ISSN 0120.

078X
Publicaciones .

Geolo_gicas
Especiales
Del
lngeomina s

By
FERNANDO ETA YO-SERNA
UNIVERSIDAD NACIONAL DE COLOMBIA

No.2,pp. 1-186,1979
Bogota - Colombia Publ. Geol. Esp.
ISSN- 0120- 078X Ingeominas
 PRESENTACION

Podni parecerles extrafio a los lectores que se publique en Colombia un tra­

bajo en ingles. Esta decision del Comite de Publicaciones del INGEOMINAS responde a
consideraciones puramente practicas. Efectivamente, traducir un trabajo tan extenso
como el del profesor Fernando Etayo significa un esfuerzo enorme; por otra parte, su
nivel y su especializacion lo destinan fundamentalmente a ser leido por profesionales
de Ia geologia. "··

Gustenos o no, el ir.gles se ha vuelto el idioma del intercambio cientifico,

una especie de latin del siglo XX. Todo profesional de las Ciencias de Ia Tierra -y me
atreverfa a decir, de las otras Ciencias-· que no quiera permanecer aislado de los ultimos
avances en su campo debe absorber mensualmente varias decenas de articulos especiali­
zados.

La inmensa mayorfa de esta produccion, que a muchos les parece desmedida,

es publicada en ingles.

De manera que a menos de contentarse con traducciones de textos que ge­

neralmente tienen varios afios de retraso o con trabajos sabre temas locales publicados
en su propio idioma, todo ge6logo que intente mantenerse razonablemente informado
debe leer en ingles.

INGEOMINAS tiene como una de sus funciones mas importantes el difundir·

los conocimientos que existan acerca de la Geologia de Colombia. En esta oportunidad,
el Instituto presenta a la comunidad geologica el excelente trabajo del profesor Fernan­
do Etayo, realizado para optar el titulo de PhD. en la Universidad de California de
Berkeley. No quiero caer en Ia academica costumbre, tan difundida en nuestro media,
de lanzarme en un elogio rimbombante de mi buen amigo Fernando; ofenderia ademas
su tradicional modestia. Sin embargo es buena que los gedlogos colombianos·sepan que
en este momenta Fernando, gracias al Convenio celebrado entre Ia Universidad Nacio­
nal y el INGEOMINAS, esta desarr ollando con ge6logos de esta ultima entidad y estu­
diantes de la primera una intensa labor de corr elacion estratigrruica de las rocas cr�ta­
ceas de la Cordillera Oriental, cuya programacion inicial es de dos ailos.

Estoy convencido del gran avance que significara este trabajo para la geologia
colombiana.

Este Instituto se ha propuesto publicar en forma sistematica todos los traba­

jos ineditos acerca de la geolog!a en nuestro pais. Esta en imprenta la tesis de doctora­
do de Daria Barre ra y seguiran, en esta colecci6n de Publicaciones Geologicas Especia­
les, otras contribuciones ineditas tanto de colombianos como de extranjeros, en espa-
. fi.ol y en ingles. Espero que los estudiosos de la geologia colombiana no vean en este
esfuerzo un intento extranjerizante sino una clara decision de robustecer el desarrollo
de las Ciencias de la Tierra en Colombia.

Michel Hermelin

Junia de 1979
 PUBLICACIONES GEOLOGICAS ESPECIALES DEL
""
INGEOMINAS

ZONATION OF THE

CRETACEOUS OF CENTRAL

COLOMBIA BY AMMONITES

By
FERNANDO ETA YO-SERNA
UNIVERSIDAD NACIONAL DE COLOMBIA
(Departamento de Geociencias, e I.C.N.M.H.N.)

MINISTERIO DE MINAS Y ENERGIA

INSTITUTO NACIONAL DE INVESTIGACIONES GEOLOGICO·MINERAS
1979

Bogota-Colombia, No, 2, pp, 1·188 ISSN • 0120 · 078X

Precio de cada ejemplar: $ 300.00
us $ 7.50

Publicacion no periodica
Fonnato de Publicacion: 21.5 x 27.5 em

Derechos Reservados por:

INGEOMINAS: lnstituto Nacional de Investigaciones Geologico- Mineras
Carrer a 30 No. 51-59
Apartado Aereo No. 4865 ·Bogota- Colombia

All the concepts emitted in this publication are of the entire responsability of the author.

Edited and printed by Ingeominas.

 CONTENTS

Page

ABSTRACT . .. . ·
. . . . . . . • • . • • . . . . . . • . . . . • . . . . . . . . • • . . . . • • • • . . . . • • , 1
:� ,_�:-i.c:}._.•.
.

INTRODUCTION 2
:-�-� . -ri':t•·,:j.!.:._- -
' - . .-;
. . • . . . . . . . . • . • • • . • . . . . . . • . • • . . . . • . . . . • . . . • • •

; , . -. �
.

ACKNOWLEDGMENTS- 3
.......
• • . . • • • • . • • • • • • . • . . • -. • . . • • • • . • . • • • •

PHYSICAL STRATIGRAPHY • . . . • '" ' . • •• • • • . . . • . • • . . • . • . . • • . • . •,.• .,�"ii.... 4

I. THE CRETACE OU S SECTION FROM VILLA DE LEIVA TO LOMA PIEDRA

GORDA, LA RGELY FROM ETAYQ-SERNA,1972. . . . . . . . . . . . . . . . . 4

PAJA FORMATION . . . . . • • , . . . . . . . . . • . . . . • , • . • : . . . . . . ... . . . . 4

SAN GIL GROUP . . . . . . . • . - . • . . . . . . . . . . . . . . . . . . . . . . • . . . . . .. . . . 4
CHURUVITA GROUP • . . . . . . ·- · . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
CHURUVITA FORMATION . . . . . . • . . • . . . . , . . • . . . • , . • . , ....... ,. 7
SAN RAFAEL FORMATION . . . . • .. . • • • . . . .. .
. . • . • . . • . . .. . . • . • ,.. 7
CONEJO FORMATION . . . . . . . . . . . . . . • . . . . • . ; .......... , .. , . . . 7
PLAENERS FORMATION. • • . . . . .. . . . • . ,. . . . ..... . . . . . . . . . ... . 8
GUADALUPE FORMATION . . , . . . . . . . . . . . . . . . . . . . . . . . . . • . . . • . . 8

II. THE CRETACEOUSSECTION IN THE APULO, VIOTA, LAS MESITAS REGION 8

LA NAVETA FORMATION . . . . . . . . . . . . . . . . . . . . . . . . . . .. . .. . ,, .. 8
VILLETA GROUP . . . . . . . . . . • . . . . • . . . . . . • . . . . . . . . • . . . . . • . . . . 8
SOCOT A MEMBER . . . . . . . . . . . . . . . . . . . . . . • • . • . . . . . . . . , , . . . . . . 9
CAPOTES MEMBER • , . . . . , • . . • . . . . , . . . . . . . . . . . . . . . . . . . . • . . . . 10
HILO FORMATION • . . . . . . . . . . . . . . . . . . . . . . . . . . • . . . . . . . . . • • .. 11
LA FRONTERA FORMATION. . . . . . . . . • . . • . . . . . . . . • .. . . . . . . . . . . 11
COMMENTS . . . . . . . . . • . . . . . • . . . . . . . . . . . • . . . . . . . . . . . . . . . . . • 11

BIOSTRATIGRAPHY . . . . . • . . . . . . . . . • • . . . . . . • . . . . . . . • . . . . . . . • . . . . . . 13

ASSEMBLAGE ZONES PROPOSED FOR USE IN COLOMBIA. . • . . . . . . • . . . 13

Du{renoyia sanctorum-Stoyanowiceras tre{{ryanus Assemblage-
zone ........................................ 13
Parahoplites (?) hubachi- Acanthohoplites (?) /eptocerati{orme
Assemblage-zone . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . • . 13
Douvilleiceras solitae-Neodeshayesites co/umbianus Assemblage-
zone........................................ 14
Platiknemiceras co/ombiana-Rinconiceras rinconi-Lyelliceras
pseudolyelli{orme Assemblage-zone . . . . . . . . . . . . . . . . . • • 14
Mammites nodosoidesappelatus-Franciscoites suarezi Assembla-
ge-zone . • . . . . . • • . . . . • . . . • , . . • . . . . . . . • .. . . . . . . 14
G/oriaceras correai-Protexanites cucaitaense - Codazziceras
scheibei Assemblage-zone ...... : . . . • . . • . . . . . . . . . . . • 15
Lenticeras baltai-Paralenticeras sieversi Assemblage-zone.. . . . . · ·15

TABLE I................................................. 16
·i-
 Page

SYSTEMATIC DESCRIPTIONS • • . • • • • . . . . . . • . • • . . • . • • . • • . . • . . · · • · • . • . • 17

Order AMMONOIDEA . . . . . . . . . • . . . . . . . • . . . . . . . . . . . . . . . . . . • . • 17
Family PHYLLOCERATIDAE.. . . . . . . . • • . . . . . . . • . . • . . . . . • . . . • . .. 17
Family GAUDRYCERATIDAE.................................. 19
Family ANCYLOCERATIDAE .................................. 20
Family PTYCHOCERATIDAE................................... 21
Family ACONECERATIDAE . . . . • . . • . . . . . . . . • . . . . . . . . • . . • . . • • .. 22
Family DESMOCERATIDAE . . . . . . . . . • . . . . . . . . . . . . • . . . . . . . .. . . . 23
Family PACHYDISCIDAE . . . . . . • . . • . . • . . . . . • . . . . . • . . • . . • • . • . .. 30
Family TROCHLEICERATIDAE. . . • . . • . . • • . • . . . . . . . . . . . . . . • • • .. . 31
Family DOUVILLEICERATIDAE.. . . . . . . . . . . . • . . . . . . . . . . • • . .. . .. 32
Family DESHAYESITIDAE . . . . . . • . • . . . . . • . . • • . . . . . . . . . . . . . • . . . 58
Subfamily PARAHOPLITINAE............. . . . . . . . . • . . . . . . . . . . . • 72
Family ENGONOCERATIDAE � . . . . . . .... • . . • . . . . . . . . . . . • . . . . . • . 74
Family BRANCOCERATIDAE ......... . . . . . . . • . . . . . • • . . . . • . . ... 78
Family LYELLICERATIDAE . . . . . . . . . . . . . • . . . . . . . . . : ... ... . .. . . 79
Family ACANTHOCERATIDAE ................................. 83
Family MAMMITIDAE. . . • . . . . . . . . . . . . . . : . . . . . . • • • . . .. . . . . .. . . 85
Family VASCOCERATIDAE. . . • . . . . • . . • . . . . . . . . . . . . . . . . . .. . . . . . 87
Family COILOPOCERATIDAE.................................. 90
Family COLLIGNONICERATIDAE. . ... .. . • . . . • . . . .. . . . . . . . . . . . . . 92
Order NAUTILIDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Family CYMATOCERATIDAE................ .................. 104

SUMMARY . . . . . . . . . . . . . • .. . . . • • . . . . . . . . . • . . . • . . . . . . • . . . . • • . • • .. 105
RESUMEN • • • . • . . . . . • • . • • . . • . • • . • . • • • • . . • . . • • • . . . ; • . • . . . . . . . • . . • 106
RESUME. .. ... . . . . . ..... . . . ... . .... .. . ... ...... ...... .. .... ... . 107
RIASSUNTO • . . . • . . . • • . • . . . . • . . . . • . . . . . • . . • . • . . • • . • • . • . . • . . . . • • . 108
ZUS.AIDIENFASSUNG . . . . . . • . . • . . . . . . • . . . • . . • . . • . . . . . • . . • • • . .. . . . . 109
REFERENCES.... .. .... .... . . . . . .... ... . . .. .. ... ... .. ... . . . ...... 110
'INDEX OF GENERA AND SPECIES. . . . . • . • . • . . • . • • . • • • . . . . . . . . . • . . . • •. • 182

ILLUSTRATIONS
Text-Fig. 1. Map of Colombia showing location of sections investigated in detail • • . . 5

Text-Fig. 2. Composite columnar sections Villa de Leiva · Apulo. • . • • . . . • • . . . . . 6

Text-Figs;�--·3 • 9. Sutures and whorl sections of the studied ammonites . . . . . • • . . . . • .. 12C

Text-Figs. 10·17. Scattem diagrams of some of the studied ammonites . . • . . . . . . . .. . . 13�

Plate 1. Ammonites from the Dufrenoyia sanctorum·Stoyanowiceras treffryanus

Assemblage zone. Figs.1-8. Upper Aptian. 15:

Plate 2. Ammonites from the Du(renoyia sanctorum-Stoyanowiceras tre(fryanus ·

Assemblage zone . Figs.1-10. Upper Aptian. 15

Plate 3. Ammonites from the Dufrenoyitz sanctorum-Stoyanowiceras treffryanus

Assemblage zone. Figs.1-2, 4-8, 10·13. Ammonites from the Parahoplites
('?) hubachi-Acanthohoplites ('?) leptocerati(orme Assemblage zone.Figs.
3, 9.Upper Aptian. --�

.-11·
Pl ate 4 . Ammo nite s fro m t he Du(renoyia sanctorum-Stoyanowiceras treffryanus.
As se mblage zo ne. F igs. 1- 3, 5, 7- 8. Ammo nite s fro m the Prahoplites (?)
hubachi-Acanthohoplites (?) Jeptocerati(orme Asse mbl age zo ne . F igs .
4, 6. Uppe r Apt ian. . . . . . . . . . • . . . . . . . . . . . . . . . . . . . • . . . . . . 158

P late 6. Ammo nites fro m the Du(renoyia sanctorum-Stoyanowiceras tre(fryanus

Assel!l blage zo ne . Fi gs. 3, 9. Ammo ni tes fro m the Parahoplfte11 (?) huba­
chi·Acanthohoplites (?) leptocerati(orme Asse mblage zo ne . l'' igs. 1- 2, 4- 8.
Uppe r Ap tian . . . . . . • . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160

P late 6 . Ammo nite s f ro m t he Duf�enoyia sanctorum-Stoyanowiceras treffryanus

Asse m blage z o ne. Fi gs. 1- 20. Upper Ap ti an • . . . . . . . . . . . . . . . . . . . 162

P late 7 . Ammo nites fro m t he Du(renoyia sanctorum-Stoyanowiceras tre{fryanus

As se mblage z o ne . Fi g. 9 . Ammo n ite s fro m the Parahoplites (?) hubachi­
Acanthohoplites (?) leptocerati{orme Asse mblage zo ne . F igs; 1- 4, 8.
Uppe r Ap ti an . Am mo ni tes fro m t he Douvilleiceras solitae-Neodeshaye-
sites columbianus Asse mblage zo ne . Fi gs. 5-7 . Lo wer Albian • . . . . . . . � 164

P late 8 . Ammo nite s fro m t he Douvilleiceras solitae-Neodeshayesites columbianus

As se mbl age zone . Fi gs. 1-7 . Lo wer Albian . . . • . . . . . . . • . . . . . . .. . . 16. 6

Plate 9 . Ammo ni tes fro m the Douvilleiceras solitae-Neodeshayesites columbianus

Asse mbl age zo ne . Fi gs. 1- 3 , 7- 8. Lo we r Al bian . Ammonites fro m t he Pla­
tiknemiceras colombiana-Rinconiceras rinconi-Lyelliceras pseudolyelli-
{orme Asse mblage zone . Fi gs. 4-6 . M idd le Albian .. . . . . . . . . , ·. ·
. . . . . . . 168

P late 10. Ammonite s fro m t he Parahoplites (?) hubachi-Acanthohoplites (?) lepto­

cerati{orme Asse mbl age zo ne . Fi gs . 2- 3. Uppe r Aptian . Ammo nite s fro m
the Douvilleiceras so/itae-Neodeshayesites columbianus Asse mbl age zo ne .
Fi gs. 1, 3-10 . Lo wer Al bian . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 0

P late 1 1. Ammo nite s fro m· t he Platiknemiceras colombiana-Rinconiceras rinconi-

Lyelliceras pseudo/yelli{orme Assembl age zone. Fi gs. 1-9. Midd le Alb ian . 17 2

P late 12. Ammonites fro m the Plaliknemiceras colombiana-Rinconiceras rinconi-

Lyelliceras pseudolyelli{orme Asse mblage zone . F igs. 1· 4 . M idd le Al bi an . 174

P late 1 3. Ammo nite s fro m t he Mammites nodosoidesappelatus-Franciscoites sua-

rezi As se mblage zone . F igs. 1-1 1, 1 3- 15 . Lo we r Turon ian . Ammonites
fro m the Gloriaceras correai-Protexanites cucaitaense-Codazziceras schei-
bei Asse mbl age zo ne. F igs. 12, 16- 19 . Lo we r Con iacian . . . . . . . . . . . . 176

P late 14. Ammoni tes from theMammites nodosoidesappelatus-Franciscoites suarezi

Asse mblage zo ne . Fi g. 5. Lo wer Tu ro nian. Ammo nite s fro m t he Gloria-
ceras correai-Prolexanites cucaitaense·Codazziceras scheibei Asse mblage
zo ne. Fi gs. 1- 4, 7- 10 . Lo we r Co ni aci an . . . . . . . . . . . . . . . . . . . . . . . . 17 8

P late 15. Ammon ite s from t he Gloriaccrus correai-Protexanites cucaitaense-Co-

dazziceras scheibei As se mblage zone. Fi gs 1- 4. Lo wer Co ni acian. . . . . . . 180

