1974. Actaeonella ovata Hacobjan

1976. Omphaloacteonella ovata Hacobjan

1977. Ovactaeonella ovata Djalilov

1982. Ovactaeonella dolium Konovalov and Poyarkova

Age. Turonian.

Occurrence. USSR (Transcauscasus).

Sogdianella peruviana (Olsson)

1934. Actaeonella peruviana Olsson

1979. Sogdianella peruviana Kollmann and Sohl

Age. Albian.

Occurrence. Peru (Amotape region).

Sogdianella aff. S. peruviana (Olsson)

1928. Actaeonella (Volvulina) spec. Gerth

1979. Sogdianella aff. S. peruviana Kollmann and Sohl

Age. Maastrichtian. 

Occurrence. Peru (Paita region).

Sogdianella praesupernata (Djalilov)

1964. Actaeonella praesupernata Djalilov

1972. A. (Sogdianella) praesupernata Djalilov

1977. A. (S.) praesupernata Djalilov

Age. Albian.

Occurrence.-USSR (Darvaza, Turkmen SSR).

Vernedia repelini (Repelin)

1902. Actaeonella repelini Repelin

Age. Cenomanian. 

Occurrence. France.

Trochactaeon sergipensis Maury, 1936

(see Peruviella gertht Olsson)

Sogdianella supernata (Pchelintsev)

1953. Actaeonella supernata Pchelintsev

1954. A. supernata Pchelintsev

1966. A. supernata Lupu

1972. A. (Sogdianella) supernata Djalilov

1974. Acteonella supernata Hacobjan

1976. Omphaloacteonella supernata Hacobjan

1977. Acteonella (Sogdianella) supernata Djalilov

Age. Cenomanian-early Turonian.

Occurrence. USSR (Armenian SSR, Transcaucasus).

Sogdianella tagarensis (Arustamov)

1966. Actaeonella tagarenensis Arustamov 

1972. A. (Sogdianella) tagarensis Djalilov

Age. Cenomanian.

Distribution. USSR (Transcaucasus).

Sogdianella tenuiflexa (Dvali)

1966. Actaeonella tenuiflexa Dvali 

1972. A. (Sogdianella) tenuiflexa Djalilov 

1976. Omphaloacteonella tenuiflexa Hacobjan
90 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Age. Turonian.

Occurrence. USSR (Georgian SSR).

Sogdianella laevis zekelii (Kollmann)

1852. Actaeonella laevis Zekeli (part) 

1965. A. laevis zekelii Kollmann 

1972. A. (Sogdianella) zekelii Djalilov

Age. Late Coniacian. 

Occurrence. Austria.

Eotrochactaeon zumoffeni (Delpey)

1939. Actaeonella zumoffeni Delpey

Age. Aptian. 

Occurrence. Lebanon.

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INDEX

[Italic page numbers indicate major references]

Page

abbreviata, Actaeonella 75

Actaeonella (Trochactaeon) 75

Nennea 75

abbreviatus, Trochactaeon 75

Trochactaeon (Neocylindrites) 75

absaloms, Trochactaeon 48

Trochactaeon (Neocylindrites) 75

abstract 1

acknowledgments 2

Actaeo n .4

Actaeon giganteus 80

Actaeon tornatlhs 2

burrowing behavior 5

Actaeonell crassa 78

gracihs 78

Actaeonella 9, 68; pis. 18-23,4, 20

abbreviata 75

acummata 75, 76

agdjahendensis 88

agncolai 20, 75

anch letai 88

armenica 88

azerbaidjamca 88

baconica 20, 75

beyrich i 76

6orneensis 6, 35, 67, 68f, 76; pis. 19, 20 

6rtarti -. 76

6r!ggs;' 35, 71, 72, 72£ 76, pis. 20, 21 

browm 24, 67, 67/i 76, pi. 18

caucasica 64, 76, 77

caucasica caucasica 76

caucasica grossouvrei 76, 77, 81, 86

caucasica schiosensis 77, 85

caucasica styriaca 77, 81

caucasica zouparriensis 77, 87

conform is 77

arnica 76, 77 

coniformis 56, 57, 58, 62, 63, 85; pi. 16

coquiensis 35, 70, 70f, 78; pis. 2, 20, 21

crassa 73, 77, 78, 81, 82, 89 

cretacea 78

cubensis 9, 35, 65, 72, 73f, 78; pis. 17, 22, 23 

cyhndracea 78

d 'Orbigny 63

decorata 65, 78

delgadoi 66, 66f, 77, 79,81; pi. 18

devil 88

dolium 88

du bertreti 79

duckettsensis 35, 69, 69f, 72, 79; pis. 19, 20 

elhpsoides 79,82

elliptica 79

elongata 70, 79

/ a&a 79 

frazierensis 50, 79

fusiformis 20, 22, 79

gagii 88

georgica 88

ghazirensis 80

gigantea 79, 80, 85

glandiformis 80, 85

goldfussi 80

gracilis 66, 79, 81

grandis 89

grossouvrei 64, 76, 77, 79, 81

inconstans 58

inflata 73, 81

involuta 81

jicarensis 7, 35, 69, 70/; 81; pis. 19, 20, 22, 23 

khalilovi 65, 89

konstantiti 89

kurdistanica 89

/ae[, [s 9,64,70, 72, 79,81,90

/aeuis 81, 82, 86 

lamarck i 80

tonga 82 

luciana 59

/uci'ano 58, 58/i 76, 82

marchmontensis 35, 77, 7If, 72, 82, pi. 21 

mardakertensis 89

matensis 82

obliquecostata 81,82

obtusa 83

occidentals 58

olwaeformis 83

olivifbrmis 20,83

orientate 83

ornata 65, 83

oi>afa 64, 89 

oviformis 35, 51, 52, 68, 83; pi. 18

packardi 51, 83

parva 83

parvus -. 53

patagonica 83

pchelincevi 84 

peconsensis 24,65

pecosensis 65, 65f, 66, 84; pis. 18, 20

peruviana 89

planilateralis 58

pompei 58, 58/i 59, 76,84 

praecaucasica 84

praesulcata 84

praesupernata 89

renauxiana 47, 80, 84, 85

repehni 89

roashensis 84

robinsoni 35, 65,69, 72, 73, 84; pis. 17,22,23 

rotundata 85

rustica 50, 79, 85

salomonis 85

Sanctae-Crucis 85

schiosensis 77, 85

silvae 85

sinaiensis 85

subglobosa 80

sublaevis 66, 67, 85

supernata 64, 89

syriaca 81, 86

szontagh i 86

tagarensis 89

tamanderensis 86

teaensis 68

tenuiflexa 89

terebellum 81,86

terebrellu m 64

terterensis 86

torrubiai 86

toucasiana 80, 86

truncata - 86

fa m ^a 86

uchauxensis 64, 76, 81, 86

valuta 87

zouparriensis 87

zumoffeni 90

sp. 6, 24,87; pis. 1, 2,22 

sp. B 74; pi. 22

sp. C 74; pi. 2 

spp. 74 

(Actaeonella) caucasica 77

roashensis 84

(Pchelincevella) caucasica 76

crassa 78

delgadoi 66, 79

/a&a 79 

gracilis 78

grossouvrei 76

pchelincevi 84

schiosensis 77, 85

styriaca 7 7

terterensis 86

zouparrifnsis 77

ornata 83

praecaucasica 84

(Pchekntsevella) sublaevis 85

(Sogdianella) gagii 88

grandis 89

khalilovi 89

A urdistanica 89

praesupernata 89

supern ata 89

tagarensis 89

tenuiflexa 89

2e/fee/u 90 

(Trochactaeon) abbreviata 75

acutissima .48, 56, 57, 75, 77

punctata 54, 56, 58, 77, 84

angustata 75

6reuis 48, 56,76, 77

burckhardti .47, 58, 76

collignoni 77

conica 76, 77

coniformis .48, 56, 77, 84

conoidea 78

cordeiroi 78

cylindracea 52, 78

gigantea .48, 58, 79, 80

gigantea 80

globosa 80

subglobosa 80

uen tricosa 80

goldfussi 80,81

Humboldti 58, 76,81

humei 81

inconstans 48, 56, 77, 81

irregularis 48, 56, 77, 81

lamarcki 82

matensis 82

occidentalis .48, 56, 77, 83

parva 83

parvus 78

planilateralis .48, 54, 56, 77, 78, 84

potosiana 48, 56, 77, 84

prisca 84

pruniformis 58, 84

quadnplicata 58, 76, 84

97
98 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

quadnphcatus 47

renauxiana 84, 85

salomonis 82, 85

subangustatus 78

vanabihs 48, 56, 77, 86

wood si 49, 87

sp. 49, 58,87 

(Volvaria) sublaevis 66, 85

(Volvulina) 89

inflata 81

laevis 77,81

schiosensis 77, 85

(Actaeonella) caucasica, Actaeonella 77

roashensi, Actaeonella 84

Actaeonelhdae, catalogue 74

diversity over time 10

evolution 20

genera and species excluded 87

phylogeny 2lf

Actaeonellina 22

Acteonellaborneensis 67

decorata 78

georgica 88

gracilis 66

grossouvret 64, 66

lamarcki 82 

oviformis 51, 52, 68

salomonis 82

supern a ta 89

syr ia ca 82 

terebrellum 64

uchauxensis 64, 86 

valuta 82

(Pchelmceuella) delgadoi 66

terterensis 86 

(Sogdianella) grandis 89

supernata 89

acuminata, Actaeonella 75, 76

Trochactaeon (Trochactaeon) 75

acuminatus, Trochactaeon (Trochactaeonl 75

acutissima punctata, Actaeonella (Trochactaeon) -54, 56,

58,77,84

Actaeonella (Trochactaeon) 48, 56, 57, 75, 77

acutissima, Mexicotrochactaeon 56, 77

Trochactaeon 48

affinities with Indo-Pacific species 35 

Africa, West 20 

agbulakhensis, Trochactaeon 75

agdjakendensis, Actaeonella 88

Ovactaeonella 88 

Omphaloacteonella 88

agricolai, Actaeonella 20, 75

Aguirre, Central, Puerto Rico, collection

localities, index map 29f 

Alaska, localities 46 

Albian Limestone, Guatenala, localities 46 

ahbeglyensis, Trochactaeon 75

Trochactaeon I Trochactaeon I 75 

Alisitos Formation, Baja California 24 

Alpine Cretaceous 7 

Ampullina 7

anchietai, Actaeonella 88 

Omphaloacteonella 88 

Ovactaeonella 88

angustata, Actaeonella (Trochactaeon) 75

angustatus, Paleotrochactaeon 75

Trochactaeon 48, 75

Trochactaeon (Neocylindrites) 75

anomalous occurrences, Late Cretaceous 24 

Antillean islands 20, 24 

Aptian to Albian time interval 9, lOf 

Aptian Age rocks, absence of actaeonellids 24 

Aquilapa, State of Chiapa, Mexico, localities 46 

aquilerae, Arctostrea 34, 58

Arctostrea aquilerae zone. Mexico 58 

Armenian SSR 22 

armenica, Actaeonella 88

Ovactaeonella 88

armentcus, Trochactaeon 75

Trochactaeon (Trochactaeon) 75

arnaudi, Mesotrochactaeon 75

Trochactaeon 75

Trochactaeon (Trochactaeon) 75

artsachensis, Mesotrochactaeon 75

Trochactaeon (Trochactaeon) 75

Assam Province, India 20 

asiaticus, Trochactaeon 85

Trochactaeon (Trochactaeon) 75 

associations, lagoonal 7 

Astrocoenia, coral 7 

Austria, associations in shales 7 

atschadjurensis, Trochactaeon 75

Trochactaeon (Trochactaeon) 75

autochthonous Trochactaeon 6

azerbaidjanensis, Eotrochactaeon 88

Trochactaeon 88

Actaeonella 88

Omphaloactaeonella 88

Ovactaeonella 88

B

babkovi. Trochactaeon 75

Trochactaeon (Trochactaeon) 75

baconica, Actaeonella 20, 75

baconicus, Paleotrochactaeon 75 

Trochactaeon 75

(Neocylindrites) 75

Baja California 20 

Barrancas Limestone Member, Mayaguez

Foundation, Puerto Rico 24, 36

Barranquitas Quadrangle, Puerto Rico, localities -35, 36

index map 29f

Barremian time interval 9, lOf, llf

barremicus. Trochactaeon 75

Trochactaeon (Neocylindrites) 20, 48, 54, 75 

Barrettia coatsi 34

gigas 71

monilifera 34

rusae 34

sp. C 34

sp. D 34

Barrettia Limestone, Stapleton Formation, Jamaica 44, 71

baylei, Actaeonella (Trochactaeon) 76

Trochactaeon (Neocylindrites) 76

belgica, Trochactaeon 76

belgiciif:, Trochactaeon (Neocylindrites) 76

Trochactaeon (Trochactaeon) 77

Bella Vista District, California, localities 46

beynchi. Actaeonella 76

Purpu ractaeon 76

Trochactaeon (Trochactaeon) 4, 49, 50, 75, 76 

bicincta, Nerinea 4

biconicus, Trochactaeon 76

Trochactaeon (Trochactaeon) 76

Big Hatchet Mountain, New Mexico 56 

biocoenosis 7 

biotic realm, tropical 4 

bivalve framework builders, rudist 4, 24 

borneensis. Actaeonella 6, 35, 67, 68f, 76; pis. 19, 20 

Acteonella 67 

Borneo 35 

Botijas Limestone Member, Pozas Formation,

Puerto Rico 36 

boutillieri, Trochactaeon 76

Trochactaeon (Neocylindrites) 20, 54, 76

brandenbergensis, Trochactaeon (Trochactaeon) 

lamarcki 82,83

Brachydontes, association with Actaeonella sp. 24

Brazil 28f, 34

brevis, Actaeonella (Trochactaeonl 48, 56, 76, 77

Spiractaeon 77

briarti, Actaeonella 76

Purpuractaeon 76

Trochactaeon 48, 76

(Neocylindrites) 54,76

briggsi, Actaeonella 35, 71, 72, 72f, 76; pis. 20, 21 

browm, Actaeonella 24, 67, 67f, 76, pi. 18

Buda Limestone, Texas, localities 45

building corals 4

burckhardti, Actaeonella (Trochactaeon) ~47, 58, 76

Neotrochactaeon 76

Trochactaeon (Mexicotrochactaeon) 34, 58, 59/i 76,

80,81,82, 84, 85, pis. 12,13

(Trochactaeon) 59

burrowing behavior, Actaeon and Actaeonella 5

Cabo Rojo, Puerto Rico, collection localities 29/i 36

Caledonia Formation, Virgin Islands, 40

California, Baja 20

Bella Vista District 46

Cenomanian distribution of Actaeonella 35

collection localities 34f, 46 

stratigraphic ranges of Western Hemisphere

species, index map 28f 

Callianassa 7

Calliostoma 64

Campanian time interval, through Maastrichtian -9, lOf 

Caprina Limestone (Edwards Limestone), Texas 45 

Caprinuloidea rudist bivalve framework sequences,

Puerto Rico 24 

Canahculata, Retusa 6

Cardenas Formation, Sar Luis Potosi, Mexico 34,46 

Caribbean region 2, 6,10 

Caribbean-Mexico region 20 

Cassidaria cretacea 49, 78

Cassis madagascarensis spinella 5

catalogue, actaeonellid species 74 

caucasica, Actaeonella 64, 76, 77

Actaeonella causica 76

(Actaeonella) 77

(Pchelincevella) 76

caucasica, Actaeonella 76

grossouvrei, Actaeonella 76, 77, 81, 86

shiosensis, Actaeonella 77, 85

styriaca, Actaeonella 77, 81

zouparriensis, Actaeonella 77, 87

caucasicus, Trochactaeon 77

Trochactaeon (Trochactaeon) 77

Cayey Quadrangle, Puerto Rico 29f, 35 

Cenomanian distribution, Actaeonella 9,10, 35 

Central Aguirre Quadrangle, Puerto Rico 29f, 37 

Central America, new actaeonellid finds 24 

Central mlier, Jamaica, localities 30/i 40-42 

Chiapas (State of), Mexico 20, 34, 35, 46 

choffati, Neotrochactaeon 80

Trochactaeon 77

(Trochactaeon) 77

Coamo, Puerto Rico, localities 29f 

Coamo P ormation, Puerto Rico 37 

abundance of Actaeonella coquiensis 35, 70 

Coamo Quadrangle, Puerto Rico, localities 37 

coatsi, Barrettia 34 

collection localities, index maps 29/-S4/' 

collignoni, Actaeonella (Trochactaeon) 77

Trochactaeon 77, 80

TVoc h act aeon (Trochactaeo n I 7 7 

columellar plaits 4 

columellar retractor muscles 2 

communis, Trochactaeon 77

Trochactaeon (Trochactaeon) 77

Turritella 6

conform is, Actaeonella 77

coma, Trochactaeon 77 

Coniacian region of Mexico 20 

conica, Actaeonella 76, 77

Actaeonella (Trochactaeon) 76, 77

Tornata 77

Tornatella 77

conicus, Purpuractaeon 49, 77

Spiractaeon 77

Trochactaeon 49, 79

(Trochactaeon) 77

coniformis coniformis, Trochactaeon 

(Mexicotrochactaeon) 77
INDEX 99

Page

Actaeonella 56, 57, 58, 62, 63, 85; pi. 16 

(Trochactaeon) 48, 56, 77, 84

Trochactaeon 48

(Mexicotrochactaeon) 54, 56, 57, 57f, 58-60,

62, 75, 76, 81, 83,84,86; pis. 10,11

comformis 77

conoidea, Actaeonella (Trnchactaeon) 78

Mitra 78

conoideus, Trochactaeon (TrochactaeonI 78

Conus schwsensis 85 

coquiensis, Actaeonella 35, 70, 70/i 78; pis. 20, 21 

corals, building 4 

cordeirm, Actaeonella (TrochactaeonI 78

Paleotrochactaeon 73

Trochactaeon 43

(Neocylmdntes) 7g. 79

Corona, California, collection localities 46

correlations, formations 24, 25f, 26f

Upper Cretaceous rocks 26f, 27f

cossmanni, Trochactaeon 80

Trochactaeon (Trochactaeon} 78

Cotui Limestone. Puerto Rico 26f, 34, 35, 38-39 

crassa, Actaeonell 7g

Actaeonella 73, 77, 78, 81, 82. 89

(Pchehncevella) 73

Volvaria 78

Volvuhna 78

cretacea, Actaeonella 78

Cassidaria 49. 78

Cretaceous deposits, Caribbean Antilles and

Central America 6 

Cretaceous faunal assemblages, salinity ranges 8f 

Cretaceous formations, correlation, Puerto Rico

and Texas 25^

Cretaceous outcrop, Cuba 3l/"

Cretaceous rocks, correlation, Jamaica 27f, 3Qf

correlation, Puerto Rico 26^

Puerto Rico and Texas 25^

Cretaceous stratigraphic record 24

cretaceus, Purpuractaeon 73

Troch actaeon 73

cnsminensis, Paleotrochactaeon 78,85

Trochactaeon 48, 78

iNeocylmdrites) 54, 73

Cuba, collection localities 3\f, 35, 44-45 

Stratigraphic ranges of Western Hemisphere

species 28^ 

cubensis, Actaeonella-^, 35, 65, 72, 73 f, 78; pis. 17, 22, 23 

Cucullaea, bivalve 7 

cumminsi, Trochactaeon 53,56,78

(Neocylmdntes) 24, 53, 53f. 55, 56, 78, 83; pis. 4, 9 

cycloideus, Trochactaeon 78

Trochactaeon I Trochactaeon I 78

Cyhchna 2, 4

Cyhchna oryza 6

cylmdracea, Actaeonella 78

Actaeonella (Trochactaeon) 52, 78

cylindraceus, Paleotrochactaeon 78

Trochactaeon 48, 78

(Trochactaeon) 52, 78

cyhndncus, Mesotrochactaeon 78

Troch actaeon 78 

(Trochactaeon) 52, 78

Cyhndritella 87

Cylindrites, and Cretaceous Actaeonellidae 20 

Cylmdntes damesi 79

Cylindrites delgadoi 78, 79

D

damesi, Cylindrites 79 

Trochactaeon 79

darwasensis, Spiractaeon 79

Trochactaeon 79, 83

(Trochactaeon) 79

dating of formations, Texas vs. Anlillean islands 24

decorata, Actaeonella 65, 78

Acteonella 78

delgadoi, Actaeonella 66, 66f, 77, 79, 81; pi. 18

Page

Acteonella (Pchelinceve/la) 66 

Cylindrites 78, 79

Trochactaeon 79

devii, Actaeonella 88

Omphaloacteonella 88

Ovactaeonella 88

Devil's River Limestone, Texas 24,45 

differential sorting after death 7 

dilatatus, Trochactaeon 79, 80

Discotectus 4

dispersal potential 9 

Distefenella 7

distribution 9

maps 1 If- 19^

pattern in tethyan realm 4

diversity 9, \Qf

dohohformis, Trochactaeon 79

dohum, Actaeonella 88

Ovactaeonella 81, 88, 89

Peruviella 88

d'Orbigny, Actaeonella 63

Dreistaetten, Austria 7 

dubertreti, Actaeonella 79

Eotrochactaeon 88

Trochactaeon (Trochactaeon) 79

duckettsensis, Actaeonella 35, 69, 69f, 72, 79; pis. 19, 20 

Durania nichdasi 34

ojauchalensis 58

E

Early Cretaceous stratigraphic record 24 

early ontogeny 9 

ecology 4 

Edwards Limestone (Caprma Limestone), Texas 45 

ellipsoides, Actaeonella 79, 82

Mesotrochactaeon 82, 83, 85

Trochactaeon (Trochactaeon) 79

Ellipsoscaph a 79

elliptica, Actaeonella 79

ellipticus, Spiractaeon 79

Trochactaeon 79

(Troch actaeon) 79

elongata, Actaeonella 70, 79

El Rayo Formation, Puerto Rico 35, 38-40 

Eotrochactaeon 20, 85, 88

azerbaidjanensis 88

dubertreti 88

gormdjukensis 89

zumoffeni 90

epifaunal life habit, Trochactaeon 5 

Erath County, Texas 4,6; pi. 1 

Euchelus 64

Euhalinikum salinity zone 1

evolution 20 

excelsus, Neotrochactaeon 79

Trochactaeon 79, 85

(Trochactaeon I 79

excluded genera and species 87

exhalant area 4

extinction 1

parallels with other groups 22

faba, Actaeonella 79

Actaeonella (Pchehncevella) 79

Voluaria 79

feeding habits 6 

ferganica, Ovactaeonella 64

ferjanica, Ovactaeonella 88

Flor de Alba Limestone Lentil, Pozas Formation,

Puerto Rico 37 

Florida Quadrangle, Puerto Rico 29f, 34, 37 

flonformis, Tampsia 58

formations, correlations 24, 25f, 26f 

framework builders, rudist bivalve 4, 24 

France, southern 20

frazierensis, Actaeonella 50, 79

Trochactaeon (Trochactaeon) 20, 22, 50, 51, 79, 85;