* * *
 ZONATION OF THE CRETACEOUS OF CENTRAL
ABSTRACT
Cretaceous rocks ranging in age from
. Upper Aptian to Lower Santonian � llve been
carefully mapped, measured and colleoted bed
by bed in two regions (it Central Colombia na­
mely_: Villa de Leiva (Department Boyaca) and
Apulo ( formerly Rafael Reyes, Department
Cundinamarca).
·
The first mentioned sequence comprises
about 2.000m of locally highly fossiliferous
shales, although limestones, sandstones and
chert beds are interspersed and sporadically
yielded fossils. At Apulo the sequence consists
dominantly of shales with locally abundant
fossiliferous calcareous "concretions" and one
intercalcation of fossiliferous graded-bedded­
sandstones (turbidites) with a total thickness of
about l.OOOm.
The lithostratigraphic terminology utilized
is that previously proposed and published by
the writer, the old terminology is reevaluated
and some new units are introduced and their
type sections and reference sections described.
Similarly the occurrence of ammonites is indi­
cated.
The core of the investigation is however
the study and description of ammonite taxa:
102 species assigned to 52 genera were thoro­
ughly investigated, of them 85% of the species
and 21% of the genera are considered to repre­
sent new taxa. A zonal scheme consisting of
7"A'iSemblage zones" is proposed, based on the
succession of ammonites in these two sections,
hopefully it will be a yardstick for the Creta­
ceous biostratigraphy of the· northern part of
South America. No close correspondence be­
tween lithostratigraphic boundaries and bios-
* *
Caminante no h ay camino ,
se h a c e camino a/ andar.
Yo u ask me th e sec re t
It only has a word,again.
But all t h is is u nrip e fruit. Our sch ola rs will some day know more than th eir masters do
now; so let us patiently continue o u r work and rema in friends.
COLOMBIA BY AMMtlNITES 1
tratigraphic units has been found, however
some taxa of ammonites wit�eculiar morpho­
logic characteristics seem to-�'associated with
definite lithofacies.
The diverse taxa of ammonites were stu­
died through dissection of their conchs in order
to determine their ontogenetic changes and all
morphological characteristics were evaluated in
an attempt to understand each taxon indepen­
dently. From the taxonomic investigations and
the known stratigraphical distribution of the
faunas it is shown once more that similar--al­
though not 'identical' -morphologic types of
ammonites recurred during Cretaceous times.
The taxonomy resulting from· the-criteria
utilized herein indicate that the described taxa
have proportionally a distinctly South American
character, however some ubiquitous elements,
several of them previously unknown from the
Cretaceous of South America are recorded; as a
possible biological explanation for this fact it is
hypothesized, that as in the case of modem ce­
phalopods the ubiquitous elements of the ammo­
noid faunas may correspond to those hatched
from small eggs (with small protoconchs) and
vice versa.
All the species described are illustrated by
drawings of their whorl cross-section and suture
line, and at least one photograph of each.
Conch measurements are expressed in
mm. The symbol D represents the diameter
of a volution. Euw is the external umbilical
width. H is the maximum height and W the
maximum width of the whorl.
*
Me ha beis preguntado que h ila e l
cru stacea entre sus patas de oro
y os re spondo: e/ mar lo sa b e.
(MACHADO, HOLUB, NERUDA, poets)
E. S u e ss.
2 F. ETAYO· SERNA
I. IN TRODUC TION
The materials that constitute the core of
this study have been worked on through several
years. Originally Professors J. de Porta and M.
Julivert suggested I undertake the summariza­
tion, according to lithologic units, of the
faunas recorded in any way from the Creta·
ceous of Colombia.
At the end of the original investigation it
was apparent that if progress was to be made in
the biostratigraphy of the Cretaceous in Colom·
bia, it had to be started {rom the beginning,
that is to say through the detailed geologic
mapping of the key areas, the building up of co­
lumnar sections, the collecting bed by bed of
the fossiliferous units, the monographic treat­
ment of the faunas recovered and the proposal
of zonal schemes based on the information ob­
tained during the field work. I have attempted
to do that and I hope that I have not contribu­
ted additional complications to the task.
It is obvious that a work of this nature
necessarily had to pass through several pauses
and obstacles, however cooperation was always
found, willingly and uncompromisingly from
the people in charge of the geological institu­
tions of Colombia.
I have collected fossils largely from two
r�ions where I have mapped in detail mySirf,
or have had the valuable cooperation of other
field geologists; the areas investigated in the stra­
tigraphical researches have been more extensive
than those studied in detail. Two regions were
selected as initial key areas, both are classical in
the geological literature of Colombia: .Jijljp_.de
L_!fY.Jl (Department Boyaca) and /wJJ]o (former­
ly called Rafael Reyes; Department Cundina­
marca). I have limited my study to beds and
fossils in the interval Upper Aptian to Lower
Santonian; two reasons have influenced my de­
cision: many classical fossils in the paleontolo­
gical literature of Colombia presumably had co­
me from beds of these ages, and second the
opening of new roads offered the possil-,ility of
exploring fresh, undisturbed outcrops. As a rule,
fossils not found in situ, or impressions only
were not collected in the early phase of the
study.
Two hypotheses arrived at as a result of
this study I think worth emphasizing: A) It has
been common to find conere lions or lens-like
calcareous ·bodies paved with ammonites, fish
remains, bivalves and gastropods. This has occur­
red in different types of host rock: clay-shale,
graded sandstones, etc. (Rod and Mayne, 1954,
p. _207; f:JJO'O·Serna, 1968). _It is suggested in
th1s repdrt. _t6pt these "bod1es" represent the
result "ot� e'chanical 'concentration, by the
.
action of submarine currents on organic remains
with similar hydrodynamic behavior; in other
words these "bodies" may represent "micro­
channels" and their fossiliferous content may
include both morphologically different or mor­
phologically similar groups that by a simple
statistical analysis could be treated as a single
taxon. I surmise also cephalopods had definite
niches in their communities. B) The distribu­
tion of cephalopods may be considered mainly
{rom two different points of view: a) necro­
planktonic, 2) planktonic; the first approach
does not have any bearing on the origin and
development of new groups or the establish­
ment of new communities of cephalopods in
areas newly invaded by the sea.
A biological reference point to keep in
mind is that most living mollusks attain their
distribution largely by the possession of at least
one life stage capable of active dispersal or to
passive transportation. The latter type of
dispersal is usually accomplished during the
larval stage, and the larval stage resulting from
development from eggs poor in yolk usually has
a long planktonic stage (Thorson, Scheltema,
Ockelman). This seems valid not only for
mollusks with a larval stage but for recent
cephalopods with a direct development (Vokes
in Lane, 1960, p. 127).
Druschitz and Khiami (1970, p. 36) have
called this early ontogenic stage in fossil cepha­
lopods am monite lla. I agree with their idea that
probably the)' did not behave differently from
thir modern relatives in mode of reproduction.
Recent N autilu s hatches from large eggs sacs.
(Willey, 1896, 1902) and its intraegg develop­
ment lasts long enough to produce large ''n au­
t iloide lla (?)" [see Stenzel, 1964, K82], but as
noted by Willey (1896, p. 222) from observing
the protoconchs of ammonites, could not some
of them have hatched from eggs poor in yolk
with a corresponding ability {or long lasting
planktonic transportation?.
Some groups of ammonoids could have
been distributed by currents in their am mon i­
te lla stage. In consequence endemic morpholo­
gic types of cephalopods could be associated
with those "transoceanic travelers", with the
latter making intercontinental correlation
feasible, while the former groups may be useful
for establishing local biostratigraphical sequences.
In consequence I do not find it necessary or even
practical to invoke a ubiquitous nature {or
many of ammonite genera. Convergent iterative
groups have repeatedly been shown among
ammonoids. Consequently I feel that the
necroplanktonic distribution of fossil cephalo·
pods has its special role in intrabasinal distribu­
tion of fossil cephalopods but the life dispersal
o. them occurs during the arnmonitella stage.
(
-�
· All the material studied in this report will
· be housed as the basic type collection of the Ins·
titulo de Ciencias Naturales Museo de Historia
Natural of the Uhiversidad Nacional de Colom­
bia, Bogota (U.N.C.M.H.N.C.P. No.).
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
ll.ACKNOWLEDGMENTS
Many thanks to Professors J. de Porta an d
M. Julivert for their valuable suggestion an d al·
ways encouraging example. I also want to thank
Professor Robert Hoffstetter who called to my
attention the importanc..of attending a foteign
Institution for advanced studies; to Dr. Jesus A.
Bueno [former Thechnical Director of the Mi·
nisterio de Minas y Petroleos of Colombia), to
Dr. Andres Jimeno rformer Director of the
Geological Survery o{ Colombia 1 who gave me
their support from the beginning of this investi­
gation; to Dr. Ramiro Lobo [Chief of the Con­
servation Branch of the Ministerio de Minas y
Petroleos of Colombia 1 for his cooperation in
obtaining a loan for my graduate studies at
Berkeley.
I have to thank the Universidad Nacional
de Colombia, the Institute de Especializaci6n
Tecnica en el Extranjero (ICETEX) and the
Scholarship Division of the Organization of the
American States (OEA) for their economic sup·
port to complete my researches. Financial sup·
port for the plates included herein was provided
by the Fondo Colombiano de Investigaciones
Cientfficas y Proyectos Especiales "Francisco
Jose de Caldas" (COLCIENCIAS).
I have to acknowledge Professors W.B.N.
Berry, D.A. Savage, J. T. Gregory, C.M. Gilbert,
for their willingness to be member of my com·
mittee; thanks are also due to Dr. Ralph W.
Imlay and again to Professor Charles M. Gilbert
for their willingness to serve on my thesis .com­
mittee.
Many friends have shared with me their
ideas and also their skill in field work, of them
G. Renzoni, D. Barrero and C. Caceres-Giron
should not be blamed for my errors. Several
friendly field aides, Messrs. C. Giraldo, H.
Peiia, A. Aleman, J. Higuera, J. Duarte, had to
bear with my stubborn insistency to work
under changing weather conditions or had to
carry the heavy load of unprepared fossils. L. F.
Peii.a, and J. Garzon were very useful in hel­
ping with the preparation of the fossils, I
thank them also.
3
The following institutions are acknow­
ledged for their gracious loan of specimens for
comparison: Museum of Paleontology, Univer­
sity of California, Berkeley [Mr. J. Peck, Jr.],
Museo Geologico, lnstituto Nacional de Inves­
tigaciones Geologico Mineras, Bogota, Colom­
bia rProf. L. F. Rincon and former Director Dr.
A. Alvarez]; the University of Michigan, the
Texas Christian University and the Academy of
Natural Sciences of Philadelphia; the British Mu·
seum (Natural History). Dr. J. Helms of the
Humboldt • Universitat in Berlin, D DR;� provi·
ded me with excellent photographs of H. Kars­
ten's original material.
I have to acknowledge two old classmates
of mine, deceased Geologist Pedro Carvajal,and
Geol. Dip!. Ruben D. Llimis, who unafraid
backed with their wealth my loan from
ICETEX; Mrs. Amparo Prada was most helpful
in my dealing with the OEA.
Many thanks to Mrs. Gloria C. de Campos
and Mrs. Ines E. de Torres, who did the compo·
sing operation of the final version of the
manuscript. I thank Mr. Carlos Ruiz for his
assistance in improving the quality of the
text-figures.
The author could not have published this
paper had it not been for the compelling inte­
rest of Dr. Alberto Sarmiento-Alarcon, and
especially Dr. Michel Hermelin, Director
Institute Nacional de Investigaciones Geologico
Mineras (INGEOli;JINAS).
I have a deep debt to Professor J. W. Dur­
ham for his highly valuable scientific advise, his
thoughtful interest and his thoroughly under·
standing of my purposes during my studies at
Berkely.
In no way I am holding anybody except
myself, for any heresy I may have fallen into.
I hope this study, as simple as I see it, to
be acceptable to my family, my teachers, my
friends and to those who preceded me in the
study of the Cretaceous of Colombia. If this
work is found to be useful to my own country,
then I will be really pleased.
4 F.
PHYSICAL STRATIGRAPHY
In the present report only those units
dealt with or that have yielded fossils conside·
red in the investigation will be discussed. A mo·
re extensive discussion will be found in Etayo
Serna (1968a, 1968b), Julivert (1968) and Ca·
ceres and Etayo-Serna (1969). The location of.
the two areas studied in detail is shown on Text
Fig. 1.
,.....__
--:-\.YTHE C�F;_TA_��OUS SECTION_!:ROM
VILLA DE LEIVA TO LOMA PIEDRA
. GORDA. LARGELY_FII,OM �.'J'AYQ
SERNA, 1972, SEE TEXT FIG. 2A
PAJA FORMATION
(LoweFHauieriViafi"-Upper-Aptian pars)
The fonnation is typically exposed on the
north bank of Quebrada La Paja where it en­
ters the Rio Sogamoso (Middle Magdalena Va­
lley), and was nrst described by 0. c. Wheeler
(Morales et al., 1958). The Paja Fonnation crops
out extensively at Villa de Leiva and surroun­
ding areas. In addition, it is exposed in a broad
band east of the Arcabuco Anticline. Morpho­
logically it forms hills, some of which are being
mined for gypsum. Essentially, the Paja is a thick
sequence of black to variegated shales with abun­
dant calcareous concretions. Frequent intercala­
tions of gypsum and thin beds of argillaceous
limestone are also present.
The formation can be divided in three
units: (a) a !���erp��E _of bla�k clay sh �e which
weather to a reodJSli color, w1th sandy mterbeds
which represent the feather edge of a thick sand­
stone body observed at this S8llle horizon to the
northeast of Villa de Leiva; (b) 8J!1J�4_1e_�<J..n e
of light colored variegated claystones, which in
its lower part, contains numerous thin beds of
sandy clay and in its upper portion, intercala­
qons of calcareous claystone and gypsum; and
(c)\ Ill! U.P,P_e.r_J..nFJ:'.'al consisting of claystone
With liOllow noiiules, some of them acquiring a
brick red hue upon weathering and others being
yellowish and whitish on fresh exposure. Only
units (b) and (c) will be discussed further in this
report. 'The lower unit attains a maximun thick·
1
ness of 380 m (1246 feet), the intermediate is
480 meters (1514 feet) thick and the upper in·
terval about 80 meters 262 feet). The total of
940 meters (3083 feet represents the maYJ·
mum known thickness o the formation; it thins
northward. At its lower contact the Paja Forma­
tion is conformable with the underlying Ritoque
formatioQ . .4IJ.d the upper contact with the San
Gil Group 4iS, gradational. Some workers place
unconformities within the Paja or at its upper
contact but studies of the present writer do not
corroborate this interpretation. The Paja is hig-
hly fossiliferous but only the ammonites from
the upper part are described in this study.
From the middle variegated claystones
unit the fauna consistis of:
ETA YO· SERNA
Toxoceratoides (Colomboceratoides) ren­
zoni n. sp.
Valdedorsella (?) colombiana n. sp.
Pseudohaploceras leivaense n. sp.
Pseudosaynella ralphimlayi n. sp.
Cheloniceras (Epicheloniceras) dognael­
virae n. sp.
Cheloniceras (Epicheloniceras) camachoi
n. sp.
Vectisites (Zambranoites) mateusi n. sp.
Acanthohoplites bigoureti{orme n. sp.
Dufrenoyia sanctorum Biirgl
Stoyanowiceras treftryanus (Karsten)
From the claystone unit with hollow no-
dules, although it is fossiliferous, no ammonites
were described [There is a misprint in Etayo
Serna, 1972, p. 43; the fossils referred to unit
(c) belong to the upper part of the unit {b)).
SAN GIL GROUP
(Upper-·Kptililf pars�Uppef'A.lbian pars)
llub.aclt£1,953) named the San Gil Group
from expbSifffi"f"'artne type locality near the
village of San Gil, Department of Santander.
From its type locality the unit can be traced in·
to the vicinity of Villa de Leiva where it is a pro·
minent ridge former) as for example in the area
south and east of Sachica. The San Gil Group is
divided into two units of formational rank. 'I'he
lower San Gil Formation consists of a lowermost
oodfofVanegated, sandy claystones with abun­
dant, small clayey nodules and sandstone inter­
beds which are locally rich in echinoid& and gas·
tropods. Above this is a series of gray, dense li­
mestones witfi-·abi.mdant bivalves and occasio·
nal ·ammonites and micaceous claystones or
siltstones. Toward the upper part of the forma­
tion there is a thick (25 meters or 82 feet) unit
of gray, thickly bedded, fined grained-quartzose
sands�one containing occasional invertebrate
fossil fragments. The uppermost unit of the for·
mation consists of 10 meters (33 feet) of dark­
gray, very micaceous, fine to medium grained
sandstones.
The Upper San Gil Formation is predo­
minantJy dark-gray shale with a few intercala­
3
tions of sandstone or limestone near the base of
the formation. The Lower San Gil measures 480
meters (1574 feet and the Upper San Gil 620
meters {2034 feet , giving a total thickness for
the group of 110 meters (3608 feet). The lo·
wer contact is gradational; the upper contact
with the overlying Churuvita Group is confor·
mable. No unconformities have been observed
within the San Gil Group.
The following fossils from the Upper San
Gil Formation have been studied:
•
Knemiceras semicostatum Sommenneier
Engonoceras (?) duartei n. sp.
Platiknemieeras sp. ind. B.
Tegoceras benavider;caceresi n. sp.
Carloscaceresiceras (?) chimuense(B.C.)
 t: A
s
•

A) VILLA DE LEIVA
-

.0<1.:::'.�
.. /
B) APULO
- i .
- i I
so 'G 100
!
..f',.; I
Anrqzoll
as ·

FIGURE l. MAP OF COLOMBIA SHOWING LOCATION OF SECTIONS INVESTIGATED IN DETAIL

FIGLH 1\ 2 FIGLRE 2

COMPOSITE COLUMNAR SECTIONS

2AJ Vl:LLA DE LEIVA 2B) APULO

(Rafael Reyes)

�
'-"-' -'-"-'-'='1

·
: ·
· < ·:-
.:
. -: :- . -
· ·;: ·
-:

l<: �-·...� f\'-...l c.1 r..n

[..1.t,.v"'�rt· ....:
····'-''•
l•lt ( 'l;n,. (>\ ., '"' "'1{t01''J. rt:;"
'7t�!).�,��. (/1-'"
tA.nRo 'C-Ot'-" rL' • c, ... \4-� \Ji �-::- \ �.
-:i �(J)'�
·;;

t
0

CD " 0
"-
"'

f � L., \, J '}� �

i'Jrf -.t :.;-1:""•

u.