pi. 8

Fredencksburg Group, Texas, localities 45 

fusiformis, Actaeonella 20, 22, 79

G

gagii, Actaeonella 88

Actaeonella (Sogdianella) 88

Omphaloacteonella 88

Sogdianella 88

galloprovinciahs, Trochactaeon (Trochactaeon) 59, 79, 80 

Garumnien, of Spain 20 

genera excluded from Actaeonellidae 87 

geographic distribution, Actaeonella and

Trochactaeon Uf-19f,20 

georgica, Actaeonella 88

Acteonella 88

gerthi, Peruvia 88

Peruviella 88, 89

ghazirensis, Actaeonella 80

Trochactaeon (Trochactaeon) 80

gigantea, Actaeonella 79, 80, 85

Actaeonella (Trochactaeon) 48, 58, 79, 80

(Trochactaeon) gigantea 80 

gigantea, Actaeonella (Trochactaeon) 80

globosa, Actaeonella (Trochactaeon) 80 

Neotrochactaeon 80

subglobosa, Actaeonella (Trochactaeon) 80 

Tornatella 80

Trochactaeon 80

ventricosa, Actaeonella (Trochactaeon) 80

giganteus, Actaeon 80

giganteus, Trochactaeon (Trochactaeon) 80 

Neotrochactaeon 79, 80

subglobosa, Trochactaeon /Trochactaeon) 80 

subglobasus, Trochactaeon (Trochactaeon) -78, 79, 80

82,86,87 

Tornatella 80

Trochactaeon 79, 80

(Trochactaeon) 80, 84

giganteus 80 

gigas, Barrettia 71

glandiformis, Actaeonella 80, 85

Neotrochactaeon 80. 85

Trochactaeon 80

(Trochactaeon) 80

Glaucon m 7

Glen Rose Formation, Texas 45 

Globiconcha maxima 78

globosa, Actaeonella (Trochactaeon)gigantea 80 

goldfussi, Actaeonella 80

Actaeonella (Trochactaeon) 80, 81

Trochactaeon 80

(Trochactaeon) 77, 80, 81, 82, 86

gormdjukensis, Eotrochactaeon 89

Trochactaeon 89

Gosau, Austria 9 

gosaviensis, Trochactaeon 55

Trochactaeon (Neocylmdntes) 81

gracihs, Actaeonell 78

Actaeonella 66, 79, 81

(Pchelincevella) 78

Acteonella 66

Trochactaeon 81, 83

gradatus, Trochactaeon 55

Trochactaeon (Neocylindrites) 81

grandis, Actaeonella 89

Actaeonella (Sogdianella) 89

Acteonella (Sogdianella) 89

Sogdianella 89

granthamensis, Trochactaeon (Mexicotrochactaeon) 60, 

61, 6lf, 63,81; pis. 13,15

Green Island inlier, Jamaica 30f, 44 

grossouvrei, Actaeonella - 64, 76, 77, 79, 81

Actaeonella (Pchehncevella) 76

caucasica 76, 77, 81, 86

Acteonella 64,66
100 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Page

Guatemala, Actaeonella browni associations 24 

Cenomaman distribution of Actaeonella 35 

collection localities 33f, 46 

stratigraphic ranges of Western Hemisphere

species 28f 

Guerrero (State of), Mexico 34 

Guinea Corn Formation, Jamaica 9, 35, 40-42 

gulistaenensis, Trnchactaeon 81

gulistanensis, Trochactaeon 81

Habana Formation, Cuba, localities 44-45 

habits, feeding 6 

head-foot mass, musculature 4 

Hispanola, new Actaeonellid finds 24 

Humboldti, Actaeonella (Trochactaeon) 58, 76, 81 

humei, Actaeonella (Trochactaeon) 81 

Trochactaeon (Trochactaeon) 81 

hydrocarbon resources, rudist bivalve framework

structures 2 

hypostracum crests I

id/evanensis, Paleotrochactaeon 81

Trochactat'on {Trochactaeon> 81

irnpressus, Trochactaeon 76,81

inconstans, Actaeonella 58 

Actaeonella lTrt>chactaeon> 48, 56, 77, 81 

Mexicotrochactaeon 56 

Indo-Pacific faunas, affinities with Western

Hemisphere gastropods 35 

inductura, parietal 2

inflata. Actaeonella 73, 81

Actaeonella / Volvulinal 81

infaunal life habit 5 

Inoceramus Shales, Jamaica 35 

inmgnis. Trochactaeon 81 

intermedius, Trnchactaeon (Trochactaeonl- 75

mterplait muscle bands, function 4 

introduction 1 

inuoluta, Actaeonella 81

irregulans. Actaeonella /Trochactat'onI 48, 56, 77, 81 

Mexicotrochactaeon 56, 57

isomorphism, and environmental setings 5 

Itieriidae 64,85

Jamaica, collection localities 3Qf. 40-44 

correlation of Upper Cretaceous rocks 27 f, 30f 

stratigraphic ranges, Western Hemisphere 

species 23f

Jerusalem Mountain inlier 30f 

localities 43, 44

khalilovi, Actaeonella -

Actaeonella (Sogdianellal -

Sogdianella 89 

Klamath River, California 46 

knnstantiti, Actaeonc/la 89

Ovactaeonella 89

Koyokuk River, Alaska 46 

kuehni, Trochactaeon iTrochactaeoni 80,81 

kurdistanica. Actaeonella 89

Actaeonella (Sogdianellal 89 

Sogdianella 84,89

Page

laevis, Actaeonella 9, 64, 70, 72, 79, 81, 90

Actaeonella fVolvuhnal 77, 81

laevis 81, 82, 86

leavis, Actaeonella 81, 82, 86

Volvaria 63-64, 81

Volvulma 81

zekelii, Sogdianella 90

lagoonal associations 7

lamarcki, Actaeonella 80

Actaeonella iTrochactaeoni 82

Acteonella 82

brandenbergeasis, Trochactaeon/Trochactaeon) 82,

83

larna'-cki Trot hactati>u '1 rochartaeot, 80, 82

85,86,87 

Spiractaeon 82

Tornatella 82

Trochactaeon 80

ITrochactaeoni 80, 82. 85, 86, 87

Trochactaeon iTrochactaeoni 9, 51

Trochactaeon /Trochactaeon) lamarcki 80, 82, 85,

86,87 

lamarcku. Tornatella 80, 82

Late Cretaceous stratigraphic record 24

Late Cretaceous species having tubercles or nodes 49

laticostata. Sevanella 48. 82

Trochactaeon iSevanellat 82

latus, Trochactaeon /Trochactaeonl 82

life mode and habits 4, 5 

Littorma 5

locality maps 29f-34f 

locality register 35 

longa, Actaeonella 82

longisulcus, Mesotrochactaeon 82, 85

Trochactaeon 82

/Trochactaeonl 85

longitudinal grooves, Trochactaeon body whorl 2 

luciana. Actaeonel/a 59

lucmno. Actaeonella 58. 58f. 76, 82

M

Maastrichtian species, Caribbean and

middle America 9, 15/! 34 

madagascarensis spinella, Cassis 5

Magueyes Formation, Puerto Rico 24, 36 

Maldon inlier, Jamaica, localities 44 

Maldon Limestone, Jamaica, abundance of 

Actaeonella coquiensis 70

localities 3Qf, 44

occurrences of Actaeonella marchmontensis 35

mammilatus, Purpuractaeon 82

irochactaeon iTrochactaeoni 82

mantle edge, character 2 

Maravillas Formation, Puerto Rico, localities -35, 37, 38 

Marchmont inlier, Jamaica 9, 3Qf. 35, 42-43 

marchmontensis, Actaeonella 35, 71, 71f, 72, 82; pi. 21 

mardakertensis. Actaeonella 89

Ovactaeonella 89

Martha's Vineyard. Massachusetts 20, 24, 35; pi. 22 

Massachusetts, localities 45 

matensis, Actaeonella 82

Actaeonella iTrochactaeoni 82 

Trochactaeon 79, 82

(Trochactaeon) 22, 51, 52, 79, 82 

maxima, Globiconcha 78

Mayaguez Formation, Puerto Rico 24 

Meiomesohalinikum salinity zone 7 

melonensis, Trochactaeon (Mexicotrochactaeon) 63, 82;

pis. 16, 17

Melones Limestone, Puerto Rico 6, 36, 38 

Mesotrochactaeon 22, 47

arnaudi 75

artsachensis 75

cyhndricus 78 

elhpsoides 82, 83, 85

Page

longisulcus 82, 85

ouremensis 83

ovalis 83

subglobosa 80

tovusensis 85, 86

vasm ikuchensis 86

Mexico, collection localities 33f, 34, 35, 45-46 

nermeid gastropods 24 

new actaeonellid finds 24 

occurrences of Actaeonella borneensis 35 

stratigraphic ranges, Western Hemisphere

species 28f 

Mexicotrochactaeon .48

acutissima 56, 77

inconstans 56

irregularis 56, 77

plamlateralis 56, 77

potosiana 77

potosianus 56 

vanabihs 56,77

(Mexicotrochactaeon) burckhardti, Trochactaeon 34, 58,

59f, 76, 80, 81,82, 84, 85; pis. 12,13

coniformis, Trochactaeon 54, 56, 57, 57f, 58, 62, 75,

76,81,83,84, 86; pis. 10,11

con iformis, Trochactaeon 77

granthamensis, Trochactaeon 60, 61, 61f, 63, 81;

pis. 13, 15

melonensis, Trochactaeon 63, 82; pis. 16,17 

nelsnni, Trochactaeon 82

palmeri, Trochactaeon - 60. 6Qf, 83; pis. 11,14 

rossi, Trochactaeon 57, 60, 62, 62f, 85; pis. 13,16 

teaensis, Trochactaeon 60, 86; pis. 13,14

Trochactaeon 9, 47, 48, 68; pi. 3

spp. 63

wrighti, Trochactaeon 59, 59f, 60, 87; pis. 11, 12

nunutis, Trochactaeon (Neocylindritesi 82

minutus, Trochactaeon 82

mirabihs, Trochactaeon 82

Trochactaeon (Trochactaeon) 82

Mitra conoidea 78

Mojado Formation, New Mexico, age 56 

monihfera, Barrettia 34

Morelos (State of), Mexico, localities 46 

morphology, features 3f 

interspecific variability 48 

muscle bands, interplait 4 

muscles, columellar retractor 2 

musculature, head-foot mass 4

Naticidae, effect on food chains 22

nelsoni, Trochactaeon (Mexicotrochactaeon) -82

Trochactaeon (Neocyhndntes) -20, 34, 55, 55f,

pis. 8, 11 

Neocyhndntes 20, 48

characteristic features 54

Neocyhndntes sp. 80, 87

(Neocyhndntes) abbreviatus, Trochactaeon 75

absalonis, Trochactaeon 75

angustatus, Trochactaeon 75

baconicus. Trochactaeon 75

barremicus, Trochactaeon 20,48,54,75 

baylei, Trochactaeon 76

belgicus, Trochactaeon 76

boutillien, Trochactaeon 20, 54, 76 

briarti, Trochactaeon 54, 76 

cordeiroi, Trochactaeon 78, 79

crismmensis, Trochactaeon 54, 78 

cumminsi, Trochactaeon 24, 53, 53f, 55, 56, 78, 83;

pis. 4, 9 

gosaviensis, Trochactaeon 81

gradatus, Trochactaeon 81

nunutis, Trochactaeon 82

nelsoni, Trochactaeon 20, 34, 55, 55/i pis. 8,11 

olwiformis, Trochactaeon 83

parvus, Trochactaeon 54
INDEX 101

Page

punctatus, Trochactaeon 20, 54, 55, 58, 59, 77,

83,84; pis. 4, 9

Trochactaeon 20, 22, 34, 47

sp. 77 

sp. A 55. 87; pis. 8, 11 

sp. B 56, 87; pi. 8 

ventricosa, Trochactaeon 84

neogastropods 22 

Neotrochactaeon 47, 80

burckhardti 76

choffati 80

excelsus 79

gigantea 80

giganteus 79, 80

glandiformis 80,85

ventricosa 80, 87

ventncosus 80

Nennea 53

abbreviata 75

bicincta 4

Nerineacea, Iteiidae 64 

nerineid-dominated shale and limestone.

Puerto Rico 34 

Neritopsis, association in shales with Trochactaeon 7 

New Mexico, collection localities 32f, 45, 56 

stratigraphic ranges, Western Hemisphere

species 28f 

Nohatatietin, Yukon, Alaska 46 

mcholasi, Durania 34

Nuevo beds, State of Chiapas, Mexico 6

O

obesus, Trochactaeon 82

Trochactaeon (Trochactaeon) 82

obliquecostata, Actaeonella 81. 82 

obliquephcatus, Trochactaeon 76, 81

obtusa, Actaeonella 83

Spiractaeon 83

Trochactaeon (Trochactaeon) 83 

obtusus, Spiractaeon 79,81

Trochactaeon (Trochactaeonl 63

occidentalis, Actaeonella 58

Actaeonella (Trochactaeon) 48, 56, 77, 83 

Ocuilapa, State of Chiapas, Mexico, localities 45 

occurrences in Western Hemisphere 22 

ojauchalensis, Durania 34

olwaeformis, Actaeonella 83

ollviformis, Actaeonella 20, 83

Trochactaeon 83

(Neocylindrites) 83

Omphactaeonella 20

Omphacteonella ovata 89

Omphaloactaeonella 64, 88

azerbaidjanica 88

Omphaloacteonella agjakendensis 88

anchietai 88

devn 88

gagu 88

ovata 88

supernata 89

tenuiflexa 89

ontogeny, early 9 

Orange County, California 46 

orientate, Actaeonella 83

Oriente Province, Cuba, exclusive occurrences 35 

ornata, Actaeonella 65, 83

Actaeonella iPchehnsevella) 83

ornatus, Paleotrochactaeon 83

Trochactaeon 83

(Trochactaeon/ 83

Orocovis, Puerto Rico 29f, 34. 37-38 

oryza, Cyhchna 6

ouremensis, Mesotrochactaeon 83

Trochactaeon (Trochactaeon)- --83

Ovactaeonella 64, 88

agdjakendensis 88

anch leh 88

armenica 88

azerbaidjanica 88

devn 88

dohum 81, 88,89

ferganica 64

ferjanica 88

konstantiti 89

mardakertensis 89

ovata 89

ovahs, Mesotrochactaeon 83

Trochactaeon 83

(Trochactaeon) 83

ovata, Actaeonella 64, 89

Omphacteonella 89

Omphaloacteonella 88

Ovactaeonella 89

oviformis, Actaeonella 35, 51, 52, 6S, 83; pi. 18 

Acteonella 51, 52, 68

packardi, Actaeonella 51, 83

Trochactaeon (Trochactaeon) 20, 22, 51, 52, 52£

68, 83; pi. 8 

Paleotrochactaeon - - - 48

angustatus - 75

bacon icus 75

cordeiroi 78

crisminensis 78, 85

cylindraceus - 78

idjevanensis 81

ornatus 83

Palmarejo, Puerto Rico 29/i 34 

palmeri, Trochactaeon (Mexicotrochactaeon) 60, GOf, 83;

pis. 11,14 

pamincum, Trochactaeon 83

parietal inductura 2 

parva, Actaeonella 83

Actaeonella (Trochactaeonl 83

Trochactaeon (Trochactaeon) 83

paruus, Actaeonella 53

Actaeonella (Trochactaeon) 78 

Trochactaeon 53, 54, 83

(Neocylindrites) 54

patagonica, Actaeonella 83

Trochactaeon 83

Pate/fa 7 

pcelinsevi, Purpuractaeon 84

pcehnsevi, Trochactaeon (Trochactaeon) 49, 50, 84 

Pchehncevella 64,65

(Pchehncevella) caucasica, Actaeonella 76

crassa, Actaeonella 78

delgadoi, Acteonella 66

/a&a, Actaeonella 79

granhs, Actaeonella 78

grossouvrei, Actaeonella 76

pchelincevi, Actaeonella 84

schiosensis, Actaeonella 77, 85

stynaca, Actaeonella 77

terterensis, Actaeonella 86

Acteonella 86

zouparnensis, Actaeonella 77

(Pchehnsevella) ornata, Actaeonella 83

praecaucasica, Actaeonella 84

(Pchelintsevella)sublaevis, Actaeonella 85

pchelincevi, Actaeonella 84

Actaeonella (Pchehncevella) 84

peconsensis, Actaeonella 35, 65

pecosercsis, Actaeonella 65, 65f, 66, 84; pis. 18, 20 

Pedalion, bivalve 7 

Peruvia gerthi 88

peruviana, Actaeonella 89

Sogdianella 89

Peru I'je/ta 88

rfo/jum 88

gerfft; 88,89

Philine 4

phylogeny 20, 2 If 

proposed 23 f

Pileolus 4

Pirenella, association in shales with Trochactaeon -7 

Plagiobrissus 5

plaits, columellar 4 

plamlaterahs, Actaeonella 58

Actaeonella (Trochactaeon) 48, 54, 56, 77, 78, 84 

Mexicotrochactaeon 56, 77

plate-tectonic theory 2 

Pleiomesohalinikum salinity zone 7 

Polyhalinikum salinity zone 7 

Polymesoda 7

polymorphism, and environmental setings 5 

pompei, Actaeonella 58, 58f, 59, 76, 84

Pondoland, South Africa 20 

posterior aperture margin 4 

post-Turonian rocks 22 

potosiana, Actaeonella (Trochactaeon) 48, 56, 77, 84 

Mexicotrochactaeon 77

potosianus, Mexicotrochactaeon 56

Pozas Formation, Puerto Rico 34, 36, 37-38 

praecaucasica, Actaeonella 84

Actaennella (Pchehnsevella) 84

praesulcata, Actaeonella 84

praesupernata, Actaeonella 89

Actaeonella (Sogdianella) 89

Sogdianella 89

prisca, Actaeonella (Trochactaeon) 84

projectiliformis, Trochactaeon (Trochactaeon) - 84

protoconch character, size 9

pruniformis, Actaeonella (Trochactaeon) 58, 84

Trochactaeon 81, 84, 85, 86

pseudocyhndraceus, Trochactaeon 84

Trochactaeon (Trochactaeon) 22, 52, 84 

Pseudomelania 7

Ptychocyhndntes 20

Puerto Real Quadrangle, Puerto Rico, localities 29f 38

collection localities 29£ 35-40

Puerto Rico, correlations of Cretaceous rocks 25 f, 26f

occurrences, actaeonellids 24

Trochactaeon I Trochactaeon) woodsi -34

stratigraphic ranges, Western Hemisphere

species 25 f 

punctata, Actaeonella (Trochactaeon) acutissima - 54,

56,58.77,84

subspecies 54 

punctatus, Trochactaeon (Neocylindrites)-----20,54,55,

58, 59,77,83, 84; pis. 4,9 

Purpuractaeon 47, 49

beynch i 76

briarti 76

comcus 49, 77

cretaceus 78

mammilatus 82

pcelinsevi 84

vediensis 86

quadratus, Trochactaeon 84

quadriphcata, Actaeonella I Trochactaeon) 58. 76, 84 

quadriphcatus, Actaeonella (Trochactaeon) 47

R

Radiohtes-Trochactaeon group 6 

radiolitid curvature 6 

Rancho Nuevo Beds, State of Chiapas, Mexico 35, 46 

ranges, stratigraphic, Western Hemisphere 28f 

Raritan Formation, Massachusetts 45 

references cited 90 

reflections of soft parts on shell 2 

reguearis, Trochactaeon 79, 84

Trochactaeon I Trochactaeon) 84

Remane's salinity zones 7 

renauxiana, Actaeonella 47, 80, 84, 85

Actaeonella (Trochactaeon) 84, 85

Trochactaeon 84

renauxianus, Trochactaeon 84

Trochactaeon (TrochactaeonI 84
102 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

Rcnauxianus, Trochactaeon 85

repehni. Actaeonella 89

Vernedia 89

resorption phenomena 5 

retractor muscles, columellar 2 

Retusa canaliculata 6 

Reves Member, Pozas Formation, Puerto Rico -34, 35-36 

Rimella 64 

Rio Bauta Member, Pozas Formation, Puerto

Rico 34, 37-38 

Rio Descalabrado Quadrangle, Puerto Rico 29f, 38 

Rio Maton Limestone Member, Robles Formation,

Puerto Rico 37 

Rio Minho, Central inlier. Jamaica 7 

roashensis, Actaeonella 84

Actaeonella I Actaeonella) 84

robmsom, Actaeonella 35, 65, 69, 72, 73, 84; pis. 17,

22,23

Robles Formation, Puerto Rico 37 

robustus, Trochactaeon 84 

Rocky Gulch, Siskiyou County, California 46 

rossi. Trochactaeon iMexicotrochactaeon) 57, 60, 62, 

62/;85;pls. 13,16 

rotundata, Actaeonella 85

rudist bivalve framework builders 4, 24

biostratigraphy 24, 25f, 26f 

rugosus, Trochactaeon 85 

rusae, Barrettia 34

rustica, Actaeonela 50, 79, 85

Sabana Grande Formation, Puerto Rico 39 

Sabana Hoyos Limestone Member, Maravillas

Formation, Puerto Rico 38 

St. Ann's Great River inlier, Jamaica 9f, 38, 39,44 

salinity ranges, Cretaceous faunal assemblages 8f 

salinity tolerance 7 

salinity zones, Remane's 7 

salomoms, Actaeonella 85 

Actaeonella iTrochactaeon) __82, 85

Acteonella 82

Troch actaeon 85

iTrochactaeon) 85

Sanctae-Crucis, Actaeonella 85

sanctaecrucis, Trochactaeon (Trochactaeon) 59,80,

84,85

San German Quadrangle, Puerto Rico 38-40 

San Luis Potosi (State of), Mexico 34, 35, 46, 58 

scalata, Valuta 85

scalatus, Trochactaeon iTrochactaeon) 85 

Scaphander .^

schwsensis, Actaeonella 77, 85

Actaeonella iPchehncevellal 77, 85 

(Volvulina) 77, 85 

caucasica 77,85

Conus 85 

Trochactaeon 85

schirabadensis, Spnactaeon 85

Trochactaeon 85

(Trochactaeon) 75. 85 

secondary lamellae 5 

sergipensis, Trochactaeon 88, 89

Sera nella 48

laticostata 48, 82

tubercu lata 86

(Sevanella) laticostata, Trochactaeon 82

Trochactaeon 20, 22, 47,48

tuberculatus, Trochactaeon 86

Sevenella 78

Shasta County, California 46 

shell character, and life habit 3f, 5 

shell morphology 65 

shell, reflection of soft parts 2 

Sierra Rica, New Mexico 56 

silvae, Actaeonella - 85 

similis, Trochactaeon 85

sinaiensis, Actaeonella 85

Slippery Rock Formation, Jamaica 40

soft parts, reflections on shell 2 

Sogdian ella 64, 88

aQgii ____ _ ____ ________88

gran dis 89

khahlovi 89

kurdistanica 84, 89

laevis zekehi 90

peruviana 89

praesupernata 89

supern ata 89

tagarensis 89

tenuiflexa 89

(Sogdianella)gagu, Actaeonella 88

grandis, Actaeonella 89 

Acteonella 89

khalilovi, Actaeonella 89

kurdistanica, Actaeonella 89

praesupernata, Actaeonella 89

supernata, Actaeonella 89

Acteonella 89 

tagarensis, Actaeonella 89

tenuiflexa, Actaeonella 89 

zekehi. Actaeonella 90

solidus, Troch actaeon 85

Trochactaeon (Trochactaeon) 85

sorting, differential, after death 7 

South Fork Cow Creek.Shasta County, California 46 

Spain, Garumnien 20 

species assigned to Actaeonellidae 74-86 

species diversity during Cretaceous lOf 

species excluded from Actaeonellidae 87-90 

speciosus, Trochactaeon 85

Trochactaeon (Trochactaeon) 85 

spinella, Cassis madagascarensis 5

Sphenodiscus sp. 34 

Spir actaeon 47

brevis 77

con icus 77

darwasensis 79

elhpticus 79

lamarcki 82

obtusa 83

obtusus 79, 81

sch irabadensis 85 

variospiralis 86

SSR, Armenian 22 

stabilis, Trochactaeon 85

Stapleton Formation, Jamaica 35, 44, 70 

Stinking Canyon, Shasta County, California 46 

Stoliczka, Volvulina 63

stratigraphy. Western Hemisphere species, ranges 28f

summary 22-35

styriaca, Actaeonella (Pchelmcevella) 77

Actaeonella caucasica 77,81

subangustatus, Actaeonella (Trochactaeon) 78 

Trochactaeon 78, 85

subburckhardti, Trochactaeon 85

subglandiformis, Trochactaeon 80, 85

subghbosa, Actaeonella 80

Actaeonella (Trochactaeon)gigantea 80

Mesotrochactaeon 80

Tornatella 80

Trochactaeon (TrochactaeonI giganteus 80 

subglobosus, Trochactaeon - -80

Trochactaeon (Trochactaeon)giganteus 78, 79, 80,

82,86,87 

sublaevis, Actaeonella 66, 67, 85

Actaeonella iPchelintsevella) - 85 

(Volvana) 66, 85

subobtusus, Trochactaeon 79

subotusus, Trochactaeon 85

subovatus, Trochactaeon 80,85 

su bpruniformis, Troch actaeon 86

Trochactaeon ITrochactaeon) 86

subrenauxi, Trochactaeon 86

Trochactaeon ITrochactaeon) 86

substrate preference 6 

Sunderland inlier, Jamaica 30/i 44, 70 

supernata, Actaeonella 64, 89

Page

Actaeonella (Sogdianella) 89

Acteonella 89

(Sogdianella) 89

Omphaloacteonella 89

Sogdianella 89

syriaca. Actaeonella 81, 86

Acteonella 82

systematic paleontology 46

szontaghi, Actaeonella 86

Trochactaeon (Trochactaeon) 86

tagarensis, Actaeonella 89

Actaeonella (Sogdianella) 89

Sogdianella 89

tamanderensis, Actaeonella 86

Tampsia floriformis zone, Mexico 58 

teaensis, Actaeonella 68

Trochactaeon (Mexicotrochactaeon)-60, 86; pis. 13,14 

Tellina, bivalve 7 

teloconch development 9 

tenuiflexa, Actaeonella 89

Actaeonella (Sogdianella) 89

Omphaloacteonella 89

Sogdianella 89

terebellum, Actaeonella 81, 86

Actaeonella 64

Acteonella 64

terterensis, Actaeonella 86 

Actaeonella (Pchelmcevella) 86

Acteonella (Pchelmcevella) 86

Tethyan biogeographic province 1

faunal associations 4

texana, Volvulina 88

Texas, Cenomanean distribution of Actaeonella 35 

collection localities 32f, 45 

correlation of Middle Cretaceous formations 25 f 

dating of formations 24 

stratigraphic ranges, Western Hemispheric

species 28 f 

Thorp Springs Limestone Member, Glen Rose

Formation, Texas 45 

time intervals, occurrences of species 9 

division of date 9, Wf 

Titaniisarcohtes limestone, Jamaica 9, 35, 42-44 

tolerance, salinity 7 

Tornata conica 77

Tornatella conica 77

gigantea 80

giganteus 80 

lamarcki 82

lamarckii 80, 82

subghbosa 80

tornatilis, Actaeon 2

Torreyites 11

torrubiai, Actaeonella 86

toucasiana, Actaeonella 80, 86

tovusensis, Mesotrochactaeon 85, 86

Trochactaeon (Trochactaeon) 79, 86 

Tra/anella 4,24

Transcaucasus 10, 20 

transylvanicus, Trochactaeon 80,86

Trichonopoly area, southern India 20 

Trochactaeon 7, 9, 46, 68, 83; pis. 3-16

abbreviatus 75

absalonis 48

acutissima 48 

agbulakhensis 75

alibeglyensis 75

angustatus 48, 75

armenicus 75

arnaudi 75

asiaticus 85

associations 6 

atschadjurensis 75

azerbaidjanensis 88

babkoi'i 75

bacon icus 75
INDEX 103

Page

barrem icus 75 

belgica 76 

biconicus 76

boutilheri 76 

bnarti 48, 76 

caucasicus 77 

choffa ti 77

colhgnoni 77, 80

communis 77 

con m 77

corncus 49, 79

con i form is 48 

cordieroi 43

cossmanni 80 

cretaceus 78 

cnsminensis 48, 78 

cumminsi 53, 56, 78 

cycloideus 78 

cyhndraceus 48, 78 

cyhndncus 78 

damesi 79

darwasensis 79, 83

delgadm 79 

dilatatus 79, 80 

doliohformis 79 

elhpticus 79 

excelsus 79, 85

gigantea 80

giganteus 79, 80

glandiformis 80

goldfussi 80 

gonndjukensis 89 

gosav lensis 55 

gracilis 81, 83 

gradatus 55 

guhstaenensis 81 

impressus 76, 81 

insigms 81

lamarcki 80 

life habit 5 

longisculcus 82 

matensis 79, 82

mirabilis 82

obesus 82

obhquephcatus 76, 81

occurrences, various time intervals \\f-\5f 

oliviformis 83

ornatus 83

ovahs- -83

pamiricum 83 

parvus 53, 54, 78, 83; pi. 9

patagonica 83

patterns of rarity and abundance, Western

Hemisphere 35 

pruniformis 81, 84, 85, 86

pseudocyhndraceus 84 

quadratus 84 

regularis 79, 84 

renauxiana 84 

renauxianus 84 

Renauxianus 85 

robustus 84 

rugosus 85 

salomonis 85

sands 6 

sch losensis 85 

schirabadensis 85

sergipensis 88, 89 

shell 3f, 5 

similis 85 

sohdus 85

specwsus 85

stabilis 85

subangustatus 78, 85 

subburckhardti 85 

subglandiformis 80, 85

subglobosus 80 

subobtusus 79, 86

subotusus 85

subovatus 80, 85 

subprumformis 86 

subrenauxi 86 

transylvamcus 80, 86 

truncatum 86 

truncatus 86 

tumidus 86

typus II 80 

vasmikuchensis 86 

zekeln 87 

Sp. 87

s.s. 22, 47

(Mexicotrochactaeon) 2, 4, 7, 9, 20, 22, 34, 47, 48,

68; pi. 3

burckhardti 34, 58, 5Sf, 59f, 76, 80, 81, 82 

84, 85; pis. 12,13

comformis 54, 56, 57, 51 f. 58-60, 62, 75, 76 

81,83, 84, 86; pis. 10,11

coniformis 77

granthamensis 34-35, 60, 61, 6lf, 63, 81;