s. 1:20;000 s. 1:10.000
 ZONATION OF THE CRETACEOUS
CHURUVITA GROUP
(Upper Albian ? pars-Turonia!" pars?)
The name "Churuvita Group" was first
proposed by Etayo Serna (1968b), with the ty­
pe locality a section on Quebrada Churuvita (a
tributary of Rio Samaci) along the Sachica­
Puente Samac& highway; According to Etayo
Serna, the Churuvita Group is comprised of the
Churuvita Fonnation and the overlying San Ra­
fael Fonnation.
The Churuvita Fonnation was originally
described by Etayo Serna (1968a) who selected
as the type locality the exposures on Quebrada
Churuvita along the Sachica-Puente Samaca
road. In the area of interest, the Churuvita For­
mation forma the highest ridge on the west
flank of the Arcabuco anticlinorium; it is also
exposed on the limbs of the Churuvita syncline
along the Sachica-Puente Samaca highway. A
tripartite division of the formation is observed;
(a) a basal unit consisting essentially of gray,
thick-bedded, quartzose sandstones with thin
interbeds of claystone and siltstone; (b) a mid·
dle interval of lenticular limestone beds, some­
times argillaceous, rich in Exogyra, which alter­
nate with claystone and siltstone as well as oc­
casional quartzose sandstones; and (c) an upper
unit comprised of massive beds of fine to me­
dium grained sandstones and equally massive
limestones interbedded with claystones and silt­
stones. In general the formation changes facies
toward the west where it becomes more arena·
ceous. The three subdivisions of the formation,
from bottom to top, measure 105 meters (244
feet), 75 meters (246 feet) and 225 meters (738
feet), respectively, giving a total of 405 meters
(1328 feet). The fonnation thins toward the
north. Contact with the San Gil Group is con·
formable at some localities and transitional at
others whereas the relationship with the overl·
ying San Rafael is everywhere conformable.
Fauna collected from the Churuvita Formation
consists for the most part of bivalve mollusks;
they are not described in the present report;
after Etayo Serna (1968, p.41) the mollusks are
the following:
Exogyra boussingaultii Dietrich (non
D'Orbigny)
Plicatulopecten concentricus Brtiggen
Ostrea syphax Coquand?
Pecten tenouklensis Coquand
Exogyra squamata d'Orbigny
Paleontological data from the underlying
and overlying strata suggest that the unit may
be assigned to the Albian pars (?)·Cemomanian.
The report of Venezoliceras (?), previously
(cp.cit)tentatively assigned to the base of the
unit iS now considered unreliable information.
SAN RAFAEL FORMATION
· (Lower Turonian)
Etayo Serna (1968a) first proposed the
OF CENTRAL COLOMBIA BY AMMONITES 7
fonnation. As type locality he chose the que·
brada whose source is 200 meters south of Chir­
cal San Rafael. The San Rafael Formation crops
out to the east of the Serrania de Arcabuco; it
is also exposed along the axis of the Churuvita
syncline. At its type locality, the formation
may readily be divided into: (a) a lower unit of
gray shales which weather to lighter tones with
minor beds of thinly interbedded limestones;
and (b) an upper unit consisting of thinly bed­
ded, siliceous siltstones with several narrow
bands of chert and toward the top, a single sha·
le bed. The lower unit of the San Rafael is 60
meters (197 feet) thick at the type section, the
upper unit is 14 meters (46 feet), giving a total
fonnational thickness of 74 meters (243 feet).
Whereas the lower unit is variable, the upper
one is of remarkably uniform thickness. From
the lower unit of the San Rafael Formation
near the axis of the Churuvita syncline along
the Sachica·Puen te·Samaca highway, several ce·
phalopods have been collected; Paramammites
colombianus n. sp., is described in the present
report.
CONEJO FORMATION
(Upper Turonian ? -Lower Senonian pars)
The Conejo Formation was first named
by Renzoni (1967; published in Etayo Serna
1968a). Its type locality is the Alto del Conejo
located on the Oicata.Chivata road, some 9 ki­
lometers (5.6 miles) due northeast of Tunja.
In the study area, the Conejo Formation crops
out on the folds forming the margins of the Sa·
maca-Sora Valley. The Formation is comprised
of three units: (a) an unnamed lower part con·
sisting predominantly of black shales with occa·
sional thin interbeds of fine·grained sandstones
or siltstones; (b) the middle, Cucaita Member
which is exposed in the core of the Cucaita an·
ticline. Its lower part is a series of calcareous
shales alternating with several sandy limestones
beds, the shales containing abundant calcareous
concretions. The member as a whole takes on a
whitish-yellow color due to the claystones to­
ward the upper part which become progressively
more arenaceous upward; and (c) an upper in·
t.;rval, also unnamed, composed of yellow·whi­
te, quartzose sandstones intercalated with dark·
gray, usually sandy shales. Near the top of the
interval are found highly fossiliferous lenses of
limestone. The total thickness of the Conejo
Formation is 370 meters (1214 feet), divided as
follows: lower interval, 80 meters (262 feet);
Cucaita Member, 170 meters (558 feet); upper
interval, about 120 meters (394 feet). In the
area of this study, the contact with both the si·
liccous siltstones of the San Rafael Formation
and the overlying Plaeners Formation are con·
fom1able. The middle, or Cucaita Member, has
yielded the following assemblage described in
the systematic part of this report.
Nice{oroceras boyacaense n. sp.
Peroniceras robertogarciae n. sp.
Peroniceras diabloi n. sp.
Peroniceras guerrai n. sp.
Prionocycloceras (?) cucaitaense n. sp.
Gloriaceras correai n. sp.
8 F. ETA YO· SERNA
Gloriaceras paulinae n. sp.
Zumpangoceras ospinai n. sp.
Zumpangoceras (?) sorae n. sp.
Buenoceras loboi n. sp.
Protexanites cucaitaense n. sp.
The uppermost part of the upper interval
contains fossils of early Santonian age, includ­
ing Lenticeras baltai Lisson.
PLAENERS FO RMATION
(Santonian pars)
According to Julivert (1968), the Plaeners
Formation was frrst differentiated by Hubach
(1931). The type section is located on the Sa­
bana de Bogota, at "Boqueron de Tabio". The
Plaeners can be traced from its type locality in·
to the study area. It forms the prominent spur
observed to the east of the Samaca-Sora Valley.
The unit consists of a series of gray, siliceous
siltstones in beds of 5-20 centimeters (2-8 inches)
exhibiting rhomboidal fracture and rich in fora­
mlnefera. The siltstones are frequently interca­
lated with beds and lenses of dark-gray chert
and occasional layers of phosphatic sandstone.
In the Quebrada San Joaquin, 50 meters {164
feet) are measured. The contact with the
underlying Conejo Formation is conformable
and with the overlying unit, gradational.
GUADALUPE FORMATION
Julivert (1968) states that Hubach in
1931 named {or perhaps more accurately, re­
defined) this unit from exposures in Cerros
Guadalupe and Monserrate,immediately east of
Bogota. The sequence forms the uppermost
·p.rominent cliffs to the east of the Samaca-Sora
valle y and represents the northern continuation
of the sandstone of the same name which is ex­
posed on the mountains surrounding the Saba·
na de Bogota. The lower part consists of beds
of siliceous argillites or siltstones, some of
which tend to fracture rhomboidally. The over­
lyingTierna Sandstone is a white, yellow-weathe­
ring, coarse grained arenite which exhibits pro·
mlnent cross-bedding toward its base. In the
studied area the argillites reach a thickness of
50 meters (164 feet) and the overlying sands·
tone 9 meters (30 feet). Contact of the lower
part with the Plaeners is gradational, and con­
tact of the Tierna Sandstone with the Guaduas
Formation is conformable.
WHE CRETACEOUS
VlOTA,
SECTION IN THE APULQ,
REGION,
- LAS MESITAS
SEE TEXT·FJG. 2B.
L new road from Anapoima to Apulo, beginning
A NAVETA FORMATION
·
�·�-?�:·.-.. '""�- ·r·
.
., ;.
The'
. •- .
.
name was originally introduced by
Hubach (1931, p.44) to describe a sandstone
horizon; it has been granted formational rank­
ing by Caceres and Etayo-Sema (1965, p.5). Its
name comes from the cement factory of that
name. The unit crops out as the core of the
Diamante anticline [Cerros del Diarnante to the
north, and Cerro del Trueno to the south 1. The
unit is well exposed on the road from La "Nave­
ta Cement factory to Ingenio Central San Anto·
nio (now Escuela Vocacional), just beyond the
bridge over the Bogota River, or on the west
limb of the El Diamante anticline where the ty­
pe section was measured. The La Naveta For­
mation consists of a sequence of white to yello­
wish quartzitic sandstones, medium to coarse
grained with frequent grains of chert; not very
compact, with interbeds of shale rich in plant
remains. The La Naveta Formation includes
three units of member rank.(!) pi_�����-�m.;
... ber; its name comes from the Ctirros D1amante;
it consists of coralline limestones interbedded
with shales and sandstones, the upper beds are
19 meters (63 feet) thick; it has been mined for
use in cement. A specimen of Thurmanniceras
sp. was found loose near the outcrop of the
unit on the east flank of the El Diamante Anti­
cline, north bank of Rio Bogota. (2) 'Th.� J?.�j�;
cal member; its name comes from Cerro de Be­
jiicanvnere it has been mined.lt is a biostrome
of bivalves, worm tubes and a few corals. It
yielded in situ: Heinzia (Carstenia) sp. The total
thickness measured without having reached the
base of the La Naveta Formation is 243 meters
(801 feet).
VILLETA GROUP
This unit was established by Hettner {1892),
it has been thoroughly reviewed by Julivert
(1968); no type locality has yet been formally
proposed. The name was introduced in this re­
gion by Hubach (1931, plate 1). Several units
with formational rank have been proposed wi­
thin the group; {1) the basal unit or Trincheras
Formation named" after its type locaiitfat Que­
brada Trlncheras. It consists of an alternation
of limestone beds very rich in bivalves; its age is
believed to be partly Barremian to Lower Ap·
�J!:!l. The lower and upper contacts are confor·
mabie at the type locality. The formation crops
out forming a band around the core of the El
Diamante anticline; towards the northeast the
formation changes into shales. The thickness
measured at the type locality is 180 meters
(594 feet). (2) The. Sos2!Lf�l1Tlatiqn. The·
.name was first used by Caceres and"Etayo-Sema
(1969, p.5); it is derived from Quebrada Soco­
ta. This unit is intended to embrace the litho·
logic sequence from the frrst graded quartzitic
and calcareous sandstone at the base of the
Socota Member (to be described next) which is
-here taken as the basal member, and extending
up to the first occurrence of porcellanite beds
here considered the basal element of the
overlying unit,_jhe_lliliL.Eormation. The type
locality of the So"CcifaF"OrfiiatiO'if'tinilong the
at km 3 + 320. Except for its basal member the
:socota Formation consists of a succession Of
shales, they are superficially variegated but dark
gray at depth; there are frequent calcareous
concretions and··:gypsum veinlets usually cros­
sing the bedding of the shales. The maximum
thickness of the unit is estimated as 150 meters
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
(495 (�et). Two members are described in de-'
tail: {iij th�Q�OTfiA �-rl�r�IT'J.'�l�.-�,E,;.f,_:�--.... .
TION measureu aoou .., me rs �" teeL1 nor�ou
of culvert No. 19 [Km 3+220) on the new road
Anapoima·Apulo.
XI Argillaceous limestone bed
or 0.49 feet.
X Variegated calcareous shales with reniform
calcareous concretions, or lens-like argill a­
ceous limestones. At base of this segment,
"fP sample C 199 yielded l
• ! lw.clites obli·
- quum (Riedel) . . . . 8 m or 26 feet.
. • •
IX Calcareous shales with lenses of argillace·
pus limestones; crushed ammonites,bival·
ves; altop sample C 200 yielded Riedeli· • � yielded Ri�delite!i_ esthersemae n. sp
"'-' tes obliquum (RiedeQ . . 1.8 m or 5�[eet�
VTII Argillaceous limestone lens . . . . . 1 m or
3.3 feet.
vn variegated shales, with intercalations of
thin layers of very fine grained calcareous
and quartzitic sandstones, with bands rich
in fragments of cephalopods and bivalves
. . . . . . . . . . . . . . . . . 8 m or 26 feet.
VI TOP Quartzitic calcareous sandstones,
very finely to finely grained in parallel la­
yers of 0.001 m (or 0.003 feet); calcareous
concretions are locally present, they con­
tain crushed ammonites and bivalves . . . .
1 1.50 m or 38 feet.
V Quartlitic calcareous sandstones, fine to
very fine grained, laterally tending to
appear massive, but regularly occurring in
parallel or wavy, fine to very fine layers
of 0.001 m (0.003 feet) thickness, locally
with calcareous concretions surrounded
by fragments of cephalopods or bivalves
or coarse quartzitic sand 15.50 m or
41 feet.
[V Light gray quartzitic, fine grained sands·
tone, massive, 1 m (3.3 feet? thickness,
with interspersed calcareous concretions
. . 1 m or 3.3 feet.
. . • '-·Du_frenoyia sanctorum Btirgl.
UI Quartzitic siltstones with intercalations of
very fine grained sandstones; predominan·
tly parallel laminae of 0.001 m (0.003 feet)
thickness. Locally there are lenses with
rounded quartz grains up_ to 0.005 m dia·
meter and crushed ammonites and bival·
ves. Some lenses of limestone also occut
7 m or 23.5 feet.
II Calcareous quartzitic siltstones to very
fine grained sandstones with subparallel
to wavy lamination; calcareous concre tions
with fragments of cephalopods and spora­
dic rounded quartz grains up to 0.001 m
diameter . 7 .30 m or 24 feet.
• . . . . • . .
BASE
Dark gray shale . . • • • .
9
REFERENCE SECTION. SOCOTA MEM·
BER. The base of the measured section is
· near the culvert at K 4 + 06.
-.. ..
Variegated shales, partly covered.
V TOP. Dark gray, quartzitic sandstone,
• . . . . 0.15 m calcareous, very fine grained to fine grai­
ned, locally medium grained. Subparallel
. lamination, with calcareous concretions
which seem to merge in to the envelopping
surrounding rock; fossiliferous: at 1.4 m
(4.6 feet) from top, sample C-220 yielded :
' G_olombiceras sarmientoi n. sp.
r. Xcantliohoplites bigoureti(orme n. sp.
i Clieloniceras(Epicheloniceras) pardoi n. sp.
at-4--m (13.2 feet) from top, sample C-222
. • . . •
5.5 m or 18'"'feeC ;· ,�" -· < · -
IV Siltstones to fine grained quartzitic sands·
tones in layers of 0.01 to 0.15 m; subpa­
rallel to wavy lamination, with tendency
. to become more massive toward its, base;
calcareous concretions 4.9 m or 16 feet.
• .
ill Siltstones to very fine grained quartzitic,
calcareous sandstones; upwards it shows
parallel lamination, towards its central
part the lamination is subparallel to wavy,
towards its base it tends to become ma·
ssive , but local lamination may be obser·
ved; at its base there is a calcareous lens
with quartz grains up to 0.05 m diameter.
Calcareous concretions occur, some are
rather calcareous lenses which merge with .
the lamination, they enclose concentra·
tions of ammonoids and bivalves. From
the top of this segment, sample C-223x
yielded :
•
Toxoceratoides (Colomboceratoides) ren .
zoni n. sp.
. • . . ·· Melchiorites palmen' n. sp.
_ Aconeceras gutierrezpalmae n. sp.
-"" Cheloniceras (Epicheloniceras) carlosacos­
tai n. sp.
Pseudoptychoceras gi/berti n. sp.
"' Eodouuilleiceras (?) tequendamai n. sp.
At 3.8 m (11.5 feet) from the top of this
portion ot the sequence, sample C-223
yielded :
.._ Eogaudryceras helmsi n. sp.
..- A <;_o_neceras gu tierrezpalmae n. sp.
Pseudoptychoceras gilberti n. sp.
� Cheloniceras (E.) carlosacostai n. sp
. • ·
• . .
7.:r-ot'25.7 feet.
II Siltstone to very fine grained quartzitic
calcareous sandstone beds, with parallel
lamination and with calcareous concre·
tions; toward the middle part the lamina·
tion becomes subparallel to wavy; locally
the grain size increases to fine and even
becomes middle size especially toward
4.5 m or 15 feet. the base of this segment where lenses of
10
coarse grained sandstone occur and the
rock tend to become massive . 7.1 m or
23.4 feet.
I Alternation of calcareous lenses up to 0.5
m (1.65 feet) thickness, which enclose
abundant fragments of cephalopods and
bivalves and also has rounded grains of
quartz up to. 0.001 m diameter, in &;I ter­
nation w1th sdtstones beds or fine gramed
quartzitic sandstones • 4.15 m or 13.7
• •
feet.
The Socota Member is a particularly inte­
resting unit; it is missing toward the southwes·
tern part of the region but it increases notably
in thickness and areal distribution northeast­
wards.
amr�tWrdJf�:·d�Alf?r��o;��rt!f·�eme��
SECTION of its basal beds was asured
adjacent to the house of Hacienda Capotes. This
name was introduced by Caceres and Etayo­
Sema (1969, p .5), without description of type
section. This unit can be recognized by the
frequent occurrence of argillaceous, thinly
bedded limestones or the abundance of highly
fossiliferous calcareous concretions up to 0 .5 m
in diameter distributed in layers; the top of this
unit is marked by the dominance of clay-shale
with frequent occurrence of small argillaceous
to slightly calcareous concretions.
Section of the basal beds of the Capotes
Member measured along the road from the casa
of Hacienda Capotes to Japon, it begins inme­
diately adjacent to the house.
VIII Argillaceous limestone • • . 0 . 5 m or 1.65
feet.
VII Variegated shales with calcareous concre­
tions; just above the underlying limestone,
sample C-255 yielded :
j Lyel/iceras pseudolyelliforme n. sp.
i Lyelliceras isaacleai n. sp .
..t:. Hypophylloceras wiedmanni n. sp
f m or 8.25 fe et
.
VI Argillaceous limestone thinning and wi·
dening irregularly; sample C-255 x yielded :
"! Neodehayesites columbianus (Riedel)
0.5 m or 1.65 feet.
V Variegated shales with interspersed calca­
reous concretions; argillaceous limestones
separated by variegated shales, with cru­
shed ammonites . . . . 1.8 m or 5.9 feet.
IV Variegated shales with lenses of limestone
up to 0.5 m (1.65 feet) thickness
2.7 m or 8.9 feet.
Ill Argillaceous limestone that tends to split
in thin lay ers . . . . . . 0.8 m or 2.6 feet.
II Variegated shales . . . 2.5 m or 8.2 feet.
F. ETA Y O - SERNA
I Argillaceous limestone . 0.8 m or 2.6 feet.
REFERENCE SECTlON of the bottom
of the Capotes Member, ·measured . aetnebegiil­
ning of trail to the power transmission tower,
just west of village of Tarapaca.
II Dark gray shale; three intercalated layers
of fossiliferous concretions. Sample C-12
yielded :
Neodeshayesites co/umb ianus (Riedel). . .
. . . . . . . . 1.2 m or 4 feet.
I Dark gray shales with very fine argilla·
ceous limestone beds and calcareous con­
cretions with crushed ammonites. 25 m
(82.5 feet) below sample C-12, sample
C-13 yielded :
Trochleiceras juliverti n. sp.
Douvilleiceras tarapacaense n. sp.
Carloscaceresiceras caceresi n. sp.
BASE . . . . . . . . . . . 60 m or 198 feet.
Hard, red-brick, Trigonia bearing limes­
tone (Trincheras Forrnation). At this
geographical point the Socota Member
is absent.
R EFERENCE _S ECTI O N for basal part of
the Capotes Member and 'underlying units; this
section was measured at Cerro Bejucal, from
the machinery house of the limestone mine,
and extending SE along the abandoned road.
VIII Gray shales, sporadic calcareous concre·
tions . . . . . . . . . . . 50 m or 165 feet.
VII Variegated shales with gypsum veinlets,
calcareous concretions and thi n fossilife­
rous limestones. At top of these shales,
sample C-169 yielded:
�- Rinconiceras rinconi n. sp.
_..,Pla tikne miceras colombiana n. sp .
.,. Carloscaceresiceras sp . . 30 m or 99 feet.
VI Variegated shales with gypsum veinlets,
. • . 2.5 calcareous concretions and thin fossilife­
rous limestones. At top, sample C-170
yielded:
'l'
Eodou villeiceras pedrocarvajali n . sp.
,.. Neodeshayesites columbianus (Riedel)
. • •
_ J)·o chleicerasjuliverti n.sp. 12 m or 40 feet.
V Variegated shales with frequent calcareous
concretions and lenses of fossiliferous li-
mestones . . . . . . . . . . . 9 m or 30 feet.
IV Variegated shales with gypsum veinlets
and locally abundant gastropods and bi­
valves bearing concretions . . . . 27 m or
. . • . . .
89 feet.
• .
III Dark gray shales . . . . 200 m or 660 feet.
II ( Socotii"'·Member). Quartzitic graded bed­
ded sandstones; large blocks of limes­
- tone . . .Du(renoyio codazziano (Karsten)
. . . . . . . . . 10 m or 33 feet.
. . . . . . .· .
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES 11