pis. 13, 15

melonensis 34, 63, 82; pis. 16,17 

nelsom 82 

palmen 60, 60f, 83; pis. 11, 14 

rossi 35, 57, 60, 62. 62f, 85; pis. 13, 16 

teaensis 35, 60, 86; pis. 13, 14 

wrighti 35, 59, 59f, 60, 87; pis. 11,12 

sp. pi. 9 

spp. 63

(Neocyhndritesl 4, 5, 7, 20, 22, 34, 47,48

abbreviatus 75 

absalonis 75 

angustatus 75

baconicus 75 

barremwus 20, 48, 54, 75 

baylei 76

belgicus 76

boutilheri 20, 54, 76 

bnarti 54, 76

cordeiroi 78, 79 

cnsmmenis 54, 78 

crisminensis-brmrti group, Europe 56 

cumminsi 24, 53, 53f, 55, 56, 78, 83; pis. 4, 9 

gosaviensis 81 

gradatus 81

minutis 82 

nelsom 20, 34, 55, 55f; pis. 8,11 

oliviformis 83 

parvus 54 

punctatus 20, 34, 54, 55, 58, 59, 77, 83, 84;

pis. 4, 9 

ventncosa 84

sp. 77 

sp. A 55, 87; pis. 8,11 

sp. B 56, 87; pi. 9

iSevanella) 20, 22, 47, 48 

laticostata 82

tubercu latus 86

(Trochactaeon) 2, 4, 7, 9, 22, 34, 47, 49, 68 

acuminata 75

acuminatus 75 

ahbeglyensis 75

armenicus 75 

arnaudi 75 

artsachensis 75

asiaticus 75 

atschadjurensis 75 

babkovi 75

belgicus 77

beynchi 4, 49, 50, 75. 76 

biconicus 76 

burckhardti 59 

caucasicus 77 

choffati 77

colhgnoni 77 

communis 77 

conicus 77

conoldeus 78

Page

cossmanni 78

cycloideus 78

cyhndraceus 52, 78

cyhndncus 52, 78

darwasensis 79 

dubertreti 79 

elhpsoides 79 

elhpticus 79 

excelsus 79

frazierensis 20, 22, 24, 50, 51, 79, 85; pi. 8 

galloprovmciahs 59, 79, 80

ghazirensis 80

giganteus 80, 84

giganteus 80

subglobosa 80

subglobosus 78, 79, 80, 82, 86, 87 

glandiformis 80

goldfussi 77, 80, 81, 82,86

guhstanensis 81

humei 81 

idjevanensis 81 

intermedius 75

kuehm 80,81

lamarcki 9, 51 

brandenbergensis 82, 83

lamarcki 80, 82, 85, 86, 87

latus 82

longisulcus 85 

mamm ilatus 82

matensis 22, 51, 52, 79, 82 

mirabilis 82 

obesus 82 

obtusa 83

obtusus 63

ornatus 83 

ouremensis 83

ovahs 83

packardi 20, 22, 24, 51, 52, 52f, 68, 83; pi. 8 

parva 83

pcehnsevi 49, 50, 84

projectiliformis 84 

pseudocyhndraceus 22, 52, 84

regu laris 84

renauxianus 84 

salomonis 85 

sanctaecrucis 59, 80, 84, 85

scalatus 85 

schirabadensis 75, 85

sohdus 85 

speciosus 85 

subpruniforrnis 86

subrenauxi 86 

szontaghi 86

tovusensis 79, 86 

truncatus 86

tumidus 86

vanospiralis 86 

nasm ikuchensis 86 

vediensis 86

woodsi 4, 7, 20, 22, 34, 47, 49, 49/1 87; pis. 4-7 

zekeln 87

(TrochactaeonI abbreviata, Actaeonella 75 

acuminata, Trochactaeon 75 

acuminatus, Trochactaeon 75

acutissima punctata, Actaeonella 54, 56, 58, 77, 84

Actaeonella , 48, 56, 57, 75, 77

ahbeglyensis, Trochactaeon 75

angustata, Actaeonella 75

armenicus, Trochactaeon 75 

arnaudi, Trochactaeon 75 

artsachensis, Trochactaeon 75 

asiaticus, Trochactaeon 75

atschadjurensis, Trochactaeon 75 

babkovi, Trochactaeon 75 

baylei, Actaeonella 76 

belgicus, Trochactaeon 77

beynchi, Trochactaeon 4,49,50,75,76 

biconicus, Trochactaeon 76 

brevis, Actaeonella 48, 56, 76, 77
104 CRETACEOUS ACTAEONELLID GASTROPODS FROM THE WESTERN HEMISPHERE

burckhardti, Actaeonella 47, 58, 76

Troch actaeo n 59

caucasicus, Trochactaeon 77

choffati, Trochactaeon 77

colhgnoni, Actaeonella 77

Trochactat'on 77

communis, Trochactaeon 77

conica, Actaeonella 76. 77

conicus, Trochactaeon 77

coniformis, Actaeonella - 48. 56, 77, 84

conoidea, Actaeonella 78

conoideus, Trochactaeon 78

cordeiroi, Actaeonella 78

cossmanni, Trochactaeon 78

cycloideus, Trochactaeon 78

cyhndracea, Actaeonella 52, 78

cyhndraceus, Trochactaeon 52,78

cyhndncus, Trochactaeon 52,78

darwasensis. Trochactaeon 79

dubertreti. Trochactaeon 79

elhpsoides, Trochactaeon 79

elhpticus, Trochactaeon 79

excelsus, Trochactaeon 79

frazierensis, Trochactaeon 20, 22, 50, 51, 79, 85.

pi. 8

galloprovinciahs, Trochactaeon 59, 79, 80 

ghazirensis. Trochactaeon 80

gigantea. Actaeonella 48, 58, 79, 80

gigantea, Actaeonella 80

globosa, Actaeonella 80

subglobosa, Actaeonella 80

ventricosa, Actaeonella - -80 

giganteusgiganteus, Trochactaeon 80

subglobosa, Trochactaeon 80

subglobosus, Trochactaeon 78. 79, 80, 82, 86, 87 

Trochactaeon 80, 84

glandiformis, Trochactaeon 80

goldfussi, Actaeonella 80, 81

Trochactaeon 77. 80, 81, 82, 86

gulistanensis, Trochactaeon 81

Humboldti, Actaeonella 58, 76, 81

humei, Actaeonella -- 81

Trochactaeon 81

idjevanensis, Trochactaeon 81

inconstans, Actaeonella 48, 56, 77, 81 

intermedius. Trochactaeon 75

irregulans, Actaeonella --48, 56, 77, 81 

kuehni. Trochactaeon 80, 81

lamarcki, Actaeonella 82

brandenbergensis, Trochactaeon 82, 83

lamarcki, Trochactaeon 80,82,85,86,87 

Trochactaeon 9, 51

latus, Trochactaeon 82

longisulcus, Trochactaeon - - 85 

mammilatus, Trochactaeon 82

matensis, Actaeonella 82

Trochactaeon - 22. 51, 52. 79, 82 

mirabilis, Trochactaeon 82

obesus, Trochactaeon 82

obtusa, Trochactaeon 83

ubtusus, Trochactaeon 63

occidentals, Actaeonella 48, 56, 77, 83

ornatus, Trochactaeon 83

ouremensis, Trochactaeon 83

Page

ovalis, Trochactaeon 83

packardi, Trochactaeon 20, 22, 57, 52, 52/1 68,

83; pi. 8 

parva, Actaeonella 83

Trochactaeon 83

parvus. Actaeonella 78

pcelinsevi, Trochactaeon 49, 50, 84

plandateralis, Actaeonella 48, 54, 56, 77, 78, 84 

potosiana, Actaeonella 48, 56, 77, 84

prisca, Actaeonella 84

projectiliformis. Trochactaeon 84

pruniformis, Actaeonella 58, 84

pseudocylindraceus, Trochactaeon 22,52,84 

quadriplicata. Actaeonella 58, 76, 84

quadriphcatiis. Actaeonella 47

regularis, Trochactaeon 84

renauxiana. Actaeonella 84, 85

renauxianus, Trochactaeon 84

sa/omonis, Actaeonella 82. 85

Troch actaeon 85

sanctaecrucis, Trochactaeon 59, 80, 84, 85 

scalatus, Trochactaeon 85

schirabadensis, Trochactaeon 75,85

solidus, Trochactaeon 85

speciosus, Trochactaeon - - 85 

subangustatus, Actaeonella - -- 78 

subpruniformis, Trochactaeon 86

subrenauxi, Trochactaeon 86

szontaghi, Trochactaeon 86

tovusensis, Trochactaeon 79, 86

Trochactaeon 9, 22, 47, 68

truncatus, Trochactaeon 86

tubercles, in Late Cretaceous Trochactaeon

species 49 

tumidus, Trochactaeon 86

vanabilis, Actaeonella 48, 56, 77, 86 

variospiralis, Trochactaeon 86

i-asmikuchensis, Trochactaeon 86

vediensis, Trochactaeon 86

woodsi, Actaeonella 49, 87

Trochactaeon 4, 20, 22. 34, 47, 49, 49/i 87; pis. 4-7

zekelu, Trochactaeon 87

Actaeonella, sp. 49, 58. 87

tropical biotic realm 4

truncata, Actaeonella 86

truncatum, Trochactaeon 86

truncatus, Trochactaeon 86

Trochactaeon (Troehactaeon) 86

tuberculata, Seuanella 86

tuberculatus, Trochactaeon (Sevanella) 86

tumida. Actaeonella 86

tumidus. Trochactaeon 86

Trochactaeon iTrochactaeonl 86

Turonian time interval 9, IQf

Turritella 6. 7

communis 6

uchauxensis. Actaeonella 64, 76, 81, 86

Acteonella 64,86

Upper Cretaceous rocks, correlations 26f, 27 f

variabihs, Actaeonella (Trochactaeon) 48, 56, 77, 86 

Mexicotrochactaeon 56, 77

variospiralis, Spiractaeon 86

Trochactaeon (Trochactaeoni 86

vasmikuchensis, Mesotrochactaeon 86

Trochactaeon 86

(Trochactaeon) 86

Vaughansfield Limestone, Jamaica 44 

vediensis, Purpuractaeon 86

Trochactaeon (Trochactaeonl 86

ventricosa, Actaeonella (Trochactaeon) gigantea 80 

Neotrochactaeon 80, 87

Trochactaeon iNeocyhndntes) 84 

ventncosus, Neotrochactaeon 80

Vernedia gastropods 85

affinity with Indo-Pacific faunas 35

Vernedia repehni 89

Virgin Islands, locality 40 

Voluta scalata 85

valuta. Actaeonella 87

Acteonella 82

Volvaria crassa 78

faba 79 

laevis 63-64.81

(Volvaria) sublaevis. Actaeonella 66. 85

Volvulina crassa 78

laevis 81

Sto/iczka 63

texana 88

(Volvulina) inflata, Actaeonella 81

laevis, Actaeonella 77, 81

schiosensis, Actaeonella 77, 85

Actaeonella -89

Vorderstoder, Austria 7

W

Washita Group, Texas, localities 45

West Africa 20

Western Hemisphere stratigraphic ranges 22, 2Sf

Woodland Shale, Jamaica 35, 44

Woodman Creek, Shasta County, California 46

Woods Hole, Mass. 6

woodsi, Actaeonella I Trochactaeon) 49, 87

Trochactaeon (Trochactaeon) 4, 20, 22, 34, 47,

49, 49/1, 87; pis 4-7

wnghti, Trochactaeon (Mexicotrochactaeon) 59, 59f. 60

87; pis. 11,12

Yukon River, Alaska 20, 46

zekelu, Actaeonella (Sogdianella) 90

Sogdianella laevis 90

Trochactaeon 87

(Trochactaeon) 87

zouparnensis, Actaeonella 87

Actaeonella caucasica 77, 87

iPchehncevella) 77

zumoffeni, Actaeonella 90

Eotrochactaeon 90
PLATES 1-23

Contact photographs of the plates in this report are available, at cost, from U.S. 

Geological Survey Library, Federal Center, Denver, CO 80225.
PLATE 1

Polished section (x 1) of oriented rock surface from roadcut immediately northwest of Morgan Mill, Erath County, Tex. USNM 305995. 

Arrow "A" points to some of the specimens of Actaeonella sp. in presumed life position, as discussed in text section on life mode. Note also 

overlying interval (arrow "fi") containing transported specimens of Actaeonella primarily oriented with axes parallel to bedding but with 

the apertures unoriented. Note also the geopetal filling of the fossils.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 1

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ACTAEONELLA
PLATE 2

FIGURES 1, 2. Trochactaeon (Mexicotrochactaeon) sp. (p. 9).