I (Trincheras Formation). Limestone in thin Ill Dark gray, siliceous silt or mud shale, it
beds . • . . • • • . . • . • . • . . • . • . • • . • • • • • breaks in small slabs. Ammonite impres­
sions are frequent; there are interspersed
-· uu,.Q �SU�
u
ISlti .This formational
name was mtro ce�n this region by Hubach
calcareous concretions up to 0.6 m In dia­
meter . . . 14.3 m or 4 7.0 feet.
. . . . • • •

(1931); its type locality is at Hilo on the road

to Cambao. Lithologically it is characterized by n· Dark gray, mud to clay shale, weakly to
an altern�tion of liditet"or porcellanite& with moderately laminated, with elliptical cal·
dark gray clay-shale . In the present report its careous concretions up to 0.5 m in diame­
base is taken at the rmt occurrence of liditic or ter . . . • • 7.7 m or 25.4 feet.
. . . . . . • •

porcellanitic beds. A good reference locality'ls

on the old road Anapoima-Apulo before entering I Dark gray, clay-shale, with petroleum odor,
the Apulo alluvial terrace. The morphologic ex­ there are interspersed limestone concre­
pression of this unit is usually steep ciifs or high tions or lenticular beds up to 0.5 m; sam­
peaks with rounded tops; the presence of brick ple C-300 yielded:
like siliceous fragments allows its presence to be
recognized in the field. Although i!n.P�-�i�� Mammites nodosoidesappelatus n. sp.
o f..a.I;Dln. P,nites are foundJr:t these rocks they were Mammites scutulatus n. sp
not studied l_�'"the present Investigation.
.

Hoplitoides hemanmojicae n. sp.

Hoplitoides {?) sp ind.
Franciscoites suarezi n. sp. ·
.

UNNAMED UNIT. Above the previous

unit there is a succession of shales, not forma· Fagesia zanellai n. sp.
lly named , they did not yield any fossils during Imlayiceras {?) ralphimlayi n. sp . • • • • • •

the field studies. 9.01 m or 30.0 feet.

�..X,B..Q��,.F,O.RMA'IlQN .This for­ .

.... R ��J;:R.E!.NG��-��CIIQ\':Ult.J.a .. frg!)._��a
mational nairie was introduced in this region by Formation� 'Iriea8tired· at H acie nda La Giraldii,
Caceres and Etayo- Serna (1969, p. S); it consists at place where the formation forms the ridge
of an alternation of thin limestones and dark alongside the Bo go ta Las Mesitas road, Km 27+
-

shales towards its base but towards the top it 832.

conaists ofporcellanites or finely stratified cherts;
the succession at several reference localities fo· Ill Dark shales, partly covered . • . • • . . • • •

llows.

REFERENCE SECTION, La Frontera For­ II TOP. Fine layers of siliceous shales in al­
matiori, road Ctimaca:vtOli;at La Cabana. K 23 . ternation with porcellanites, in layers do­
minantly 0.1 m (0.33 feet), with intercala·
XII Dark gray, siliceous shale, finely lamina· tions of limestone beds up to 0.5 m (1.6
ted . . • • • 1.4 m or 4.62 feet.
. . . . . . . . feet) . . 12.5 m or 41 feet.

XI Alternation of porcellanite& in layers up I Fin ely laminated dark gray shales with in·
to 0.15 m (0.49 feet) thick with gray to terspersed lenses of limestone up to 0.5 m
maroon mud shale 4 m or 13.2 feet. or 1.6 feet thickness, very compact, pyri·
tized, some of them are fossiliferous; sam­
. . . •

X Gray shales. • . . . . . . 1.5 m or 4.9 feet. ple C-264 collected 18 meters (59.40 feet)
below the top of this interval yielded :
IX Creamy p orcellanites, sticky when wet,
with impressions of ammonoids . . . . . • . Hoplitoides lagiraldae n. sp .
1:6 m or 6.3 feet.
·

Sample C-263 coiiected 11.2 m ( 37 feet)

Vill Gray silt-shale very rich in compressed below the preceding sample yielded:
impressions of "Inoceramus ".
Mammites fugax n. sp
. • . . . . . .

2.3 m or 7.59 feet. . . 39 m or 129 feet.

Vll Maroon mud-shale and porcellanites • . . •

0.9 m or 3.0 feet. ROAD LEVEL

VI Creamy porcellanites with abundant im· The laminated clay shales overlying the
pressions of ammonites and bivalves, with Frontera Formation yielded sample C-134 :
some cherty concretions; the rock is stra·
tified in layers up to 0.25 m thick Prionocycloceras longispinatum (Basse).
Prionocycloceras portarum n . sp .
• • • • • •

5.1 m or 16.80 feet.

Codazziceras scheibei (Riede l)
V
.

Gray silt-shale with elliptical calcareous Codazziceras fina n. sp.

concretions 1,5 m or 4 .90 feet.
COMMENTS
• • • • . . •

IV Creamy porcellanite stratified in layers up

to 0.25 m thick, with mud shale intercala· With respect to the li thostratigraphic
tions; the porcellanite& tend to break in scheme utilized in the present report a further
small brick-like blocks.4.5 m or 14.8 feet. comment is necesary. Some of the fossils illus-
12 F. ETA YO · SERNA
trated in several of the early paleontological
works on Colombian Cretaceous: I.Lea (1841);
E. Forbes (1845) ; H. . Karsten.(1858) ; A. D'- Or�
bi!Wr(!B""�2)i L_. �jec:lel (19 38), �arne from th. e ·
Apulo , Vwta, Las Mesitas reg1on. The first
attempt to present a stratigraphic sequence of
the Cretaceous rocks of this region is that of E.
Hubach ( 1931), in a paper in which a geologic
map and a columnar section are presented ; fos­
sils were collected and preliminarily studied but
they were not described or illustrated and only
a brief account of them was later given by J!m.;__
istroffer .(t9�E?1 presumably based on the mate�
nil re'TerreCtO by Hubach . Hubach (Op. Cit.,
p .5 5 ) discussed the stratigraphic sequence
studied by him and the faunas with preliminary
identifications by C. Rhein listed in the text.
Of his lithologic units ("Horizons" in Hu­
bach's terminology) discussed , the m ost fre­
quently quoted has been that called "Horizonte
de Colombiceras (rY)" about which Hubach
wrote (Trad ., lb., p. 57) "It has been possible to
study carefully the 'Horizonte de Colombiceras'
only in its upper part. These levels of the upper
part h ave been divided in the level of Hoplites­
Acanthoceras (IVc), in the level of ammonitife­
rous concretions (lVb) and in the level of Co­
lombiceras (IVa). The highest of these levels
1
which form together about 50 to 70 meters ot
the upper part of the horizon, consists of a
bank of lime composed of ammonites of the
group Acanth oceras-Hoplites, that can not be
separated satisfactorily. Furthermore, ch&rac­
teristic species of Hoplites and of Desmoceras:
D. Hopkinsi F., D. Mayorianus '! are observed".
However and unfortunately the levels
were numbered upside down in the stratigra­
phic column on the left side of the geologic
map accompanying the paper, that is level (IVa)
appears as the highest and level (rYe) as the
l owest, but the expression "Nivel de Acantho­
ceras-Hoplites" remains attached to the highest
level.
Breistroffer (19 36, p. 1 5 6) �ave a list of
the fossits='ot-Hubach's levels thJS way : "Le
Gargasien correspond a !'horizon a Colombice­
ras (IV) de M. Hubach. Sa partie inferieure
(rYa) renferme, a Viota-Capote-Apulo, Du{re­
noya (?) nov. sp. (gr. Codazzklna Krst. sp., sub
Colombiceras Karsteni ap . Rhein). Sa partie
m oyenne (rYb) presente, a Cerro del Capote ,
Douuilleiceras solitae d'Orb.sp.( ::multinodosum
Rhein in litt.). Sa partie superieur (rv c) mon­
tre, i San-Antonio, Phylloceras cf. Buchwnum
Forl)�s sp. (sub Desmoceras Mayorianus ? ap.
Rhein), · Pseudohaploceras Hopkinsi Forbes sp. ·
et Du{renoya nou. sp. (interm. entre D. luren­
sis Kil sp. et D. texana Burckh .).Enfin, Ia locali·
te de Cerro del Capote (IVb-c) a encore donne
Brancoceras cf. aegoceratoides Steinm. (affini­
tes avec 1' Albien du Perou ). "
Here it is to be observed that there is a
precise correspondence between Hubach's enu­
meration of levels in the text and Breistroffer's
usage of the roman numeration.
Hubach (Op. Cit., p. 45-46) had written
"In the Viota region, ascending the Cerro del
Capote , it is found, on the upper part of the
horizon of Colombiceras, first a bank of limes­
tone 30 em thick, consisting exclusively of the
species of ammonite Colombiceras karsteni
Marcou . . . the h orizon of Hoplites -Acan­
thoceras is above that of the h orizon of Colom·
biceras. The m agnitude of the sediments that
separate both h orizons does not seem conside­
rable, at most about 50 meters."
Eviden tly ��l:!istr?ffer was �rong when he
used the expressiOn " I h orizon a Colombiceras
(IV) de M. Hubach . Sa partie inferieure (IVa)"
because level (IVa) of Hubach's terminology
represents the base of the upper part of the
Horizonte de Colombiceras; unfortunately Hu­
bach used both the words horizon and level
(lb., p . 45,46) for two different things.
�
. Sub�<!u �� l �iirgl ll.953, p . 13) wrote :
"M. lmiSttciffer \ 19'"3"6 , . . ), i n ve rted the
position of these horizons considering rvc,
after Hubach the lowest, as the highest." One
can not be sure if BUrgi trusted only the co·
lumnar section next to the geologic map on
Hubach's paper; h owever Burgi called the three
levels of Hubach (rYa, IVb, rYc) as "nivel de
Colombiceras" and proposed to include it
{1955, p. 13,14) as the basal level of the Albian
� Incidentally in a previous paper Etayo-Serna
mcorrectly wrote base of the Aptian instead of
Albian; 1964 , p. 20J. Accord in g to my studies
Btirgl was correct with respect to the age assig­
ned to the three levels but not in the taxonomic
treatment of the fossils.
Burgi (1955) also recognized good
upper Aptian fossils, according to his lists
(Ibid., p. 12) but he did not illustrate them; in
his paper on the Apulo region (=Rafael Reyes)
Biirgl did not include Aptian beds in his "Nivel
de Colombiceras", but later (1961, p. 1 04 , p.
1021 Fig. 8) he included under this term both
Aptian and lower Albian beds. Julivert first
observed (1968, p. 222) that Biirgl had conti­
nued talking about the Nivel de Colombiceras
when referring to a resistent sandy or calca­
reous horizon, which stands out prominen­
tly in the midst of the uniform lutitic succession
constituting the Villeta Group; Julivert was re­
ferring to the composite Fig. 22 (Biirgl, 1961 a)
and Fig. 8 (BUrgi, 1961c).
A furiJ.'ler point must be stressed here:
.
eastward from the region here discussed there is
a well known. se"cjiience of SaD.dstones orig!nally
named "Ar�nisca . �� Une" by H!!l2!!-£h (1931,
columnar section�orCngft't side of the . geologic
map , plate 1), this unit was subsequently subdi·
vided in three parts, the upper calle d the l.!.r!e
sand�ton�. and the lower one the Ubaque
sandstone , the unit in between was left u n'na�
med (Campbell and Biirgl, 1965, p. 578, Fig 3);
both named units were however considered
Albian in age. One of the preceding two tenns
was used bt' Burgi (1963, Fig.8, P· 720) to
replace his ' Nivel de Colombiceras' of Fig. 22
and 8 (1961a, 1961c) that are the same illustra·
 ZON ATION OF THE CRETACEOUS OF CEN TRAL COLOMBIA BY AMMONITES
tion a6 appeared on lJ. 720 (1963) and on
which the new term "Ubaque-Sandstein-Hypa­
chanthoplites Niveau" was Introduced. The
above discussed terminology is not followed in
1-�-B-I_osT AP H_Y___-[.
the present study.
_R_A_
.....
R_
__
T_W
The following subdivision of the litholo­
gic sequences investigated in this report, based
on their faunal content (mainly ammonites),
that is their zonation, attempts to be in agree·
ment with the recommendations of the Intema·
ttonal Subcommission on Stratigraphic Classifi­
cation: "An International Guide to Stratigra­
phic Classification, Tenninology, and Usage"
(1972). All the proposed zones in this scheme
are considered ••Assemblage-zones (Cenozones)".
The purpose-of estibliSJITng Oils zonation IS to- - ­
facilitate th e geological work currently being
done on the Cretaceous rocks of the Cordillera
Oriental Colombiana, and I am not attempting
to establish a rigid" Archetypical" scheme. I feel
it is important to have a reference element, an
objective starting point whose value it is hoped
will be challenged by others,
In other words, the subdivisions are
based on the succession of the paleontological
material collected in situ in the field and from
successive sequences of rocks as measured and
mapped at localities of easy access and unequi·
vocal identification. The lower boundary of the
units is established at the fll'St known occurren·
ce of new taxa, especially genera; the upper
limits of the unit are consequently determined
by the occurrence of the next new groups at
the base of the overlying unit. Correlation or
time equivalence with the zonal scheme esta­
blished in Europe is not attempted in greater
detail other than the mere comparison of the
common occurrence of certain groups of
ammonites considered significant enough to
permit usage of the stage terminology for the
Cretaceous rocks as first proposed by_ d'Orbigny
and Coquand.
THE F O LLOWING ASSEMBLAGE ZO·
NES ARE HERE PROPOSED FOR USE IN CO·
LOMBlA.
As far as these studies demonstrate, the
zonal species have their rtrst appearance along
with the fU"St record of Cheloniceras (Epichelo­
nicerCI$), Vectisites (Zambranoites), Riedelites,
Juandurhamiceras and GargasicerCI$, Cortezice­
nzs, Pseudoptychoceras, Toxoceratoides (Colom­
boceratoides).
AS TYPE LOCALITY for the base of this
zo�e the beds immediately above the white,
argillaceous (tuffaceous?) layer rich in compre·
sse� specimens of ''Procheloniceras, cf. Proche­
lomceras albrectUJustriae Hoh." are selected.
These were previously recorded in Etayo-Sema
(1968), "Estacion 10e", columnar section Fig.3,
and "Est.lO", km. 28 on the road Sachica-Puen-
13
te Samac&, geological map. Lithologically this
zone is represented by La Paja Formation, s�g�
i!
� nt E as described by'"Etayo•Seiiia:,--1968. A
rE'lt�
CE LOCALITY Cor the zone is the
previously described sections of the_ Socot8
Member on the road Anapoima-Apulo.
Species referred to the zone described in
this paper are :
Du{renoyia sanctorom Biirgl
Du{renoyia boteroi n. sp.
Du{renoyia codazziana (Karsten)
Stoyanowiceras treffryanus . (Karsten)
Stoyanowiceras cintatum n. ·sp.
Cheloniceras (Epicheloniceras) dognaelui­
rae n. sp.
Cheloniceras (Epicheloniceras) camachoi
n. sp.
Cheloniceras (Epichelonicenzs) carlosacos-
tai n. sp. -�
Cheloniceras (Epicheloniceras) jimenoi n.
sp.
Cheloniceras (Epicheloniceras) barreroi
n. sp.
ChelonicerCI$ (Epicheloniceras) pardoi n.
sp .
ChelonicerCI$ (Epicheloniceras) leonhard­
triedeli n. sp.
Vectisites (Zamur-anoites) cadenai n. sp.
Vectisites (Zambranoites) crozi n. sp.
Vectisites (Zambranoites) zambranoi n. sp .
Vectisites (Zambranoites) duquesi n. sp.
Vectisites (Zambranoites) mateusi n. sp.
Riedelites obliquum (Riedel).
Riedelites (?) quebradanegra n. sp.
Riedelites esthe�emae n. sp.
Juandurhamiceras juandurhami n. sp.
Juandurhamiceras giraldoi n. sp .
Gargasiceras interiectum (Riedel).
Gargasiceras pulcher (Riedel).
Corteziceras cortezi n. sp.
Pseudoptychoceras gilberti n. sp:
Toxoceratoides (Colomboceratoides) ren­
zoni n. sp .
Aconeceras gutierrezpalmae n. sp.
Melchiorites palmeri n. sp.
Eogaudryceras helmsi n, sp.
Eogaudryceras (Eo te tragonites} cabaricoi
n. sp.
Pseudohaploceras leivaense n. sp.
. Valdedo�ella (?) colombiana n. sp.
Colombiceras sarmientoi n. sp.
Eodouvilleiceras (?} tequendamai n. sp.
Colombiceras foreroi n. sp.
�.P..lil.�..aJ.Jw!HI.Gbi.· &gatfJahQuli��&
.J!_p tocerati{orme Ass:_m�lSe - zone
As far as known to me these species have
their most occurrence in this part of the sequen­
ce; the absence of Cheloniceras (Epichelonice­
ras}, Du{renoyia, Stoyanowiceras, Vectisites
(Zambranoites), Aconeceras, Eogaudryceras,
seems indicative also.
AS TYPE LOCA LITY for the base of
this zone the clay-shale beds with abundant
argillaceous-calcareous concretions at point
C-127 on the road Anapoima-Apulo is design-
 14 F. ETA Y O
ated. In the lithostratigraphic nomenclature it is
represented by the unnamed sequence of shales
overlying the Socota Member and below the
Capotes Member of the sequence in the region
of Apulo, Viota, Las Mesitas,
Species referred to the above named zone
Parahoplites (?) hubachi n. sp.
Parahoplites (?) tristan n. sp.
Acanthohoplites bigoure ti{orme n. sp.
Acanthohoplites luisperezi n. sp.
Acanthohoplites seunesiforme n, sp.
Acanthohoplites pluricostatum n. sp.
Acanthohoplites eleganteante n. sp.
Acanthohoplites (?) leptoceratiforme n.sp.
Acanthohoplites (?) serpenti{orme n. sp.
Riedelites (?) filosum n. sp.
Corteziceras latecostata (Riedel)
Melchiorites media Riedel
Both preceding zones represent the pP�r
.1\Rtian of Colombia. The tenn Aptian proposea
by· d'Orbigny (1840, I, p. 631) is here adopted;
its previous use in the stratigraphy of the Creta·
ceous of Colombia has been thoroughly reviewed
by the author (Etayo-Sema, 1964). I find a ge­
neral agreement, in broad sense, between my
succession of faunas and those discussed by
Breistroffer (1947), Casey (1961), and Thomel
(1964) for the European uPper Aptian.
As far as known these two species have
their first occurrence in this part of the sequen·
ce. All the genera recorded for the preceding
zones are absent, but this is not so when com·
pared to Europe; the new species listed below
occur for the first time.
AS TYPE LOCALITY for the zone is
selected the same type locality as for the basal
· part of the Capotes Member, however litholo·
gical characteristics and fossils do n.:�t necessa·
rily correspond ; as a REFERENCE LOCALITY
the same reference locality as for the Capote&
Member is designated. Other species belonging
to this zone are also given:
Ho/cophylloceras hernandezi n. sp .
Douui/leiceras abozagloi n. sp.
1Jouuilleiceras tarapacaense n. sp.
Eodouuilleiceras pedrocarvajali n. sp.
Neodeshayesites karsteni (Marcou)
Neodeshayesites albertoalvarezi n. sp.
Neodeshayesites cingulatum n. sp.
Neodeshayesites contracta (Riedel)
Carloscaceresiceros monteroi n. sp.
Carloscaceresiceras caceresi n. sp.
Trochleiceras (Trochleiceras)
. juliuerti n.
sp.
Trochleiceras(Jaumaportaiceras) hof{ste·
tteri n. sp.
· SERNA
Elatilmemiceras .colombiana-R inconiceras
rinciini�Ljielliceras pseud01ye1Ufor"!e
_5ssembly���
-
.
Some of the preceding species also occur
in this zone but the faunal spectrum changes in
favor of Lyelliceratidae and Engonoceratidae.
AS TYPE LOCALITY of this zone is
chosen the "Estacion 33a" previously recorded
by the writer (1968), columnar section Fig. 9,
and on the geological map annexed to that
paper. Lithologically at its type locality this
zone is represented mainly by "Nivel 8" of the
"Segment B" of the Upper San Gil Formation.
At the type locality the name-bearing species of
the zone have not been recorded yet. AS REFE·
.
RENCE SECTION where the species occur are,
first the clay-shale concretions bearing succe­
ssion found at Cerro Bejucal, extending SE
along the abandoned road and from the machi·
nery house of the limestone mine.
Other species generally occurring in
the zone are:
- Engonoceras (?) duartei n. sp.
-Xnemiceras semicostatum Sommermeier
�-J(nemiceras pegnai n. sp.
�-Engonoceras olgaluciae n. sp.
tTegoceras benavidescaceresi n. sp.
):-Lyelliceras pseudolyelliforme n. sp.
' Lyel/iceras isaacleai n. sp.
� Ralphimlayites apuloense n, sp.
The last two assemblage-zones discussed
represent the lpwer Albian and the base of the
Middle Albian jn Colombia. The tenn Albian
proposed-by� a'Or6lEliY"-<1B43, p. 463) isliere
stressed for its usefulness. There are similarities
···betWeen the· faunas from Pe:rU described by Be­
navides Caceres (1956), from Madagascar by
Collignon (1950) and discussed by Breistroffer
(1952), and with those of the European Albian
(France, England), but the similarity with the
latter areas is less marked than in the case of the
Aptian.
Between the preceding zone and the fo·
llowing zone there is a "barren zone" where
ammonites have not been collected or found.
The dating of these rocks based on other groups
of mollusks are not within the scope of the work.
ite
_loJf!_!!J!!!. �:fr��f2l!��agel�n:s�:;;�'!.��El!es -·
-
=>·-·· · ..
As far as known to me the name-giving
...
'-"=
species have their fU'St occurrence at this part of
·
-
--
the sequence.
The section exposing rocks of La Fronte·
ra Formation1 at La Cabafia, km 23 of the road
Cumaca·Viota is selected as TYPE LOCALITY
of the zone; A REFERENCE SECTION is the
type locality of the San Rafael Formation,
previously described by the writer (1968; geo·
logic map).
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
Other species occurring in thla zone am :
- ·· Mammites scutulatus n .sp. •
--Hop/itoides hemanmojicae n. sp.
... Fagesill zanell6i n. sp.
Imlayiceras (?) ra/phimlayi n. sp.
_ -Prionocycloceras portarum n. sp.
- Mammites {ugax n..sp.
- Hop/itoides lagiraldae n. sp.
- Paramammites co/ombitznus n. sp.
1bis zone ia here considered to represent
tile lower Turonian of Colombia. The Turonian
S taie';--p�l(i'd'"'by d'Orbigny (1842) 1a aJao
adopted; its previous use in the geological litera­
tuft! on Colombia has already been discussed by
the author (1964). The affinities of the listed
fossils am given in the systematic descriptions.
G/oriaceras con-eai-Pro texanltes
cucartilense:Codliizrceras scheroei
�m�l!£e.zone
As far as known these species occur for
the first time in this part of the sequence.
TYPE SECTION of the zone Is the expo·
sores of the Cucaita Member at "Estaciones 6-9"
on the road Cucaita· Piedra Gorda as previo�sly
recorded by the author (1968, p. 45, Fig. 13,
and annexed geologic map); a REFERENCE
LOCALITY is the less accessible outcrops of
the beds above the La Frontera Formation
(Coordinates �42000W-979000N, on Plancha .
M6·1·A, Cundmamarca of the I.G.A.C.).
Other species conside�ed belonging to this
zone are:
15
..aloriaceras paulinae n. sp.
...Peroniceras solorzanoi n. sp.
...Peroniceras dillbloi n. sp.
...Peroniceras guerra/ n. sp.
....Prionocycloceras longispinatum (Basse)
..Prionocycloceras (?) cucaitaense n. sp.
-Nice(oroceras boyacaense n. sp.
-zumpangoceras ospinai n. sp.
.-Zumpangoceras (?) sorae �_. sp.
This zone is here considered to represent
the lower Coniacian. The term Coniacian as
proposeci"""by"'d0qu8iia (1857, p. 748) is here
adopted. Similar faunas occurring abroad are
discussed in the taxonomic part of this report.
Lenticeras baltoi-Paralenticeras sieuersi
Assem6lage-zone
The name-giving species have previously
been discussed by the wnler (1969). .
..
·· .
.
.
The TYPE LOCALITY for the .zone Is
the outcrop of the upper part of the' Conejo
Formation and below the Upper Lidita�'Forma•
tion, about 100 meters south of well .B·20, as
depicted by the writer on the Geologic map of
the Villa de Leiva Area, near the road ·Puente
Samaca-Cucaita (1968). The Santonian . Stage
also proposed by Coquana1t857,-�p� - 749) is
here stressed, however the basis for Its recogni­
tion is not especially strong when the poverty of
ammonites now known from Colombia is consi­
dered.
The proposed zonation is summarized in
Table 1.
16 F. ETAY O · SERNA

TABLE I

ASSEMBLAGE ZONES PROPOSED FOR USE IN COLOMBIA

z
<
not represented
z
-

0
E-<
z Paralenticeras siiwersi
<
Len ticeras baltai
Cll

z
not represented
<
0
-
.

< Gloriaceras correai Peroniceras diabloi Niceforoceras

z
-

0 Protexanites cucaitaense Prionocycloceras (?) cucaitaense

0
Codazziceras scheibei Zumpangoceras - ospinai

z
< not represented
z
-

0
� Mammites nodosoidesappelatus Hop litoides lacabagne Fagesia zanellai
0
E-< Franciscoites suarezi Mammites fugax Paramammites

0
+ not represented
<

Platiknemiceras colombiana L_y_e lliceras carvajalorum

z Rinconiceras rinconi Knemiceras pegnai

<
- - LyeUi9�as pseudolyeli.iforme Engonoceras ( ?) duartei
-�
..::I Neodeshayesites columbianus R:�ZIJ_himlayi_t�s Jaumaportaiceras
<
Carloscaceresiceras caceresi

Douvilleiceras solitae Neodeshayesites karsteni

Parahoplites (?) hu bachi Acanthohop lites ( ?) leptoceratiforme

z Juandurhamiceras R iede lites

<
-
Stoyanowiceras treffryanus Corteziceras A coneceras
E-<
Zambranoites Colombiceras
(:I._

< Colorn boceratoides Gargasiceras

.

Pseudoptychoceras Epicheloniceras
Dufrenoyia sanctorum Melchiorites� Pseudohaploceras
 ZONATION OF THE CRETACEOUS OF CENTRAL C OLOMBIA BY AMMONITES 17

SYSTEMATIC DESCRIPTIONS half, they project onto the venter fanning an

adoral arc. The pleats increase their width slow­
Phylum MOLLUSCA ly but steadily towards tho venter.
Class CEPHALOPODA
There are shallow but wide constric­
� �
Order AMMONOIDEA
Family P LL RATIDAE tions that approximately follow the trend of
Genus I Ph]l
o te
uess , 1866 S the ribs but project strongly forward as a lingui·
fonn arc on venter; they are weU developed on
Phylloceras Suess, 1866, p. 7. the internal mold but only as broad and low
areas on the surface of the test. On the adoral
TYPE SPECIES. A mmonites heterophyUus portion of the last whorl the shell Is preserved
J. Sowerby, original designa­ and the pleats are well defined as flattened belts
tion. of growth striae wider than the intercostal
spaces. These bel ts stre tch approximately to the
Phylloceras. buchjqnu �;(Forbes) ? adumbilical third of the flank .where they fade
PI. 8, F1g. 4; Text- 1g. 3C. ou t and only fine raised growth lines extend
toward the umbilicus. The periumbilical area
1844 Ammonites Buchilzna Forbes, p. 177, seems smooth. Strigations are visible especially
Fi�· a,b, under the species name. around the umbilicus.

? 1858 Ammonites Buchilznus Forbes, Karsten, The suture line has L. trifid�,., ,
_

p . 84.
DIAGNOSIS. This species Is characterized
1881 A. rmmonites.] BuchitJna Fbs., Hyatt, by: A) its sinuous, low ribs especially developed
p. 3'10. on the external half of the flank, B) the lingui­
fonn projection of the constrictions on 'ventei:
1883 [Phylloceras] BuchitJna Forb., Uhlig, on the intemal mold, C) the broadly rounded
p . l82. section of whorls on venter.

A mm. [onitesl Buchiana

' '
·

1907 Forbes, P�r- Meuuremenu. D Euw H W DfEuw % D/1{%

vinquiere , p . 5 0. Hypotype 14 3 ?7 8 21 ?60

1910 Ph. fylloceras ] Buchianum Forbes, COMMENTS. This peculiar species has
Fallot, p. 70. been synonymized with P. uelledae (Michelin)
d'Orb. [ Fallot, 1910, p. 70; Gignoux, 1920, p .
1920 Ph. [ylloceras] BuchitJnum Forbes, 91], but Spath (1923, p. 17) h as given it
Gignoux, p. 91. specific standing because "It i s more inflated
throughout" ; in Forbes' species the periventral
1923 Ammonites Buchiana, Forbes, Bose, and ventral zones fonn a broader arc than in P.
p. 125. velledae. Phylloceras moriezense Sayn (1920,
p. 194, pl. 1, Figs. 2,3) has a similar develop·
1923 Ph. [ylloceras] buchitJnum (Forbes ), ment of the ornamentation but the whorl
Spath, p. 17. section seems less inflated and its ribs are clearly
rounded in contrast to those of Forbes' species.
1924 Ammonites buchiana Forbes, Sherborn, However on both species the ribs are slightly
p . 925. projected on venter.

? 1936 Phylloceras cf. BuchitJnum Forbes, OCCURRENCE. Capotes Member.

Breistroffer, p. 156.

? 1955 Phylloceras cf, BuchitJnum Forbes, ex Genus \Holcophylloceras]Spath, 1927.

Breistroffer, :aurgl, p. 13.
Holcophylloceras Spath, 1927, p . 38.
1964 Phylloceras buchianum (Forbes), Etayo
Serna, p. 101. pars. TYPE SPECIES , Ph l oce ras mediterraneum
yl
Neumayr, original designa­
Holotype B.M.N. H. . tion.
Hypotype C-217. (U.N.C.M.H.N.C.P.Hy. N .1)

I am referring to this species a group

of small, inflated and Involute conchs; they ha·
ve a broad venter and convex sides that round Holotype C-13 (U.N.C.M.H.N.C.P.Ht. N. 1)
abNptly into a deep umbilicus. Paratype C·13 (U.N.C.M.H.N.C.P. Pt. N.1)