1. Aperture view (x 50) of a specimen, showing heterostrophic protoconch and early development of shell. Guinea Corn 

Formation, Central inlier, Jamaica, at locality 61. USGS 29972, USNM 305996.

2. Aperture view (x 50) of a specimen, showing both heterostrophic protoconch and the proportionally rapid increase in 

whorl diameter compared with height of early growth stages. Guinea Corn Formation, Central inlier, Jamaica, at 

locality 61. USGS 29972; USNM 305997.

3. Actaeonella sp. (p. 9).

Aperture view (x 30) of a specimen, showing heterostrophic protoconch and initial stage of envelopment. "Titanosarcolites limestone," Marchmont inlier, Jamaica, at locality 97. USGS 30026; USNM 305998.

4. Actaeonella sp. (p. 9).

Aperture view of a juvenile specimen (x 45), showing heterostrophic protoconch and early developmental stage. "Titanosarcolites limestone," Marchmont inlier, Jamaica, at locality 97. USGS 30026; USNM 305999.

5. Actaeonella sp. C. (p. 74).

Apertural view (x 1) of a specimen from along the Yukon River, Alaska, at locality 170. USGS 2179; USNM 306203.

6. Actaeonella cf. A. coquiensis Sohl and Kollmann (p. 71).

Polished section (x 1) of a specimen of Melones Limestone from the Puerto Real Quadrangle, Puerto Rico, at locality 31, that 

shows a concentration of Actaeonella. USGS 27014; USNM 305600.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 2

,«2-"^w&k ***Kii>-t**f-Hr*» * - -^V 'v35^'-vfli /'^ fc^/^-f^^w-v * ^mm^ / ^ --f^ js^e ^^.^ ^s??^
PLATE 3

FIGURES 1-4. Views of the heterostrophic protoconch (x 350) of a specimen of Trochactaeon (Mexicotrochactaeon) (p. 9) from the Guinea Corn 

Formation, Central inlier, Jamaica, at locality 61. USGS 29972; USNM 306202.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 3

TROCHACTAEON (MEXICOTROCHACTAEON)
PLATE 4

FIGURES 1-4. Trochactaeon (Neocylindrites) cumminsi Stanton (p. 53).

1. Camera lucida drawing (x 1.5) of an axial section though a specimen from the Washita Group, Texas, at locality 136.

USGS 11853; USNM 306001. 

2, 3, 4. Camera lucida drawings (x 1) of transverse sections through specimens from the Devils River Limestone, Texas, at

locality 134. USGS 1941; USNM 306002-306004. 

5. Trochactaeon (Neocylindrites) punctatus (Maldonado-Koerdell) (p. 54).

Camera lucida drawing (x 1) of an axial section through a specimen from the State of Chiapas, Mexico, at locality 151. USGS 

11467; USNM 306005. 

6-8, 12. Trochactaeon (Trochactaeon) woodsi (Rennie) (p. 49).

6-8. Camera lucida drawings (x 0.5) of axial sections through specimens from the Cotui Limestone, San German

Quadrangle, Puerto Rico, at locality 39. USGS 30512; USNM 306006, 306016, 306017. 

12. Camera lucida drawing (x 0.75) of axial section through a specimen from the Cotui Limestone, San German

Quadrangle, Puerto Rico, at locality 39. USGS 30512; USNM 306007. 

9-11. Trochactaeon (Trochactaeon) cf. T. (Trochactaeon) woodsi (Rennie) (p. 50).

Camera lucida drawings (X 0.5) of axial sections of specimens from the Pozas Formation, Orocovis Quadrangle, Puerto Rico, 

at localities 25 and 26. USGS 29099 and 30532; USNM 306008-306010.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 4

TROCHACTAEON (NEOCYLINDRITES) AND T. (TROCHACTAEON)
PLATE 5

FIGURES 1-5. Trochactaeon (Trochactaeon) woodsi (Rennie) (p. 49).

1, 3. Back and aperture views (x 1.5) of a specimen from the Sabana Grande Formation, San German Quadrangle, Puerto 

Rico, at locality 43, which shows raised strong sinused growth-line lamellae at the shoulder nodes. USGS 30343; 

USNM 306011.

2. Aperture view (x 1) of a specimen from the Cotui Limestone, San German Quadrangle, Puerto Rico, at locality 39. 

USGS 30512; USNM 306012.

4. Aperture view (x 1) of a specimen from the same locality. USGS 30512; USNM 306013.

5. Polished axial-section view (x 1) of a specimen from the same locality. USGS 30512; USNM 306007.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 5

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TROCHACTAEON (TROCHACTAEON}
PLATE 6

FIGURES 1, 2, 4, 5. Trochactaeon (Trochactaeon) woodsi (Rennie) (p. 49).

1. Aperture view (x 1) of a specimen from the Pozas Formation, Barranquitas Quadrangle, Puerto Rico, at locality 5. 

USGS 29364; USNM 306014.

2. Back view (x 1) of a specimen from the Cotui Limestone, San German Quadrangle, Puerto Rico, at locality 39. 

USGS 30512; USNM 306015.

4. View of polished axial section (x 1) from the same locality. USGS 30512; USNM 306016.

5. Back view (x 1) of a specimen from the same locality. USGS 30512; USNM 306018. 

3. Trochactaeon (Trochactaeon) cf. T. (Trochactaeon) woodsi (Rennie) (p. 50).

Back view (x 1) of a small specimen from the Pozas Formation, Orocovis Quadrangle, Puerto Rico, at locality 26. 

USGS 30532; USNM 306019.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 6

TROCHACTAEON (TROCHACTAEON}
PLATE 7

FIGURES 1-3. Trochactaeon (Trochactaeon) cf. T. (Trochactaeon) woodsi (Rennie) (p. 50).

1, 3. Aperture and back views (x 1) of a specimen from the Pozas Formation, Orocovis Quadrangle, Puerto Rico, at locality 25.

USGS 29099, USNM 306020.

2. Cut and polished axial section (x 1) of a specimen from the same locality. USGS 29099; USNM 306009. 

4, 5. Trochactaeon (Trochactaeon) woodsi (Rennie) (p. 49).

4. Aperture view (x 1) of a specimen from the Cotui Limestone, San German Quadrangle, Puerto Rico, at locality 39. USGS 

30512; USNM 306021.

5. Aperture view (x 1) of a specimen from the Cotui Limestone, San German Quadrangle, Puerto Rico, at locality 29. Arrow 

points to attachment scar of a Requienia sp., a rudist bivalve. USGS 30512; USNM 306022.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 7

 '

--AEi' 0TR''

TROCHACTAEON (TROCHACTAEON)
PLATE 8

FIGURES 1-5. Trochactaeon (Trochactaeon) frazierensis (Anderson) (p. 50).

1. Aperture view (X 1) of the holotype from the Turonian of Shasta County, Calif., at locality 162. CAS 10664.

2. Aperture view (x 1) of the holotype of Actaeonella rustica Anderson from the Turonian of Shasta County, 

Calif., at locality 162. CAS 10666.

3. Aperture view (x 1) of a specimen from the Turonian of Shasta County, Calif., at locality 165. CAS 53169.

4. Aperture view (x 1) of the paratype from the Turonian of Shasta County, Calif., at locality 162. CAS 10665.

5. Aperture view (x 1) of a specimen from the Turonian of Shasta County, Calif., at locality 165. CAS 53169. 

6, 7, 8. Trochactaeon (Neocylindrites) sp. A. (p. 55).

6, 8. Back and aperture views (x 2 and X 1V:>) of a specimen from the El Rayo Formation, Sabana Grande Quadrangle, Puerto Rico, at locality 36. USGS 29075; USNM 306023.

7. Aperture view (x 1.5) of a specimen from the same locality. USGS 29075; USNM 306084. 

9-14, 18, 19. Trochactaeon (Neocylindrites) nelsoni Sohl and Kollmann, n. sp. (p. 55).

9. Aperture view (x 4) of a paratype from the Pozas Formation, Orocovis Quadrangle, Puerto Rico, at locality 

28. USGS 30534; USNM 306024.

10. Aperture view (x 4) of a paratype from the same locality. USGS 30534; USNM 306025.

11. Aperture view (x 4) of a paratype from the same locality. USGS 30534; USNM 306026.

12. Aperture view (x 1%) of a paratype from the Pozas Formation, Florida Quadrangle, Puerto Rico, at locality 

23. USGS 28742; USNM 306027.

13. Aperture view (x 4) of the holotype from the Pozas Formation, Orocovis Quadrangle, Puerto Rico, at locality 

28. USGS 30534; USNM 306028.

14. Aperture view (x 4) of a paratype from the Pozas Formation, Florida Quadrangle, Puerto Rico, at locality

24. USGS 29372; USNM 306029.

18, 19. Back and aperture view (x 1) of a paratype from the same locality. USGS 29372; USNM 306030. 

15-17. Trochactaeon (Trochactaeon) packardi (Anderson) (p. 51).

15, 16. Aperture and back view (x 1) of the holotype from the Turonian of Siskiyou County, Calif., at locality 163.

CAS 10663.

17. Cut and polished axial section (x 1) of a specimen from the Turonian of Orange County, Calif., at locality 

167. CAS 733.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 8

TROCHACTAEON (TROCHACTAEON) AND T. (NEOCYLINDRITES)
PLATE 9

FIGURES 1-6. Trochactaeon (Neocylindrites) punctatus (Maldonado-Koerdell) (p. 54).

1. Back view of a specimen (x 2) from limestones near Ocuilapa, State of Chiapas, Mexico, at locality 148. USGS 

11467; USNM 306031.

2. Aperture view of a specimen (x 1) from the same locality. USGS 11467; USNM 306032.

3. Polished axial section of a specimen (x 1.5) from the same locality. USGS 11467; USNM 306033.

4. Aperture view of a specimen (x 1) from the same locality. USGS 11467; USNM 306034. 

5, 6. Aperture and back views (x 2) of a specimen from the same locality. USGS 11467; USNM 306035. 

7, 8. Trochactaeon (Neocylindrites) sp. B. (p. 56).

7. Latex cast of an incomplete specimens (x 1.5) from the Mojado Formation, Hidalgo County, N. Mex., at locality 147. 

USGS 1694; USNM 306036.

8. Latex cast of a specimen (x 1.5) from the same locality. USGS 1694; USNM 306037. 

9-19. Trochactaeon (Neocylindrites) cumminsi Stanton (p. 53).

9. Aperture view (x 1.5) of a syntype from the Devils River Limestone, Val Verde County, Tex., at locality 134. USGS

1941: USNM 306038. 

10, 11. Back and aperture views of a specimen (x 2) from the same locality. USGS 1941; USNM 306039.

12. Aperture view (x 1) of a syntype (x 1) from the same locality. USGS 1941; USNM 306040.

13, 14. Aperture views (x 4) of the cotypes of Trochactaeon parvus Stanton from Edwards Limestone at the mouth of 

Bartons Creek near Austin, Tex. USNM 103123a, b.

15. Back view of a syntype (x 1) from the Devils River Limestone, Val Verde County, Tex., at locality 134. USGS 1941; 

USNM 306041.

16. Front view of the lectotype (x 1) from the same locality. USGS 1941; USNM 306042.

17. Polished axial section of a specimen (x 1.5) from the Washita Group near Big Lake, Reagan County, Tex., at locality 

136. USGS 11853; USNM 306001.

18. Back view of a syntype (x 1.5) from the Devils River Limestone, Vale Verde County, Tex., at locality 134. USGS 

1941; USNM 306043.

19. Aperture view of a syntype (x 1) from the same locality. USGS 1941; USNM 306044.
GEOLOGICAL SURVEY

15

PROFESSIONAL PAPER 1304 PLATE 9

/

/..v

12

. A

17

14

TROCHACTAEON (NEOCYLINDRITES)
PLATE 10

FIGURES 1-17. Trochactaeon (Mexicotrochactaeon) coniformis (Bose) (p. 56).

1. Aperture view (x 3) of a specimen from the Cardenas Formation, State of San Luis Potosi, Mexico, at locality 157. 

USGS 28216; USNM 306045.

2. Aperture view (x 1.5) of a specimen from the same locality. USGS 28216; USNM 306046.

3. Aperture view (x 3) of a specimen from the same locality. USGS 28216; USNM 306047. 

4, 5. Aperture and back views (x 2) of a specimen from the same locality. USGS 28216; USNM 306048.

6. Aperture view (x 1.5) of a specimen from the same locality. USGS 28216; USNM 306049.

7. View (x 1.5) of an incomplete specimen from the same locality, showing form of columellar plaits. USGS 28216; 

USNM 306050.

8. Aperture view (x 1.5) of a specimen from the same locality. USGS 28216; USNM 306051.

9. View (x 1.5) of an incomplete specimen from the same locality, showing development of columellar plaits. USGS 

28216; USNM 306052.

10. Aperture view (x 1.5) of a specimen from the same locality. USGS 28216; USNM 306053.

11. Aperture view (x 1) of an especially large specimen figured by Myers (1968, pi. 7, fig. 3) from the Cardenas Formation, 

State of San Luis Potosi, Mexico. UT (WSA) 155 54a.

12. Aperture view (x 1.5) of a specimen from the Cardenas Formation, State of San Luis Potosi, Mexico, at locality 157. 

USGS 28216; USNM 306054.