The adapical part of the last p reserved
whorl of the hypotype has low, broad, subdued
�leats especially visible with oblique illumina­
tion. These pleats h ave a sinuous trend, they
describe an arc on the inner half of the flank
and a broad adoral concavity on the adventral
In the juvenile the conch is inflated, na·
rrowly umbilicated, the umbilicus is deep; the
conch Is divided into protruding segments by
biconcave constrictions that are Ungulate on
venter, when the shell is preserved one adapical
ridge parallels the contrictions; the omamenta-
18 F. ETA YO · SERNA
tion consisfli only of rme growth lines that tend
to form almost imperceptible bands that para·
Del the constrictions and ridges. There are seven
constrictions on the preserved last whorl.
The section of the whorl is subquadrate,
slightly wider than high, it has flat parallel flanks
which are weakly beveled at the periventral
margins producing a narrow venter; the umbilical
wall is oblique on the adapical segment of the last
whorl, it is subvertical on the adoral portion.
There are seven constrictions on the last preser­
ved whorl. 'The suture line has subrectangular
bifid saddles and wide, slightly asymmetrical
tripartite lobes.
In the adults the whorl section is subqua­
drate , slightly wider than high; the flanks are
flat but round into a narrower feebly arched
venter; the periventral z one is feebly beveled.
The umbical wall is subvertical and the umbili­
cal rim is rounded.
The conch is divided into segment"$ by fi.
ve wide and shallow constrictions on the inter­
nal mold,these are represented by very feeble
low zones on th e shell and are rimmed ad orally
by a slightly thickened ridge. 'These constric·
tions follow the trend of the growth Iirae.
'The ornament consists of raised growth
lirae that gather and crowd in broad , flat,
rib·like sheaves; now and then these sheaves
may raise as low ribs, they begin at the umbili­
cal seam where they form a minute adoral con­
cavity just near the umbilical seam from which
point they trend prorsiradiate up to the adum­
bilical third of the whorl where they bend
abruptly to form and arc on the middle and
p art of the ad ventral thirds of the flank; on th�
periventral margin they forin a conspicuos
adoral concavity and on the venter form an
acute adoral arc. The internal mold in covered
by low undulations corresponding to the
sheaves of striae.
The suture line has phyUoid bifid saddles
with narrow bases; E/L is bifid and its external
branch is also bifid and its lower branch contacts
the midventral saddle.
DIAGNOSIS. This species is characterized
by: A) the conch divided into protruding seg­
ments between deep sinuous constrictions, lin­
gulate on venter in the young, B) flattened seg­
menbi of the conch between attenuated cons­
trictions in the adult, C) its falcoid sheaves
of lirae stressed adventrally, and D) its qua­
drate whorl section with feebly beveled peri-
ven�argins. i '
Meuurementa D EuW H W E(Euw •fJ)fH'%
Holotype 21 6 10 11 24 48-
Pmtype 13 3 7 8 23 64
- COMMENTS. The early whorls of this spe­
cies resemble Valdedorsello getulina (Coquand)
as Ulustrated by Pervinquiere (1907, p. 15 1, pl.
6, Fi g. 16), but the latter has a depressed whorl
section. The Te trogonites-Iike aspect of the adult
whorl is a mere chance morph ological similarity
because in contrast to the latter genus the Co­
lombian species has L tripartite.
These striking differences between the
juvenile and adult whorls are of common occu­
rrence in Phylloceratidae (see Wiedmann, 1963,
p.186); the specimens I consider conspecific
came from the same concretion and are similar
in the development of the subquadrate whorl
section .
Of the Cretaceous species referred to this
genus, Holcophylloceras guettardi (Raspall) [in
Wied man�,. 196 3, p. 258, ,Pl. 17, Fig. Sa, 8b] ,is
most sirnil.ar to my spec1es. In the European
species the whorl section is rounded (Wiedmann,
ibid, text-Fig. 62a). In the Colombian species the
growth striae are accentuated adumbilically
contrary to those of the Mallorcan species.
This species is named in honor of the zoo·
logist Jorge Hernandez, of the Universid ad Na­
cional de Colombia.
OCCURRENCE : Capotes Member. Lower
Albian.
Genus l Hypophyllocer3Salfeld, 1924.
Hypophylloceras Salfeld , 1924, p.5
TYPE SPECIES. Phylloceras onoense Stanton,
original designation.
Jil'llPDlWiacerg§ wjed :W
IJ.i n. sp.
Pl.10, Fig. 3; Text-Fig� ·
? 19 3 8 Phylloceras velledae Mich. var seresiten­
se Perv., Riedel, p. 8, pl. 4, Figs. 6,7;
pl. 12, Fig. 7, pl.14. Fig. 1.
? 1968 Phylloceras serecitensis sensu Riedel
(non Perv .), Etayo-Sema, 1968, Fig.3,
Table I.
Holotype . C-255 (U.N.C.M.H.N.C.P.Ht. N.2).
The conch is involute, narrowly umb ilica­
ted, the umbilicus is rimmed by a narrow plano·
concave surface. The adumbilical half of the
whorl has an almost smooth surface on which
with adequate illumination, the extension of the
ribs are visible as rme threads. The adventral
third of the whorl-side is convex and covered
with fme, rounded ribs, separated by subequal
intercostal spaces up to twice as wide as the ribs.
The boundary between the callused and ribbed
<::csurfaces is irregul ar through ou t t.p,e le n gt.tt o f the
· last chamber. Toward the adapical segment of
the last preserved whorl there is a more promi­
nent rib that ex tends to the umbilical wall . The
trend of the ribs is straight or feebly concave
adorally across the umbilical rim. The ribs des·
cribe a wide adoral arc on the fianks, a weak
adoral ·concavity on the imprecise periventral
area, and then croBS the venter and bend sUngb·
tly forward. At the boundary of the callused
surface of the nank the ribs seem to be of uni·
form width, they thin below the glazed zone
 ZONATION OF TilE CRETACEOUS
and tend to fonn bundles some of which beco·
me more prominent near the adumbilical third
of the whorl side that appelll'll as a surface cove­
red with irregularly distributed pleats. The sec­
tion of the whorl has a broadly rounded venter
and flattened diverging flanks which at the urn·
bllical rim attain their ,!Daximum width and
then tum abruptly into the. oblique umbical rim.
No living chamber is available.
DIAGNOSIS. This species is characterized
by: A) the callused adumbilical half of the whorl,
B) the channel-like intercostal spaces, the broa·
dly arched trend of the fme rounded ribs on the
flanks, and their feeble projection on venter,
C) the whorl section with broadly rounded
venter and central third of the flank flattened,
and D) the plano concave adumbilical surface.
Meuurementl D Euw H W DfEuw % D/H.04
Holotype 46 6 24 18 13 54
COMMENTS. On Riedel's specimen of
Phylloceras velledae Mich. var. seresitensis Perv.,
from Utica, not far from the region studied in
the present report, (see especially Riedel's illus·
tration of the internal whorl, pl.4, fig.6) the
adumbilical half of the whorl is eroded so the
callused co,yering and urd�rlying fme . ribbing
can not be recognized, but Riedel remBl'ks:
"Los flancos estan cubiertos de numerosas listas
finas que, naciendo aproximadamente a media
altura de las espiras, siguen derecho bacia el
!ado extemo, donde forman un arco apenas
perceptible hacia adelante". Riedel commented
that his specimen does not' show the falciform
trend o"f the ribs as in Ph. velledae Michelin (see
the figured cast of the neotype of this species in
Wiedmann, 1963, pl.12). The lectotype of Ph.
seresitense Pervinquiere (1910, p.9, Fig. 2, pl. 1,
Fig. 1; se·e Wiedmann, 1963, p.222) shows a
flattened surface of the flanks that descends
funnelwise to the umbilicus, this surface is
reduced to a mere rim around the umbilicus in
Riedel's figured specimen and in the present
specimen.
Wiedmann, the last reviewer of the
group (1963), recognized three "subspecies" of
Ph. seresitense based on the section of the whorl
and the width of the umbilicus. In the present
species the umbilicus is narrow and the section
of the whorl, like that figured by Riedel (1938
pl,. 12, Fig. 5) has a broadly rounded periventrai
shoulder rather than beveled shoulders as seen
i� the lectotype of PerVinquiere (1910, pl. 1,
F1g. lb) and the hypotype figured by Wiedmann
(1963, pl.21, Fig. 1b).
. The. flattening of the flanks in the Colom-
bian sp�cunens produces a middle third of the
whorl 11de that diverges from the venter and
�en �p�ptly rounds into the umbilicus; in Per­
\'Ulqulere s illustrations, both middle and
adumbilical thirds of flanks are flattened but
sho"! . a. tendency to converge toward the
umb�1cal rim. Wiedmann used the flattening of
the s1d�s as a specific character (1963, p.202·
203, F1g_. 47b): Phy/locerrl$ (Hypophylloceros)
subserec1tense Wiedmann. The trend of the ribs
OF CENTRAL COLOMBIA BY AMMONITES 19
In the latter species is radial to slightly concave
adorally (?) (op.cit., pl. 15, Fig. 6) on flanks.
The subspecies P. (H.) seresitense tanit Pervin·
quil�re (Wiedmann, p. 228, fig. 54) has a bicon­
cave type of ribbing.
This new species is named in honor of
Dr. Jost Wiedmann, of the Tiibingen Universitat,
West Germany.
OCCURRENCE. Capotes Member. Lower
Albian. Paja Fonnation (Arcillolitas abigarradas
Segment E.). Upper Aptian.
Family G UJR c
Genus \E<(aud c
��path,
� ;; eri1Sj3 TIDAE
1927
Eogaudrycerrl$ Spath, 1927, p. 66.
TYPE SPECIES. Lytoceras numidum Coquand,
original designation.
d!:.J!�!.!I§: lJelr.rfin. sp.
-Jf'l!!:&
[s;Fig. 14, TexT-fg. 31.
Holotype. C-223. (U.N.C.M.H.N.C.P. Ht.No. 3).
The only specimen available was disected
back to the first whorl of the phragmocone;
the first whorl has an arched venter which is
divided into segments by prominent rounded
flanges which are accompanied by flattened
shallow furrows just paralleling them on both
sides across the venter; in the intervening spaces
there aie fine widely separated lirae. The whorl
section at this stage is subcircular although it
has tendency to be wider than high. The second
whorl has reniform outline with wide evenly
arched venter and convex sides that round
smoothly into the umbilical wall, in a wide,
rounded transitional zone without an edge.
The shell is ornamented by acute lirae
that describe a wide adoral concavity from the
umbilical seam up to the beginning of the
flanks, then they described a wide arc on the
flank to finally straightly cross the venter. The
striae tend to gather in bundles toward the
periumbilical rim, these bands of striae are.
separated by narrow and depressed areas or
furrows which however may not be called
constrictions, consequently the lntemal mold is
characterized by slight transverse swellings. The
lirae are not of uniform strength and on the
internal mold they are represented by fine
grooves whereas the interspaces appear as flat·
tened and much wider bands. Some slightly
wider and accentuaded transverse furrows are
interspersed. On the adapertural part of the
second whorl and the entire third whorl the
bundles of lirae are conspicuous along the
umbilical rim and the interbundle spaces are
better defined as smooth, relatively wide bands;
the bundles tend to group five lirae, but two
bundlea may fuse together and finer Iirae may
develop also. On the third whorl the section is
reniform, much wider than high and rather
flattened on venter; toward the aperture of the
fourth preserved whorl the section is rectangu·
lar with rounded comers; there seems to be an
20
increasing overlap of the whorls with age.
DIAGNOSIS. This species is characterized
by: A) the change in shape from subcircular to
rounded rectangular cross-section in the ontoge·
ny, B) the bundle fonning convex Urae, and
C) the lack of 'constrictions' in later stages of
growth.
Meuurementa D Euw H W DfEuw %
Holotype 17 8 11 1& 41
COMMENTS. This species differs from
Eogaudryceras vocontianum (Fallot) [ 1921, p,
2 33 pl. 2, Fig.2] because the lirae are radial ('!)
on bank of his species, (Fall ot, op.cit., p.235)
and the constrictions are accentuated, further·
more its maximum thickness is toward the um­
bilical margin whereas in the Colombian
species it is located toward center of the flank.
The specimens of Fallot's species referred to by
Wiedmann (1962, p. 40, pl.2, Figs. 2,6,8) as
showing variation of the width of the section
(ib., p.41), clearly show a high vaulted section
also. From Eogaudryceras numidum var. inter·
mediLJ (Fallot) f1920, p. 235, 1?1.2, Fig.3: lecto·
type, Breistroffer, 1947, p.56j the Colombian
species differs because its cross section profile is
more evenly rounded and wider and the less
accentuded constrictions on its flanks attenuate
very early.
This species is named in honor of Dr.
J. Helms of the Humboldt Universitat in Ber·
• TYPE SPECIES.
lin, DDR.
OCCURRENCE. Socota Member. Upper
Aptian.
Subgenus
. j
(Eo etraponites:[D
19 7.
1nreistroffer,
Eotetragonites Breistroffer, 1947, p. 56.
TYPE SPECIES. Lytoceras (Tetragonites) de­
pressus Raspail, original desig­
nation.
..fogaudr:x,ceras (Eotetrag,onites2 cabaricoi,..
n. sp.
Pl. 6, Fig.ll; Text-Fig. 3M, 3N.
Holotype. C-513. (U.N.C.M.H.N.C.P. Ht. N.4).
On the earliest whorl observed (the se·
cond?) the section is reniform, wider than high,
it has a broadly rounded venter; the whorl sides
are convex and round into a vertical umbilical
wall. The ornament on the internal mold con·
sists of rounded prominent undulations which
fonn feeble adoral arcs on the whorl side, they
are separii�d by constriction-like low zones; on
the test fine transverse lirae are visible on venter.
On the_ th.�. _whorl· [of the paratype ] the oma­
m�nt consiits of bundles of striae, but wide
constrictions that follow the pattern of the
bundles develop on the opposite sides of the
whorl without crossing the venter. On the third
whorl, the whorl section is subquadrate and
has a clear tendency to become higher than de-
F. ETAYO · SERNA
pressed; the sides and venter are nattened.
DIAGNOSIS. This species is characterized
by A) the ontogenetic changes of the section of
the whorl from renifonn in the inner whorls to
subquadrate In the outer whorls, and B) the
sporadic strong constrictions that do not cross
the venter.
D{H % Meuu�ements. D Euw H W D(Euw % D{H%
61 Holotype 26 6 11 14 24 44
COMMENTS. This species is comparable
to Eotetragonites jacobi ambigua Breistroffer
f= Guadryceras in termediLJ Fallot, 1920, pl. 2,
Fig. 6, 10), from the French Gargasian but on
the latter species the constrictions are more nu­
merous and frequent than in the Colombian spe­
cies which furthennore is more robust and invo­
lute.
This species is named in honor of Mr. Ma·
nuel Cabarico, of the Geological Survey of Co­
lombia.
OCCURRENCE. Socoti Member. Upper
Aptian.
Family
ncM -c Genus
Toxocerat6ides Spath, 1924, p. 78.
Toxoceras royeriLJnum d'Or­
bigny (I.C.Z.N.?).
Subgenus-? {QiJo}itbog��}!!!J n. subge-
nUS
::- 7Q not/a,. o'kJ , fp<J-�1! I _) ;) 2 Y
TYPE SPECIES. Tox ergJoides (.Colomboce·
,!g_tofJE
· es) re'!.zoni n. sp.
DIAGNOSIS. Coiling as in Toxoceratoides
but differs from the latter genus by the deve·
lopment of sculpture characterized by a retar­
ded development of the subdue lateral trituber­
culation: early ribs without tubercles, followed
by the appearence on both sides of venter of
slender spines, subsequently lateral tubercles
appear and much later the nipple-like peridorsal
tubercles show up. The suture line is much sim·
plified, it has massive subrectangular saddles
and a subtrifid L.
Toxoceratoides (Colombocerqtoide§)
iJ
renzo Jl. sn.
·
Pl."6', ig. 19; Text-Fig. 3. 0.
Holotype. LB-10 (U.N.C.M.H.N.C.P.Ht.N.6.)
The earliest portion of the slightly curved
shaft has a suboval whorl section: flattened to
feebly rounded dorsal area that merges into the
• evenly rounded sides which interconnect thro·
ugh the narrow arched venter.
The ornament consists of prominent
raised rounded ri&s that thicken toward venter
and that cross the venter prorsiradiate or des·
cribe an inconspicuous adoral arc; on the dorsal
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
area the ribs describe a wide adoral arc and on
both sides of this arc two narrow adoral conca­
vities are formed; the ribs then trend obliquely
and slightly arcuately across the flanks and fl.
nallY they cross normal to th� ventral medial
axis where they become depressed and even
very attenuated but not ln terrup ted. The inter­
costal spaces are channel like with rounded
bottom and usually subequal in width to the
ribs but occasionally are a bit wider. The moilt
adapical preserved segments of shaft h ave ribs
of uniform strength that bear low, rounded to
compressed bullae on either side of the vef\tral
median depression;· now and then however a
fmer, almost ventrally nontuberculated rib
occurs, likewise stronger and ventrally noded
ribs are intercalated .
With maturity a clear differentiation of
the ribs occurs: one non stressed rib alternates
with one strong rib which bears one spinose
tubercle on the periventral margin and a second
mammiform tubercle at the boundlllY betwen
the peridorsal and central thirds of the flank;
the costal whorl section acquires a beveling of
its profile on the adventral third of the flank.
With further growth the primlllY ribs develop
an additional, smalle r, mammiform tubercle
near the rim of the dorsal side but the ribs are
nearly interrupted on venter, developping a weak
channel along the ventral median line; two
intercalary ribs may occur on mature stages.
Later the section of the whorl becomes roun·
ded subrectangular, the segment of the ribs
between the ventral and lateral tubercles is
flattened, with a tendency to split longitudinally .
DIAGNOSIS. This species is characterized
by A) the rapid increase in width of the shaft,
B) the ontogenetic development of its ornament
from undifferentiated barely tuberculate · ribs,
not interrupted on venter, to an alternation of
trituberculate primary ribs and almost non tu·
bereulate intercalatory ribs all of which become
attenuated on venter, and C) the progressive
development -of the lateral tubercles.
COMMENTS. This species differs from
.Toxoceratoides royerianus ("d'Orbigny) as emen·
ded by Casey (1961, p. 77) because the ribs do
not divide on the dorsum or if they do no more
than two rible ts appear; they differ also in the
suture line which is less complicated in the Co·
lombian species. However this difference may
be due only to the diameter of the whorl at
which the sutu:res were drawn.
. This species is named in honor of the geo·
logqt Giancario Renzoni.
OCCURRENCE. Paja Formation (Arci·
llolitas ablgard
•
ra as. Segment E.). Upper Aptian.
Tox�ceru toides (Colomboceratoides} ren­
zom n. sp.
. (M orph I).
Text-Fig. 3P.
Paratype. (U.N.C.M.H.N.C.P.Pt.N.2.)
21
The shaft Is slightly arched. The early sec­
tion of the whorl is vaulted; the dorsal area ia
arched with the sides rounding outwardly up to
the center of the whorl then evenly rounding
toward venter which is flat.
The ornament consists of prominent
rounded ribs which cross the dorsal area linear­
ly or with a slight adoral bow, they cross the
flanks forming f�rst a weak adoral concavity on
the dorsal rim and then trending prorsiradiate
or describing a slight arc, or even forming an
adoral concavity ; on venter they become inte­
rrupted -on the internal mold- or very atte­
nuated on exterior of the test around the ven·
tral medial line; the end of the ribs converge
toward the siphonal line forming a w ide V with
its apex pointing adorally . All ribs •develop
rounded to spinose periventral tubercles. Though
sporadically one rib may show greater strength,
the ribs gain strength progressively and do not
develop a regular alternation of strong and
weak ribs. The intercostal spaces are deep and
twice as wide as the ribs on the internal mold,
but they appear shallow and narrower when the
shell is preserved. ·
· .
NOTE : This morph is notabl_e- for the steady
growth of the section of the shaft, by
the strong, uniform and unituberculate
ribbing and the early stressing of the
ventral tuberculation.
OCCURRENCE. Socoti Member. Upper
Ap tian.
Family P Y HOCERATIDAE.
(}
Genus � 1 Pseudoptychoceras n.
.=:: £o.plyrh11u -""A. �
TYPE SPECIES. Pseudoptychocerrzs gilberti n.
sp.
DIAGNOSIS. Straight or slightly undula·
ted, slowly increasing initial shaft, ornamented
with prominent, oblique annular rounded ribs,
separated by conspicuous constriction-like inter­
costal spaces; the section of the whorl is ellip­
tical to almost circular, the first shaft bends to
a sh orter (?) branch which leaves an opening
("needle hofe") before appressing against the
:lorsum of the first shaft; this second branch is
ornamented with annular acute ribs distant
from each other. The suture line has short sub·
rectangular elements, relatively undisected, L
Is trifid.
COMMENTS. This new genus differs
from Ptychoceras because of its loop forming
hook and its trifid L; from Euptychoceras
(Breistroffer, 1947) because the fanner Is
smaller and has strong ornamentation on both
shaft and hook.
Pseu doeachoceras gjlberti, n.
sp.
P[G, li'ig. 18; Text-Fig.3S, 3T.
Holotype. C-223 (U.N.C.M.H.N.C.P. Ht. N. 6).
Paratype. (U.N.C .M.H.N.C.P.Pt. N.3) .
•
There are three fragments of the shaft,
two of them with part of the hook preserved.
The shaft is straight, progreSBively tapering, It
22 F.
has a subelliptical to subcircular section: the
dol'Bal region is wide, flattened, abruptly chan­
ging into the feeble convex sides which evenly
round toward the narrow arched venter. There
are broadly rounded, low ribs, slightly thicke­
ned on venter. On the dorsal area the ribs are
barely visible, they cross it as fme lirae normal
to the dorsal-central axis; at the margins of the
dorsal region the ribs become well defmed and
cross the flanks of the shaft in an arc convex to­
ward the hook; they cross the venter normal to
the ventral-central axis; the intercostal space,
constriction-like in appearance, is prominent
on flanks and venter and subequal to slightly
narrower than the ribs.
The beginning of the hook Is indicated by
three narrower ribs which appear as a bundle at
the dorsal margin; of the three ribs, that nearest
to the hook is the most prominent and the one
adjacent to it Is the weakest but this relation­
ship may be inverted on other specimens; a
second group of two ribs follows on the hook
and thereafter only single ribs appear; on the
hook the ribs tend to taper acutely toward the
margin of the dorsal area and to broaden on
venter; toward the beginning of the hook the
ribs sometimes develop a weak tendency to
depress along the ventral-central line.
DIAGNOSIS. This species is characterized
by: A) the change of tbe whorl section through
development from circular to subovate, and B)
its peculiar broad ribbing which gave it a worm­
like aspect.
COMMENTS. This species diffeD from
the type species of the genera Ptychoceras and
Euptychoceras as discussed under the genus.
OCCURRENCE. Socota Member. Upper
Aptian.
Pseudoptychoceras gilberti n. sp.,
rMorph I]
1'1.6, Fig. 17.
Paratype. (U.N.C.M.H.N.C.P.Pt. No.4).
One complete hook (?) and a section of
tbe shaft. The dorsum is flattened and rounds
smoothly into the slightly bulging flanks which
in tum grade into a narrowly arched venter.
There are very broad rounded ribs separated by
constriction-like interspace& which cross the
flanks obliquely and project slightly on venter;
both rib and constriction fade out at the mar­
gins of the donal area. The hook has only single
ridge-like rib8'""whicb taper toward, the dorsal
area and are rounded on venter, the adapical
aide of these ridges is steep whereas the adoral
side Is elongated giving that part of the shell the
appearance of consisting of interlocking seg­
ments.
NOTE : These fragments have the section of
their shafts more inflated around the
dorsal zone, the ribs are strong and
rounded and cross the venter oblique­
ly, and the ribs form interlocking rid-
ETA YO · SERNA
ges on the living chamber.
OCCURRENCE. Socota Member. Upper
Aptian.
=il
··---�am A�
�Y:_�� C�
O!I!i
N� C�
E� E!ii
Rif. �A
iD
T:il
A:: !i(jE
m,--
uenus @co1eceros(Hyatt, 1M3.
r;rco��c�ros H'yaft, 19oa·;·p.loo.
•.••,. • •,.,....,.• • • � • •. . • -.. :.: - • -�.. . ·� . .• . - 4 ' • .- -. . . . •..- · .··-:-· ..�
TYPE SPECIES. Ammonites nisus d'Orbigny,
monotypic.
"ficw "ros�tierrezealmf.- n.sp .
.6, Fig. 15;exi-Figs. 3�3Y, 3Z.
Holotype.C-223 (U.N.C.M.H.N.C.P. Ht. N.7)
Paratype. (U.N.C.M.H.N.C.P. Pt. N.5)
A) Up to 10 mm diameter. The fil'Bt whorl
of the phraginocone is inflated and has a whorl
section only slightly higher than wide, evenly
rounded and with a wide ventra] area. On the
specimen studied the internal mold of the fiDt
whorl shows a rounded rib parallelled on each
side by a rather shallow constriction. On the se·
cond whorl the section is ovoid and has a narro­
wer ventral zone. On the third whorl the flanks
become flattened and the ventral zone fastigate
and slightly shouldered at its margins, the umbi·
Iical wall is short but vertical with a blunt um­
bilical rim. The ornament consists of bundles of
falcate pleats, they trend prorsiradiate to weakly
concave from the umbical rim to the boundary
between the adumbilical and middle thirds of
the flank where they bend sharply, fonning cres·
centic adoral concavity and fmally projecting
on venter. The pleats are stressed and slightly
raised about the involution line where the sharp
acute bend occurs. The space between the plea�
is narrower than the pleats themselves.
B) Up to 20 mm diameter. The most im·
portant aspect of the conch at this diameter is
that the umbilicus has not increased its width at
all, in spite of the great increase in the height of
the whorl; the sides are noticeably flattened
and the venter is planifastigate.
The suture line has E/L divided by a lobe
into two branches of which tbe external one is
the widest and it is also bisected; the internal
branch is subdivided into three ramose bran·
ches. The L/U2 has the internal branch larger
than the rest of the saddles; the external lobe is
subsymmetrical.
DIAGNOSIS. This species is characterized
by: A) its ontogenetic development changing
from inflated involute to septicarlnate oxycone;
the internal mold is almost planlfastigate, and
B) iUI strongly falcate Urae and the tendency of
these to become stressed about the centrolateral
,cute bend.
M euuremental D Euw H. W D/Euw0h, D /H Ofo
Holotype 19 2.. 11 5 10 57
Paratype 9 2 5 2 22 55
COl\!MENTS. This species can be differen-
 ZONATION O F THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
tlated from Aconeceras nisus d'Orbigny (1841,
p.184, pl. 55, Figs. 7, 8, 9; Sarasin 1893, p. 153,
pl. 4-6, Fig. 9a, b, c, Text-Fig. 1, 2) irrespective
of size, mainly by the more massive and rectan­
gular base of E/L in the Colombian species. On
Sara&in's Figs. 2,3 (ib., p. 153) and in d'Orbigny's
Fig. 9 of plate 55, the io_temal branch of E/L is
stronger than the exteni81 branch of the same
saddle, but in the Colombian specimens the re·
verse 1s true.
On Aconeceros nisoides (Sarasin) [ 1893,
i.155, pl.4-6, Figs. lOa, b,c, Text-Flg.3; Casey,
1. p. 125, pl.26; Figs. 3-5, Text-Fig. 41a-c],
96
againthe internal branch of E/L is the predo­
minant (see also Fig. 4 le, p.127, Casey, op.cit.);
the growth striae in the Colombian species are
more acute than in the French species (see
Casey� op .cit., fig. 4 1c). The specimen figured
by Somay (1955, p.l2, pl.l, Fig.14) has
a juvenlle suture with E/L symmetrically bifid.
Incidentally , immature whorls of the
Colombian species bear constrictions that are
absent on mature whorls;
': ·
. 135, pl. 9, Fig. 9a, b, 10) has a higher and
OCCURRENCE. Socota Member. Upper
Aptian.
Family
IMeChiorite{
Genus
Melchioriie� sp�tit. i923, p.33 -
---
_.,.... ,. . . .. .... . ., . · · ' · . __... . ... . .
TYPE SPECIES.
*e�htBrites Tex
media Riedel
: : ig. 9;, 3 R.
1938 Melchiorites e merici Rasp. var. media
· Riedel, 1938, p. 17, pl. 4, Figs. 9 · 1 1
pl.12, Fig. 7, pl. 14, Fig. 3 .
· · 1957 Melchiorites emerici meaius Riedel,
Burgi, p. 134.
1964 Melchiorites emerici medius Riedel,
Etayo-Serna, p. 111.
non 1965 Melchiorites inca (Forbes), Btirgl, in
.. Campbell arid BUrgi, L. C. D. 8393.
1!:1.65 Melchiorites emerici media Riedel,
BUrgi, in Campbell and Btirg� L.C.D.
839"3.
l{olotype. Riedel's figured specimen, 1938, pl. 4,
.. . ·•
. . . . .. Fig. 10-11.
�!otype.C-127 (U.N.C.M.H.N.C.P.Hy.N.2.).
· ,.t'e l ocality . Utica (Cundinamarca).
� umbilical rim and one or two shorter intercala­
Two specimens are available, one of them
. .· · ...
deformed. The conch is moderately involute· the
section of the whorl is subquadrate, feebly �on­
vex tq Oattened and subparallel on adumbilical
halt of. flanks, then slightly converging to the
feebly arched venter; the umbilical wall short
is
vertical, and rounds abruptly Into the flanks. In,
23
both specimens the adapical third of the last
preserved whorl is septate, the remaining part of
the conch represents the living chamber; there
are constrictions which begin at the umbilical
seam, they are falcoid and wide, they describe
an adoral arc on middle of flank and they pro­
ject in a strong linguiform fashion on venter.
On the test of the hypotype C-127 there are
faint falcoid sheaves of stressed growth striae
forming thin, flat ribbons that project on venter
paralleUing the constrictions; on the living cham­
ber this ornament is visible also on the internal
mold where the ribs still retain a uniform thick­
ness.
DIAGNOSIS. This species is characterized
by A) Its compressed section of whorl, slightly
taller than wide, and B) its peculiar sheaves of
rme striae that form low, flat falcoid bands on
test.
Meuu:rementa. D Euw H W D(Euw % �-jH%
Hypotype 18 5 7 7 28 38
COMMENTS. The type species of the ge­
nus, "Ammonites melchioris" Tietze (1872, p.
narrower vaulted whorl section than Riedel's
species. Riedel has .discussed other comparable
DE MOCEPc � ATIDAE
Spath, 1923.
species.
OCCURRENCE. Unnamed unit above the
Socota Member and below the Capotes Member.
Upper Aptian.
- �- ..
A mmonites melchioris Tiet­ .Me.l£hiorites121Jlmruj n. sp.
ze, original designation. PI. 6, Figs. 6, 10; Text-Fig. 3$
'? Holotype. C-223 (U .N.C.M.H.N.C.P.Ht.N.8).
t:Fig. 3 Q Paratype. (U.N.C.M.H.N.C.P.Pt.N.6.).
A) Up to 20 mm fC-2001. Whorls slightly
wider than high (at D : l1 , H : 7 , W:9); each whorl
covers approximately half of preceding whorl.
The inner whorls have rather close, narrow but
conspicuous oblique constrictions; on the ada­
pica! segment of the last preserved whorl of this
specimen the constrictions begin at the umbili­
cal seam, slant across the umbilical wall, then
bend and widen near the middle of the flank,
and next project forward as a linguiform arc,
that is most narrow at the ventral median line.
The constrictions are preceded adapically by a
rib which starts near the middle of the flank
and on venter it projects in the same fashion as
the constrictions; a very shallow depressed zone
adorally precedes the rib also.
The ornament becomes distinctive with
growth. It consists of fine rounded riblets some
of which begin in a subdued manner on the
tory ribs appear about middle of the whorl; the
large ribs are falcoid, all ribs are well developed
on venter. The constrictions trend across the
ribs. The re are 8 constrictions on the last pre·
served whorl and 8 riblets on the space between
the two last constrictions; the intercostal dis·
tance is slightly larger than the width of the
ribs. The umbilical wall is short, vertical, it
 24 F . E TA Y O · S E R N A
rounds abruptly into th e convex sides.
B) Up to 30 rmn [C-223]. One incomplete
specimen septated throughout is available. The
early whorls are smooth. The phragmocone is
sculpt..ued with periodic, widely spaced radial
flared ribs. These prominent ribs begin at the
umbilical margin in a subdued manner; they
trend obliquely on the adumbilical third of the
whorl or form a tenuous adoral concavity. At
the boundaey with the central third of the whorl,
the ribs make a sharp bend and then up to the
ventral zone they fonn an adoral concavity.
Next they cross the venter in a linguiform ado­
ral arc; they are emphasized adventrally . The
section of the whorl is subcircular, slightiy wi·
der than high (H: 10m.m; W: llmm), especially
on the internal whorls; the venter is broadly
rounded grading smoothly into the convex
flanks which grade abruptly into the vertical
umbilical wall.
On the internal mold and also observable
on the exterior of the test there is a constric­
tion just adoral to the flared rib; a flattened
area develops adapical to it. Between two of
the prominent ribs there are numerous fme ribs
(up to 10-14 on venter) which are divided into
two types: 1) slightly prominent, low, fine, top
rounded ribs that begin slightly above the urn·
bilical rim, their trend is that of an elongated S,
like the flared ribs, and 2) fine intercala..T)' ribs
. which rise slightly higher on whorl and only ra­
rely bifurcate from the primary ribs. The inter·
costal space is at least twice as wid e as the ribs,
sometimes greater; the umbilical rim is smoot!":;
with growth the bifurcation or trifurcation of
the ribs seem rather common. On the adoral·
most preserved segment the ribs gather in bund­
les · fanning a low triangular periumbilical thic·
kenning which extends form the center of the
flank to the umbilical rim but this does not
occur on the preceding whorls.
. DIAGNOSIS. This species is characterized
·
by . A) its subrounded whorl section, B) the in·
, creasing number of fine ribs through the onto·
· geny, and C) by its strong, flaring linguiform
ribs;
Miluurements. D Euw · H w D{Euw % D/H %
Holotype - 31 12 7 1 39 ? the ridges. The sulcus immediately adapical to
· Paratype 22 7 1 7 32 ? the flared ribs is usually the most prominent on
. the adumbilical half of the whorl side, but the
.
COMMENTS,: Both specimens here COfilSi·
dered conspecific have to be compared indepel)�
·dently to other species. In regard to the smaller
specimen, the .closest species known to me is the
one figured by Pervinquiere (1907, p. 155, p.
156, Text-Fig. 61, pl. 6, Figs. 21a-b, 22a-c) as
"Puzosia AnglGdei Sayn"; this thick form shows
fine ribs mostly near the constrictions when:as
in my specimens the ribbing is more uniform
and perc.eptible inwards, down to the involution
line; in the latter feature the Colombian species
is more like the holotype (?) figured by Sayn
(1890, p. 173, pL. 2, Fig. 13a, b, c) ; this could
possibly be due to preservation though.
Pervinquiere (ib., p. 156) compared his
specimens to "Puz. Emerici Raspail" though he
remarked on the flattened flanks and the grea­
ter development of the last who;�l in the latter.
In regard to the large specimen (holotype},
this is somewhat similar to "Desmoceras akus·
chaense Anthula" as figured by Jacob (1905, p.
402, pl. 12, Fig. 1; =Valdedorsella Brei&troffer,
1947), but the French species is much more in·
volute and its flared ribs have a peculiar splitting
at that point where the Colombian species have
a centro-lateral acute adoral bend. The African
and European species referred to Valdedorsella
by Breistroffer ( 194 7, p. 60) are much more in·
flated and narrowly umbilicated than the speci·
mens from Colombian here assigned to Melchi­
orites. The constrictions on the latter group of
species is Puzosia-like. This interesting Colom·
bian species also resembles Melchiorites media
Riedel, but the latter lacks the discrete ribbing
on the external part of the whorl and its whorl
section is ogival rather than rounded.
OCCURRENCE. Socota Member Upper
Aptian.
Genus · \Valdedorsella)Breistroffer,
1947.
: Valdedorsella Breistrofer,
.... --1947,
- p. 60. . :
'' --t. -·�..,..-�p:•• ,,_.._.,•..., , ., .::,•,j,..p...r• '""- ,...-.,., ,. · , � · .... •�•-· '• •-·· ----...· -
TYPE SPECIES Desmoceras akuschaense An·
� tula, origtilal designation •
Valdedoryella f71 colQ.mbia� n. sp.
'M:3, Fig. 10; Text-Figs. 3V, 3W. .
Holotype. Santa Ana (U. N. C. M. H. N. C. P. Ht.: '
N.9).
Paratype. Santa "Ana (U.N.C. M. H,. N. C. P. Pt.
N. 7).
A) Up to 35 mm D. [Sta. Anal. 'The sec­
tion of the whorls is suboval, inflated. The test
is ornamented with sinous, flared ribs (27 per
whorl), which correspond to prominent ribs on
the internal mold. There is a constriction on
both sides of the ridge on the internal mold,
they follow the trend of the flared ribs but
attenuate on venter. On the test the constric·
tions are indicated by smooth bands parallel to
reverse is true on the adventral half of the
whorl side. Between two of the ·prominent ribs
there are numerous (:up to 8), rounded ribs,
separated by intercostal spaces one and a half as
wide as the ribs. 'These ribs on venter represent
both primary ribs that originate above the umbi·
Iical rim and secondary ribs that arise from the
primary at about the midpoint of the Oanks. At
the same time the frrst pair of ribs immediately
. adoral to constrictions tend to separate therrisel·
ves from the rest which gather to form a low
bundle at the umbilical rim.
B) Up to 60 mm D [Sta. Ana]. The deve·
lopment of the ornament continues similar to
the previosly described stage except that there
 ZON A'riON OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
seems to be a tendency to decrease the number
of the ribs on venter between the constrictions,
from 8 to 6 or 5. The living chamber has been
preserved as a quarter of a volution.
DIAGNOSIS. This species is characterized
by: A) the decreasing number of the ribs be­
tween contrictions from...S to 5 through growth,
B) the stressed ribs gathered in bundles toward
the umbilical margins, ands C) the inflated
whorl section.
Meuurements D Euw H W DfEuw % D/H %
Holotype 57 22 22a 20a 38 38
Puatype 37 ? ? ? 7 7 Fallot r Fallot, 1920, p. 259, pro. "Puzosia Ma­
COMMENTS. The disposition of the
constriction in this species is close to that des·
cribed by Jacob (1905, p. 402, pl. 12, Fig. 1)
for ''Desmoceras akuschaense Anthula" although
in the French species the flare is divided at the
bend at the midpoint of the flank. This species
resembles "Desmoceras " sparsicosta Boese
(1923, p. 107, pl. 7, fig. 1-3) from Mexico; the
latter, however, possesses intermediate ribs that
bifurcate twice at differente levels on flank
(Boese, ibid., p. 107), whereas in the Colom­
bian species the bifurcation occurs only once;
the ribbing is much more prominent on theMexi­
can species, which in addition, is less inflated
than the Colombian species.
OCCURRENCE. Socota Member. Upper
Aptian.
==� pseu l�c }.!:1!���-�oo:�_
doha er s
�� ��� p a
Pseudohaploceras Hyatt, 1900, p. 570.
TYPE SPECIES. Haploceras lip toviense Zeus­
chner, in Uhlig 1883, original
designation.
� uf!l!jaeJ�ceras leivaense.l}· sp.
P �g:-'1, Text-fig. au, au.
Holotype. VI-10-17 (U. N. C. M. H. N. C. P. Ht.
N. 10).
The inner whorls are apparently smooth.
The section of the second whorl is vaulted.
Flanks diverging from the rather narrow venter,
reach their maximum width near the external
part of the adumbiliclil third of the flank and
then grade abruptly into the vertical umbical
wall. The last whorl has a cross section much
the same but more inflated.
The test of the last preserved whorl is
�ivided into seven trapezoidal areas by flared
nbs which begin irregularly on the adumbilical
thir� of the flank although the growth lines
project obliquely onto the umbilical wall. The
intercostal distance between flaring ribs increa­
ses toward the aperture; between two of the
fiared major ribs there are from 7 to 9 finer
�alcoi� ribs which begin as stressed growth
lines. Independently or in bundles (barely
bulgmg), and which become clearly defined just
adumbilical to mid-venter; they are rounded
25
and are half as wide as Ule intercostal spaces;
some of Ulem widen irregularly on venter. On
the living chamber Ule ribs become very accen·
tuated especially where Uley fonn bundles but
they do not develop real bullae.
DIAGNOSIS. This species in characteri·
zed by: A) its ogival whorl section, B) the in·
creasing width between its flaring ribs or cons­
trictions through grow th , and C) the progressive
stressing of the primary ribs on the flanks.
COMMENTS. There is some resemblance
between this species and "Puzosia Douvillei"
th erom d'Orbigny" H. D ouville (19 16, p. 103,
pl.13, Figs. 1 to 7)'J , especially "in the tendency
of ribs to gather in prominent bundles adumbi·
lically without, however, Conning heavy bullae
as in Uhligella (Jacob, 1907). The African spe·
cies shows more discrete and finer ribbing. Mo·
re similar in appearance, however are several of
the specimens figured by Boese from Mexico as
differen t species of "Desmoceras " in which the
tendency of the ribs to gather in umbilical bun­
dles is much more accentuated ; one of the most
similar species would be "Desmoceras alzatei"
Boese (1923, p. 114, pl. 8 , Fig. 15-18) but the
Mexican species has closer, less distinct ribbing
than the Colombian species.
OCCURRENCE. Paja Formation. (Arci­
llolitas abigarradas;segment E). Upper Aptian .
·-ae-nus -- · · _-r:c��o�a�
� �e-;�c!{.3n g ;;. _;
.· . J
· __
_
TYPE SPECIES. Carloscaceresiceras caceresi n.
sp.
DIAGNOSIS. Moderately evolute conchs,
with low ogival whorl section; the umbilical
wall is vertical and merges abruptly into the
flanks. The ornament consists of sinuous to bi·
concave growth striae gathered in bundles or
weak riblets which sometimes produce on the
internal mold a ribbon-like subdued costation,
particularly visible on venter. There are cons·
trictions preceded by a prominent rounded
swelling visible both on test and the internal
mold but which develop on the adventral
third of the flank only; adorally the constric·
tion is margined by a blunt ridge extending
across the flank down to the umbilical edge, it
is not noticeable on the test. The constrictions
are accentuated progressively with growth,
from almost incospicuos on the early wh·oris to
very wide on the living chamber. The constric·
tions cross normal to the ventral median line.
The apertural border is plain and follows the
path of the growth striae.
The suture line has E shorter than L,
which is deep and very incised.
COMMENTS. . This _new __genus . .difters
_
JI(J.f_fi }3_eudanticeras m_i��!... 19_95) because of
_
its lomr ·mer wider whorl section, its more
evolute whorl and the strengthening of the
internal sculpture during growth. It differs from
Pseudorbulites [ Casey , 1961, p.145, ex Breistro·
26 F. ETAYO · SERNA

ffer, 196 3, p.74, nom. nudum type species Meuurement 0 Euw H W Vw O{Euw O{H W{Vw
UhligellD convergens Jacob, 1907, p.29, pl.2, % % ,Ofo
fig. 25a, b,c} because the latter lacks flanges C- 1 3• 1 26 6 12 10 6 24 48 50
bordering tlie constrictions, has narrow and c- 13· 2 19 4 9 8 4 21 47 50
falcate constrictions and has a funnel-like C- 13· 3 26 6 13 10 5 24 62 50
umbilicus. From Puzosia (Bayle , 1878) it c- 1 3- 4 20 4 10 9 4 20 1>0 44
diffen in the lack of linguifonn projection of C· 13· 5 17 3 9 7 1 17 52 1
the constrictions on the venter and the suture c- 1 3·12 25 6 13 7 1 24 62 1
line with only feebly retracted suspensive lobe. c- 13-13 29 7 14 11 6 24 48 45
c- 13·14 18 4 9 8 4 22 60 50
This genus is named in honor of geologist c- 13·1 fi 28 7 14 10 6 25 60 60
Carlos Caceres. C- 1 3· 1 7 29 7 14 10 6 24 48 60
c- 1a-1s 30 6 16 11 6 20 40 1>4
C- 1 3·19 32 8 16 7 ? 26 50 7
C· 1 3·20 35 9 17 14 7 26 48 50
C· 1 3·22 34 9 16 13a 7 a 26 47 40
C- 1 3·23 37 10 17 13 8 27 47 61