13. Axial section (x 3) of a specimen from the same locality. USGS 28216; USNM 306055.

14. Axial section (x 2) of a specimen from the same locality. USGS 28216; USNM 306056.

15. Axial section (x 2) of a specimen from the same locality. USGS 28216; USNM 306057.

16. Axial section (x 3) of a specimen from the same locality. USGS 28216; USNM 306058.

17. Aperture view (x 3) of a specimen from the same locality. USGS 28216; USNM 306059.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 10

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TROCHACTAEON (MEXICOTROCHACTAEON)
PLATE 11

FIGURES 1-4. Trochactaeon (Mexicotrochactaeon) coniformis (Bose) (p. 56).

1. Axial section (x 2) of a specimen from the Cardenas Formation, State of San Luis Potosi, Mexico, at locality 157. 

USGS 28213; USNM 306060.

2. Axial section (x 2.5) of a specimen from the same locality. USGS 28216; USNM 306061.

3. Axial section (x 2.5) of a specimen from the same locality. USGS 28216; USNM 306062.

4. Axial section (x 2.5) of a specimen from the same locality. USGS 28216; USNM 306057. 

5. Trochactaeon (Mexicotrochactaeon) palmeri Sohl and Kollmann, n. sp. (p. 60).

Axial section (x 1.5) of a paratype from the "Titanosarcolites limestone" of the Jerusalem Mountain inlier, Jamaica, at

locality 111. USGS 29466; USNM 306063. 

6, 7, 10. Trochactaeon (Neocylindrites) nelsoni Sohl and Kollmann, n. sp. (p. 55).

6. Axial section (x 4) of a paratype from the Pozas Formation, Orocovis Quadrangle, Puerto Rico, at locality 28. USGS 

30534; USNM 306064.

7. Axial section (x 4) of a specimen from the Cotui Limestone, San German Quadrangle, Puerto Rico, at locality 40.

USGS 30347; USNM 306065. 

10. Axial section (x 2) of a paratype from the Flor de Alba Limestone Lentil of Pozas Formation, Florida Quadrangle,

Puerto Rico, at locality 22. USGS 28629; USNM 306066. 

8, 9. Trochactaeon (Mexicotrochactaeon) wrighti Sohl and Kollmann, n. sp. (p. 59).

8. Axial section (x 1.5) of a paratype from the Cardenas Formation, State of San Luis Potosi, Mexico, at locality 152. 

USGS 24392; USNM 306067.

9. Axial section (x 2.5) of a paratype from the same locality. USGS 24392; USNM 306068. 

11. Trochactaeon (Neocylindrites) sp. A. (p. 55).

Axial section (x 2.5) of a specimen from the El Rayo Formation, Sabana Grande Quadrangle, Puerto Rico, at locality 36. 

USGS 29075; USNM 306069.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 11

8 \ f 9 

TROCHACTAEON (MEXICOTROCHACTAEON) AND T. (NEOCYLINDRITES)
PLATE 12

FIGURES 1-6, 10. Trochactaeon (Mexicotrochactaeon) wrighti Sohl and Kollmann, n. sp. (p. 59).

1. Aperture view (x 1.5) of a paratype from the Cardenas Formation, State of San Luis Potosi, Mexico, at locality 

152. USGS 24392; USNM 306070.

2. Aperture view (x 1) of a paratype from the same locality. USGS 24392; USNM 306071.

3. Axial section (x 1) of a paratype, showing (arrow) fold of palatal wall. Same locality. USGS 24392; USNM 306072. 

4, 5. Back and aperture views (x 1V2) of the holotype from the same locality. USGS 24392; USNM 306073. 

6. Axial section (x 1V2) of a paratype from the same locality. USGS 24392; USNM 306074. 

10. Aperture view (x 1) of a paratype from the "Titanosarcolites limestone," Marchmont inlier, Jamaica, at

locality 99. USGS 30462; USNM 306075. 

7-9, 11-13. Trochactaeon (Mexicotrochactaeon) burckhardti Bose (p. 58).

7, 9. Aperture and back views (x 1) of a specimen from the Maldon Limeston, Maldon inlier, Jamaica, at locality 119.

USGS 29955; USNM 306076.

8. Axial section (x 1) of a specimen from the Guinea Corn Formation, Central inlier, Jamaica, at locality 64. 

USGS 29978; USNM 306077.

11. Back view (x 1) of a specimen from the Maldon Limestone, Maldon inlier, Jamaica, at locality 119. USGS 

30018; USNM 306078.

12. Aperture view (x 1) of a specimen from the Guinea Corn Formation, Central inlier, Jamaica, at locality 67. 

USGS 29981; USNM 306079.

13. Aperture view (x 1) of a specimen from the Maldon Limestone, Maldon inlier, Jamaica, at locality 119. USGS 

30018; USNM 306080.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 12

TROCHACTAEON (MEXICOTROCHACT AEON)
PLATE 13

FIGURES 1-3, 6, 14. Trochactaeon (Mexicotrochactaeon) teaenis Sohl and Kollmann, n. sp. (p. 60).

1. Axial section (x 1) of a paratype from the Cardenas Formation, State of San Luis Potosi, Mexico, at locality 155. 

USGS 28187; USNM 306081.

2. Axial section (x 1) of a paratype from the Cardenas Formation, State of San Luis Potosi, Mexico, at locality 156. 

USGS 28218; USNM 306082.

3. Axial section (x 1) of a paratype from the "Titanosarcolites limestone," Jerusalem Mountain inlier, Jamaica, at

locality 109. USGS 30011; USNM 306083.

6. Axial section (x 1) of a paratype from the same locality. USGS 30422; USNM 306084. 

14. Axial section (x 1.5) of a paratype from the El Rayo Formation, San German Quadrangle, Puerto Rico, at locality

48. USGS 30354; USNM 306085. 

4, 5. Trochactaeon (Mexicotrochactaeon) rossi Sohl and Kollmann, n. sp. (p. 62).

4. Axial section (x 1) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, locality 55. USGS 

29454; USNM 306086.

5. Axial section (x 1) of a paratype from the "Titanosarcolites limestone," Marchmont inlier, Jamaica, at locality 99.

USGS 30462; USNM 306087. 

7-9. Trochactaeon (Mexicotrochactaeon) burckhardti Bose (p. 58).

7. Axial section (x 1) of a specimen from the Guinea Corn Formation, Central inlier, Jamaica, at locality 64. USGS 

29978; USNM 306088.

8. Axial section (x 1) of a specimen from the Maldon Limestone, Maldon inlier, Jamaica, at locality 119. USGS 29955; 

USNM 306089.

9. Axial section (x 1) of a specimen from the Guinea Corn Formation, Central inlier, Jamaica, at locality 58. USGS

31487; USNM 306090. 

10-13. Trochactaeon (Mexicotrochactaeon) granthamensis Sohl and Kollmann, n. sp. (p. 61).

10. Axial section (x 1) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 64, USGS 

30044; USNM 306091.

11. Axial section (x 1) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 60a. USGS 

30043; USNM 306092.

12. Axial section (x 1) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 68. USGS 

29982; USNM 306093.

13. Axial section (x 1) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 69. USGS 

29987; USNM 306094.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 13

11 12 13 

TROCHACTAEON (MEX1COTROCHACT AEON)
PLATE 14

FIGURES 1-5, 8. Trochactaeon (Mexicotrochactaeon) palmeri Sohl and Kollmann, n. sp. (p. 60).

1. Aperture view (x 1) of a paratype from the "Titanosarcolites limestone," Jerusalem Mountain inlier, Jamaica, 

at locality 108. USGS 30421; USNM 306095.

2. Back view (x 1) of a paratype from the same locality. USGS 30421; USNM 306096.

3. Aperture view (x 1) of a paratype from the Habana Formation, Santa Clara Province, Cuba, at locality 129. 

USGS 27278; USNM 306097.

4. Aperture view (x 1) of the holotype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 65. 

USGS 29979; USNM 306098.

5. Aperture view (x 1) of a specimen from the Habana Formation, Santa Clara Province, Cuba, at locality 129.

USGS 27278; USNM 306099. 

8. Axial section (x 2) of a paratype from the El Rayo Formation, San German Quadrangle, Puerto Rico, at locality

42. USGS 31488; USNM 306100. 

6, 7, 9-16. Trochactaeon (Mexicotrochactaeon) teaensis Sohl and Kollmann, n. sp. (p. 60).

6, 7. Back and front views (x 1) of a paratype from the El Rayo Formation, San German Quadrangle, Puerto Rico, at

locality 48. USGS 30354; USNM 306101. 

9, 10. Aperture and back views (x 1) of a paratype from the same locality. USGS 30354; USNM 306102.

11. Back view (x 1) of a paratype from the same locality. USGS 30354; USNM 306103.

12. Axial section (x 1.5) of a paratype from the same locality. USGS 30354; USNM 306104.

13. Aperture view (x 1.5) of the holotype from the same locality, USGS 30354; USNM 306105.

14. Axial section (x 1) of a paratype from the "Titanosarcolites limestone," Jerusalem Mountain inlier, Jamaica, at 

locality 109. USGS 30422; USNM 306106.

15. Aperture view (x 1) of a paratype from the Cardenas Formation, State of San Luis Potosi, Mexico, at locality 

156. USGS 28218; USNM 306107.

16. Aperture view (x 1) of a paratype from the "Titanosarcolites limestone," Jerusalem Mountain inlier, Jamaica, 

at locality 109. USGS 30011; USNM 306108.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 14

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14 15 

TROCHACTAEON (MEXICOTROCHACTAEON}
PLATE 15

FIGURES 1 11. Trochactaeon (Mexicotrochactaeon) granthamensis Sohl and Kollmann, n. sp. (p. 61).

1. Axial section (x 1) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 64. USGS 

30044; USNM 306091.

2. Aperture view (x 1) of a paratype from the same locality, showing the nature of the inner lip callus. USGS 30044; 

USNM 306108.

3. Aperture view (x 1) of a small paratype from the same locality. USGS 30044; USNM 306109.

4. Aperture view (x 1) of a paratype from the same locality, which has a proportionally low spire. USGS 30044; USNM 

306110.

5. Back view (x 1) of a paratype from the same locality. USGS 30044; USNM 306111.

6, 7. Front and back views (x 1) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 

68a. USGS 29985; USNM 306112.

8. Side view (x 1) of a paratype, showing repair of a shell breakage. Guinea Corn Formation, Central inlier, Jamaica, 

at locality 64. USGS 30044; USNM 306113.

9. Axial section (x 1) of a specimen from the same locality. USGS 30044; USNM 306114. 

10, 11. Aperture and side view (x 1) of the holotype from the same locality. USGS 30044; USNM 306115.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 15

TROCHACTAEON (MEXICOTROCHACT AEON)
PLATE 16

FIGURES 1-9. Trochactaeon (Mexicotrochactaeon) rossi Sohl and Kollmann, n. sp. (p. 62).

1, 2. Back and aperture views (x 1) of a specimen figured by Myers (1968, pi. 7, fig. 4) as Actaeonella coniformis Bose.

UT (WSA) 15585. 

3. Aperture view (x 1) of a paratype from the "Titanosarcolites limestone," Marchmont inlier, Jamaica, at locality

99. USGS 30462; USNM 306116. 

4, 5. Back and aperture views (x 1) of a paratype from the same locality. USGS 30462; USNM 306117.

6. Axial section (x 1) of a paratype from the same locality. USGS 30462; USNM 306118.

7. Aperture view (x 1) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 78. USGS

30141; USNM 306119. 

8, 9. Back and front views (x 1) of the holotype from the "Titanosarcolites limestone," Marchmont inlier, Jamaica, at

locality 99. USGS 30462; USNM 306120. 

10-13. Trochactaeon (Mexicotrochactaeon) melonensis Sohl and Kollmann, n. sp. (p. 63).

10, 11. Back and front views (x 1) of a paratype from the Melones Limestone, Cabo Rojo Quadrangle, Puerto Rico, at 

locality 13. USGS 30565; USNM 306121.

12. Axial section (x 1.5) of a paratype from the Melones Limestone, Puerto Real Quadrangle, Puerto Rico, at locality 

33. USGS 29354; USNM 306122.

13. Aperture view of the holotype (x 1) from the Melones Limestone, Cabo Rojo Quadrangle, Puerto Rico, at locality 

13. USGS 30565; USNM 306123.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 16

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PLATE 17

FIGURES 1, 4, 5. Actaeonella robinsoni Sohl and Kollmann, n. sp. (p. 73).

1. Axial section (x 1%) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 69. USGS 

29987; USNM 306124.

4. Axial section (x 1V2) of a paratype from the same locality. USGS 29987; USNM 306125.

5. Axial section (x 2) of a paratype from the same locality. USGS 29987; USNM 306126.

2. Actaeonella cubensis Kollmann and Sohl, n. sp. (p. 72).

Axial section (x 3) of a paratype from the Habana(?) Formation, Oriente Province, Cuba, at locality 125. USGS 19465. 

USNM 306127.

3. Actaeonella n. sp. A. (p. 73).

Axial section (x Vz) of a specimen from the Maravillas Formation, Coamo Quadrangle, Puerto Rico, at locality 19. USGS

26834; USNM 306128. 

6-8. Trochactaeon (Mexicotrochactaeon) melonensis Sohl and Kollmann, n. sp. (p. 63).

6. Axial section (x 1%) of a paratype from the Melones Limestone, Cabo Rojo Quadrangle, Puerto Rico, at locality 13. 

USGS 30565; USNM 306122.

7. Axial section (x 1V2) of a paratype from the same locality. USGS 30565; USNM 306129.

8. Axial section (x 1V2) of a paratype from the same locality. USGS 30565; USNM 306130.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 17

ACTAEONELLA AND TROCHACTAEON (MEXICOTROCHACTAEON)
PLATE 18

FIGURES 1-8. Actaeonella browni Sohl and Kollmann, n. sp. (p. 67).

1, 2. Back and aperture views (x 2) of the holotype from Albian Limestone, Peten, Guatemala, at locality 160. USGS 

28327; USNM 306131.

3. Back view (x 2) of a paratype from the same locality. USGS 28327; USNM 306132.

4. Aperture view (x 2) of an incomplete paratype from the same locality. USGS 28327; USNM 306133.

5. Back view (x 1) of a paratype from the same locality. USGS 28327; USNM 306134.

6. Aperture view (x 2) of a paratype from the same locality. USGS 28327; USNM 306135.

7. Aperture view (x 2) of a paratype from the same locality. USGS 28327; USNM 306136.

8. Aperture view (x 2) of a paratype from the same locality. USGS 28327; USNM 306137. 

9-14. Actaeonella pecosensis Stanton (p. 65).

9. Aperture view (x 6) of a small specimen from the Edwards Limestone of Williamson County, Tex., at locality 146. 

146. UT 1036TX6.

10. Back view (x 1) of a paratype from the Devils River Limestone, Kinney County, Tex., locality 139. USGS 11885; 

USNM 77561a.

11. Back view (x 2) of a specimen from the Devils River Limestone, Val Verde County, Tex., at locality 134. USGS

1941; USNM 306138. 

12, 13. Back and aperture views (x 1%) of the holotype from the same locality. USGS 1941; USNM 77560.

14. Axial section (x 1%) of specimen from the same locality. USGS 1941; USNM 306139. 

15, 16. Actaeonella oviformis Gabb (p. 68).

Back and aperture views (x 1) of the holotype from Cottonwood Creek, Shasta County, Calif. ANSP 4323. 

17-20. Actaeonella delgadoi (Choffat) (p. 66).

17-20. Aperture views (x 2) of specimens from the Walnut Formation at unspecified localities in Texas. Renfro collection.

USNM 306140, 306141, 306142, 306143. 

21, 22. Actaeonella cf. A. sublaevis Briart and Cornet (p. 66).

21. Aperture view (x 1) of a specimen near Kent, Culberson County, Tex., at locality 133. USGS 1926; USNM 306144.

22. Back view (x 1) of an incomplete specimen from the Edwards Limestone of Williamson County, Tex., at locality 

146. UT 1036 TX5.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 18

17 20 

ACTAEONELLA

22
PLATE 19

FIGURES 1-8 Actaeonella duckettsensis Sohl and Kollmann, n. sp. (p. 69).

1. View (x 1) of incomplete paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 62. USGS 

29973; USNM 306145.

2. Axial section (x IVk) of a paratype from the "Barrettia Limestone," Green island inlier, Jamaica, at locality 121. 

USGS 30007; USNM 306146.

3. Aperture view (x IVk) of a compressed paratype from the "Titanosarcolites limestone," Marchmont inlier, 

Jamaica, at locality 97. USGS 30026; USNM 306147.

4. Aperture view (x 1V2) of a paratype from the same locality. USGS 30026; USNM 306148.

5. Aperture view (x 1) of a paratype from the same locality. USGS 30026; USNM 306149. 

6, 7, 8. Aperture, back, and basal view (x 1%) of the holotype from the "Titanosarcolites limestone," Marchmont inlier,

Jamaica, at locality 106. USGS 30455; USNM 306150. 

9-15. Actaeonella jicarensis Sohl and Kollmann, n. sp. (p. 69).

9, 10. Back and aperture views (x 2) of a paratype from the El Rayo Formation, Sabana Grande Quadrangle, Puerto

Rico, at locality 36. USGS 28664; USNM 306151.

11. Back view (x 2) of a paratype from the same locality. USGS 28664; USNM 306152. 

12, 13. Aperture and front views (x 3) of a paratype from the same locality. USGS 28664; USNM 306153.

14. Aperture view (x 2) of the holotype from the same locality. USGS 28664; USNM 306154.

15. Axial section (x 3) of a paratype from the same locality. USGS 28664; USNM 306155. 

16-21. Actaeonella borneensis Nuttall and Leong (p. 67).

16. Axial section (x 1) of a specimen from near Ocozocuautla, State of Chiapas, Mexico, at locality 150. USNM 306156.

17. Aperture view (x 1) of a specimen from near Ocozocuautla, State of Chiapas, Mexico, at locality 149. 

USNM 306157.

18. Aperture view (x 1) of a specimen from the same locality. USNM 306158.

19. Axial section (x 1) of a specimen from Santa Clara Province, Cuba, at locality 130. USGS 27295; USNM 306159.

20. View (x 1) of an incomplete specimen from near Ocozocuatla, State of Chiapas, Mexico, at locality 149. USNM 

306160.

21. Axial section (x 1) of a specimen from the same locality. USNM 306161.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 19
PLATE 20

FIGURES 1, 2, 5, 8. Actaeonella coquiensis Sohl and Kollmann, n. sp. (p. 70).

1. Axial section (x 4) of a paratype from the Coamo Formation of Berryhill and Glover (1960), Central Aguirre 

Quadrangle, Puerto Rico, at locality 16. USGS 29090; USNM 306162.

2. Axial section (x 3) of a paratype from the same locality. USGS 29090; USNM 306163.

5. Axial section (x 2V£) of a paratype from the Stapleton Formation, Sunderland inlier, Jamaica, at locality 122.

USGS 30014; USNM 306164. 

8. Axial section (x 2V£) of a paratype from Coamo Formation of Berryhill and Glover (1960), Central Aguirre

Quadrangle, Puerto Rico, at locality 16. USGS 29090; USNM 306204. 

3, 6. Actaeonella duckettsensis Sohl and Kollmann, n. sp. (p. 69).

3. Axial view of a paratype (x 1V&) from the "Barrettia Limestone," Green Island inlier, Jamaica, at locality 121. USGS

30007; USNM 306146. 

6. Axial view of a paratype (x 4) from the Slippery Rock Formation, Central inlier, Jamaica, at locality 53. USGS

30032; USNM 306165. 

4. Actaeonella briggsi Sohl and Kollmann, n. sp. (p. 72).

Axial view of a specimen from the Pozas Formation, Barranquitas Quadrangle, Puerto Rico, at locality 4. USGS 29365;

USNM 306166. 

7. Actaeonella jicarensis Sohl and Kollmann, n. sp. (p. 69).

Axial view (x 3) of a paratype from the El Rayo Formation, Sabana Grande Quadrangle, Puerto Rico, at locality 36.

USGS 28664; USNM 306155. 

9. Actaeonella cf. A. coquiensis Sohl and Kollmann (p. 71).

Axial view of a specimen (x 1.25) from the Melones Limestone, Puerto Real Quadrangle, Puerto Rico, at locality 32. 

USGS 30537; USNM 306167.

10. Actaeonella pecosensis Stanton (p. 65).

Axial view (x \Vz) of a specimen from the Devils River Limestone, Val Verde County, Tex., at locality 134. USGS 1941; 

USNM 306139.

11. Actaeonella borneensis Nuttall and Leong (p. 67).

Axial section (x 1.25) of a specimen from the upper Rancho Nuevo Beds, State of Chiapas, Mexico, at locality 149. 

USNM 306161.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 20

ACTAEONELLA
PLATE 21

FIGURES 1-6. Actaeonella marchmontensis Sohl and Kollmann, n. sp. (p. 71).

1. Aperture view (x 6) of a paratype from the "Titanosarcolites limestone," Marchmont inlier, Jamaica, at 

locality 97. USGS 30026; USNM 130668.

2. Aperture view (x 6) of a paratype from the same locality. USGS 30026; USNM 130669.

3. Aperture view (x 4) of a paratype from the same locality. USGS 30026; USNM 130670.

4. Aperture view (x 4) of the holotype from the same locality. USGS 30026; USNM 130671.

5. Aperture view (x 4) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 64. 

USGS 30044; USNM 130672.

6. Aperture view (x 4) of a paratype from the "Titanosarcolites Limestone," Marchmont inlier, Jamaica, at

locality 97. USGS 30026; USNM 130673. 

7-12, 15. Actaeonella coquiensis Sohl and Kollmann, n. sp. (p. 70).

7. Aperture view (x 3) of a paratype from the Coamo Formation of Berryhill and Glover (1960), Central Aquirre 

Quadrangle, Puerto Rico, at locality 16. USGS 30358; USNM 306186.

8. Aperture view (x 2) of a paratype from the same locality. USGS 30358; USNM 306187.

9. Aperture view (x 2) of the holotype from the same locality. USGS 30358; USNM 306188.

10. Aperture view (x 4) of a paratype from the same locality. USGS 30358; USNM 306189.

11. Aperture view (x 2) of a paratype from the same locality. USGS 30358; USNM 306190.

12. Aperture view (x 5) of a paratype from the same locality. USGS 30358; USNM 306191. 

15. Back view (x 5) of a paratype from the Coamo Formation of Berryhill and Glover (1960), which shows borings 

in shell. Central Aguirre Quadrangle, Puerto Rico, at locality 18. USGS 30541; USNM 306192.

13. 19-23. Actaeonella cf. A. coquiensis Sohl and Kollmann (p. 71).

13. Back view (x 1) of an incomplete specimen from the Melones Limestone, Puerto Real Quadrangle, Puerto 

Rico, at locality 32. USGS 30537; USNM 306193.

19. Axial section (x \Vz) of a specimen from near central Soledad, Santa Clara Province, Cuba, at locality 127. 

USGS 27266; USNM 306194.

20. Axial section (x 1%) of a specimen from the same locality. USGS 27266; USNM 306195.

21-23. Axial sections (x 1) of specimens from the Melones Limestone, Puerto Real Quadrangle, Puerto Rico, at 

locality 32. USGS 30537; USNM 306196-306198.

14. 16-18. Actaeonella briggsi Sohl and Kollmann, n. sp. (p. 72).

14. Aperture view (x 3) of a paratype from Pozas Formation, Barranquitas Quadrangle, Puerto Rico, at locality 4.

USGS 29365; USNM 306199. 

16, 17. Aperture and back view (x 2) of a paratype from El Rayo Formation, San German Quadrangle, Puerto Rico, at

locality 48. USGS 30354; USNM 306200.

18. Aperture view (x 2) of the holotype from the Pozas Formation, Barranquitas Quadrangle, Puerto Rico, at 

locality 4. USGS 29365; USNM 306201.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 21

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PLATE 22

FIGURES 1, 2, 5. Actaeonella robinsoni Sohl and Kollmann, n. sp. (p. 73).

1. Aperture view (x IVk) of a paratype from the Cardenas Formation, State of San Luis Potosi, Mexico, at locality 158. 

USGS 28219; USNM 306174.

2. Axial section (x 1) of a paratype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 70. USGS 29988;

USNM 306175. 

5. Axial section (x 1) of the holotype from the Guinea Corn Formation, Central inlier, Jamaica, at locality 69. USGS

29987; USNM 306176. 

3, 4. Actaeonella n. sp. A. (p. 73).

Axial section and aperture view (x 1) of a specimen from the Maravillas Formation, Coamo Quadrangle, Puerto Rico, at 

locality 19. USGS 26834; USNM 306177.

6. Actaeonella sp. (p. 74).

Axial section (x IVk) of a specimen from the Barrancas Limestone Member of the Magueyes Formation, Barranquitas 

Quadrangle, Puerto Rico, at locality 12. USGS 31363; USNM 360178.

7. Actaeonella sp. B (p. 74).

Aperture view (x 4) of a specimen from glacial erratic cobbles of sideritic ironstone concretions on Martha's Vineyard, 

Mass., at locality 131. USGS 9568; USNM 360179.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 22

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ACTAEONELLA
PLATE 23

FIGURES 1, 6. Actaeonella n. sp. A. (p. 73).

1. Axial section (x 1) of a specimen from the Maravillas Formation, Coamo Quadrangle, Puerto Rico, at locality 19.

USGS 26834; USNM 306180. 

6. Axial section (x 1) of a specimen from the Pozas Formation, Orocovis Quadrangle, Puerto Rico, at locality 26.

USGS 30532; USNM 306181. 

2-5, 7, 8. Actaeonella cubensis Sohl and Kollmann, n. sp. (p. 72).

2. Aperture view (x 1M>) of a paratype from the Habana(?) Formation, Oriente Province, Cuba, at approximately 

locality 125. USGS 19464; USNM 306182.

3. Aperture view (x 1V2) of the holotype from Habana(?) Formation, Oriente Province, Cuba, at locality 125. USGS

19465; USNM 306183. 

4, 5. Axial sections (x 3 and x 6) of a paratype from the same locality. USGS 19465; USNM 306184.

7. Aperture view (x 2) of a paratype from the same locality. USGS 19465; USNM 306185.

8. Aperture view (x 2) of a paratype from the same locality. USGS 19465; USNM 306186.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1304 PLATE 23

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