l
Holotype. C - 170 20 (U.N.C.M.H.N.C.P.Ht.
· C• 1 3·26 41 12 18 14 9 29 44 64
N.ll C· 1 3·27 42 11 20 16 9 26 47 60
Paratypes. C· 1 · 1,2,3- (U.N.C.M.H.N.C.P.Pt. C- 1 3-28 42 12 20 16 9 28 47 60
8,9,10). C· 13·29 48 13 21 18 1 1 27 44 60
C- 13-30 48 14 20 17 1 0 29 41 59
A) Up to 20 mm (C-13·1,2 ). The first C- 1 3·33 34 8 16 13 8 23 47 61
whorl after the protoconch is subcircular in s�c· C- 1 3-34 50 17 21 7 1 34 42 ?
tion, the second whorl is slightly higher than C· 13·36 53 17 23 18 9 32 43 50
wide but already has a steep umbilical wall; on C- 1 3·36 68 20 21 ? ? ? 34 7 ?
the third whorl the vaulted shape of the whorl C- 1 3·37 52 21 18 · u 18 36 40 60
section is reached. The test has growth striae C·170· 1 20 6 12 8 4 25 60 50
that extend obliquely from the umbilical rim C-1 70· 2 22 6 12 9 5 23 M 56
and describe a feeble concavity on the umbilical C-170· 3 30 6 16 10 5 20 53 60
third of the flank ; on the central third of the C-170· 4 32 7 16 12 7 22 60 68
flank they fonn an adoral arc and on the adven· C-170· 7 29 7 14 10 6 24 48 60
tral third of the flank they fonn one acute con- C-170· 9 41 10 20 15 8 24 48 63
cavity, next they project mode!ately on the
• -
C-170·11 48 14 21 18 9 29 46 50
venter. C-170·12 46 13 20 17 9 29 44 53
C-170·13 46 12 ,20 17 10 26 ' 43 58
B) Up to 26 mm diameter (C- 13-3]. Up to C-170·14 44 12 20 16 9 27 45 56
this size no special change occurs on the test C·170·16 44 15 19 16 8 34 43 60
ornament except that the umbilicus has already C-170-16 44 13 19 16 9 29 43 66
begun to widen. The growth striae tend to form C-170·17 46 14 20 17 9 30 43 63
irregular bundles which may leave slight bands C-170-1 8 64 18 22 18 1 0 33 40 65
or rib-like impressions on the internal mold; C-170·19 60 14 23 18 12 28 46 65
some striae appear more accentuated than others C-170·20 62 19 26 21 1 1 30 42 62
-
especially on venter. C-170·21 57 17 - 24 20 1 1 31 44 56

� C-1 7D-22 76 27 29 28 1 5 36 38 63
C) Up to 50 mm D (C-13- The specimens C·17D-23 79 31 31 28 13 39 39 46
are septate; weak elongated S·h e constrictions
appear; the remainder of the conch corresponC:.s
to the living chamber. Approaching the begin- COMMENTS. Same as for the genus.
ning of the living chamber the constrictions be-
come progressively more accentuated ; the cons-
r£!
A Peruvian species of ammonites descri·
trictions are preceded adapically by a flange-like bed by Gabb (1881, p. 268, pl. 3 7 , fig.l, 1 as
bulging on the periventral margins and vent.er "A [mmonites� Hyatti", has (ibid.) "su ace
only, but the adoral margin of the constrictions smooth, or on y marked by indistinct lines of
fonn a short vertical ·wall on the internal mold. growth, which bend forward on the inner mar-
Very wide and ·slightly arched constrictions tP.nd gin of the whorl, slightly backwards. on the
to divide the internal mold corresponding to the middle, and then fonnard towards the dorsum .. ' �
living chamber into triangular segments, and on Lisson (1937, p . 23) renamed this species "Des-
these there are htdications of tenuous ribs that mocera$_ Hyattum , and pointed out (ibid p.25)
"

cross the venter, they are narrower than the in-
tercostal spaces and extend down to the umbi-
lical seam. These ribs correspond to bundles of
growth striae on the test. The living chamber
occupies half the length of a whorl. On venter
there are ridge-like thickenings of the test co·
rresponding to the constrictions on the inter·
nal mold, but the thickenings are restricted to
the venter and are not noticeable on the flanks.
DIAGNOSIS. The same as for the genus.
that it lacks constrictions and ribs. These charac-
teristics recall those of the genus Carloscacere-
siceras; however the stratigraphic position of
Gabb's species remains undetermined (Benavi--
deS'-Caceres, 1956, p. 439).
OCCURRENCE. Capotes Member Lower
Albian.
---...
.,e.ar�oscgce!]§Jc!lro� maat!lmLn . sp.
Pl. 0. Fig. 5; Text-figs. 4E; 10.
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
Holotype. C-13-33 (U. N. C. M. H.
N.12).
Very similar to the previously described
species, but in the fanner the elements of the
suture lin e are higher and the saddles are more
finely zigzagged-lncised;"'in the preceding spe­
cies the adumbilical and central thirds of the
flank are Oat, the adventral third of the flank is
slightly beveled. In Car/oscaceresiceras monteroi
n. sp. the flanks are faintly convex and further­
more they show more indication of ribbing on
the internal mold .
OCCURRENCE. Capotes Member-Lower
Albian.
This species is named in honor of geologist
Juan Montero:
�eslo�ace
V1
�ceresjcerns C? J £hjmuense, (Bena·
res)
Pl. 11, fig. 9; Text-figs. 4J, 4K.
Hypotype. VL.33a. (U.N.C.M.H.N.C.P. Hy. N.3).
1956 Desmoceras chimuense Benavides-Caceres,
p. 438, pl. 4 1, figs. 5·8 .
1964 cf. Desmoceras chimuense Benavides·Ca·
ceres, Etayo-Sema p. 37, fig. 9.
One internal mold in black calcite . The in,
nennost whorl observed is higher than wide in
section, it has flat flanks which round abruptly
into the short but vertical umbilical wall. The
ventral zone is not discernible ; the visible part
of the flanks shows fine growth striae which
form .a shallow arc from the umbilical seam to
the involution line. The last whorl preserved
has flattened sides which feebly diverge from
th e evenly arched venter; the section reaches its
m!D4mum width about the umbilical third of
the who.rl where the sides round abruptly into
the yertical umbilical wall. On the surface of
th is mold, vague indications of constrictions are
..
Present but . the exact trend of them can not be
aec.u:rately detennined because of the state of
pre�rvation.
M.�asurementl D Euw H W D{Euw % D/H %
VL"33;. at 7� 19 33 30 25 .
.. . COMMENTS. In suture line and whorl
$ection this specimen resembles that described
by Benavides�Caceres as Desmoceras chimuense
B�navides·C�ce s, 1956, p. 43�, pl. 41, fig. 5·
�
8, '!'he _spec1es IS referred tentatively to Carlos·
cQcereslceras. .
·. ·.· OCCURRENCE . Upper San Gil
.
Forma·
tion, r.,evel 8. Middle Albian.
[�[����..�:.'fCorteziceraij���--g����.:. ... :
!YP· .· �. S Jit�IES. Corteziceras corte
zi n.g., n.sp.
- ZvJN
· · ctfc.ff.r,
··
.
� -l'1f7t
�
. • . DIAGNOSIS. C nch evolute, of mode
·
siZe , The early whorls are finely striate rate
su.,sequent whorls are divided into segmen d the
ts by
27
N. C. P. Ht. flaring and between them accentuated ribs
ribs
occur; the latter may bifurcate at different
levels on the flank. The more prolonged the
initial smooth stage, the less strong the orna­
ment becomes; all the intermediate ribs are
prominent, however. The ribs tend to gather in
bundles which reach the umbilical rim tangen­
tially but they do not develop any umbilical
tubercles. There are two constrictions especially
noticeable on the internal mold, parallelling
each flaring rib, of these the adapical constric·
tion is accentuated in early ontogentic stages
but tends to weaken with growth to th·e benefit
of the adoral constriction.
COMMENTS. Corteziceras differs from
Uhligella Jacob 1907 (Type species: D. clansa­
yesense Kilian) because it lacks nodes or swe·
llings on the umbilical margin. It differs from
Zurchere/la Casey (1954, type species: D. zuer·
cheri Jacob) because of the weakness of the rib·
bing and greater involution of the latter. I
.
This genus is named in honor of Santiago
Cortez, Colombian paleontologist ·
�{r;,fLr:zs cortezj,.n. sp.
lfC3�-ng.2, Text-1ig. 4C.
.
Holotype. Santa Ana, A.. P. G. · · (U.N.C;M.�.
N.C.P. Ht. N. 13)
? 1938 Uh/igella ziircheri Jacob &Tobl., Riedel,
p.20, pl. 4, figs. 12, 13, pl. 14, f.4.
? 1957 Uhligella ziirchen' Jacob 8. Tobler;
Biirgl, p. 135, pl. 8, fig.S.
The conch is discoidal, moderately umbi."
Heated. The inner whods are smooth, they are
covered only by sigmoidal growth striae; the
latter are concave ado rally on the umbili�al wall,
trend obliquely on the adumbilical third of the
flank, then tum feebly adapically describing a
weak arc on the central third of the flank. On
the adventral third of the Dank they trend su­
bradially and describe an adoral concavity and
finally they barely project onto the venter.
Fascicules of growth lines near the middle
of the flanks pass into ribs that become conspi·
44 cuous on the venter; at 25 mm diameter they
are already recognizeable. Periodically one of
these ribs becomes more prominent, especially
on venter, but it shows the same trend than the
other ribs; such flange-like ribs are represented
on the internal mold by a rib also, but, imme­
diately adoral to it there is a deep furrow that
runs parallel to the rib but which is represented
on the exterior by a flattened band.
Adapical to the rib is another less deep
sulcus on the internal mold corresponding to a
flattened band similar to the adoral one. Be·
'
tween two of these prominent ribs there are
usually 5 finer but also stressed ribs of which
those two occupying a more central position
usually extend down to about the adumbilical
third of the flank, whereas the other two join the
latter at about the central third of the flank;
28 F. ETA Y O - SERNA
this is particularly noticeable on the exterior of
the shell .
The whorl section is higher than wide and
is widest toward the adumbilical margin; the
venter is arched, the umbilical wall is subvertical
and rounds smoothly into the slightly convex
Danks. On the internal mold the ribs do not give
the impression of being as inclined as they are
on the test. The intercostal spaces are twice as
wide as the ribs.
DIAGNOSIS. This species is characterized
by A) the progressive ontogenetic development
of the orname tation on which the stres� of the
ribbing accentuates with growth from the ven·
tral third on the early whorl towards the adum­
bilical third on later whorls, B) the triangular
segments of the test with four to five ribs on
venter, and C) by its relatively evolute conch.
Meuurementll . D Euw H W E/Euw % D/H%
Sta.Ana.A.P.G.No. 64 20 20 18 40 40
COMMENTS. With the name of" Uhligella
zurcheri Jac. It Tob.'' Riedel described (1938, p.
20, pl. 4, fig. 13, U, pl. 14, fig. 4) a species of
ammonites as having two or three weak ribs
between two prominent ribs, however on the
illustration it is possible to count up to 5 of the
"weak ribs". This is proven by direct observa­
tion of Riedel's original specimen (S.G.N.C.P.
�o.OS1). Riedel's specimen came from the sa­
me general area as my specimens and I consider
, them tentatively conspecific.
On the specimen figured by Jacob (1906,
p . 9, pl. 2 , fig. 1a, 1b) the more numerous prin­
cipal ribs aie slightly sinuous only and the trian­
gular surfaces they circumscribe have the shape
of a narrow truncated pyramid, whereas in my
specimens, because of the exaggerated inclina·
tion of the principal ribs this surface has the sha­
pe of an inequilateral trapezoid , furthermore the
arrangement of the ribs on Jacob's specimen is
as if the central intercalatory ribs were branching
about the .!Piddle of the flank; the ribbing is
uniformly thin in Jacob's species.
The specimen figured by Biirgl ( 1 9 5 7 , pl.
8 , fig. Sa, Sb) corresponds to Riedel's original
of plate 4, fig. 12, 13, but the picture is only
8{10 of the natural size.
The Mexican "Uhligella" reesidei Hump·
hrey (1949, p. 152, pl. 18, fig. 3,4), is a much
smaller and·flattened species on which the streng·
thening of the ribs occurs earlier than in the Co­
lombian species;they share,however, the strength
of the ribbing and the tendency to produce a
wide umbilicus. Casey's suggestion of generic
status for this species and similar Mexican species
( 1949, p . 336; 1961, p.161} has, in my opinion,
good biogeographic support. A similar observa­
tion was made by Boese (1923, p. 109) when
describing his "Desmoceras " from Mexico
(trans) : "Among the European forms. . . . all
are distinguished by their lack of prominent
intermediate ribs".
OCCURRENCE. Socota Member. Upper
Aptian.
Corteziceras cortezi n. sp.
fMorph I]
Pl. 3, Fig. 7.
Paratype . (U.N .C.M.H.N.C.P.Pt.ll)
Only one specimen is available. The test is
thick, the section of the whorl is vaulted with
sides diverging from the arched venter and gain­
ing their maximum width at the umbilical rim.
The early visible whorls are smooth except for
barely perceptible fine growth lines. The last
preserved whorl is completely septate; the orna·
ment on this whorl consists of primary ribs
which gather together about the umbilical rim,
the secondary ribs bifurcate from the primary
ones at about the middle of the flank but the
adoralmost of the secondary ribs may itself bi­
furcate at U1e adventral third of the flank; these
groups of ribs occur between two prominent ribs;
the latter ribs cross the flanks in an elongate S.
On the internal mold there is a channel like
constriction, twice as wide as the flaring rib
which develops immediately adapical to the lat­
ter; this constriction corresponds either to a
constriction or to a flattened band on the test.
The umbilical margin is rounded and the umbi·
lical wall is subvertical on the last preserved
whorl.
NOTE : This morph is notable for A} its rather
low section, B) the frequent presence
of constriction (9 per whorl ?), C) the
fasciculation of the primary ribs around
the umbilicus which, however, do not
fonn nodes, and D) its quick accen­
tuation of the ornamentation and short
smooth earlier stage.
Meuurementlr. D Euw H W DJEuw % D/H%
Sta.Ana.A.P.G.No. 41 15 16 14 36 36
COMMENTS. This "morph" bears some
resemblance to "Uh ligella reesidei" Humphrey
( 1945, p. 153, pl. 18,Fig. 3, 4), but the Mexican
species is more compressed, the adumbilical and
central thirds of the flank are flattened and the
bundles of ribs are not as strong and persistent
as in the Colombian "morph". The Mexican
species has attenuated constrictions which are
parallelled by more strongly developed ridge-like
ribs ( : flanges) contrary to the Colombian
"morph". This "morph" differs from the bolo·
type because of its more numerous constrictions
and more irregular branching p attern of the ribs.
OCCURRENCE. Socota Member · Upper
Aptian.
r1ezife� latecostata.
WI.
(Riedel)
, Ftg. 6, Text-Ftg. 4F.
1\ '
1938 Uh ligella latecostata Riedel, p.2J, pl.5,
figs. 1-3 .
19f.7 Uhligella latecostata Riedel,Biirgl, p. 1 35.
 ZONA'l10N OF THE CRETACEOUS OF
1964 Uhllgello latecostll tll Biedei, Etayo-Sema,
p.lll.
1966 Uhligell4 14tecostatlJ Riede�BUrgl ln Btlrgl
and Campbell.
Holotype. Rledel's- l!peeimen
Hypotype. (U.N.C.M.H.N.C.P. Hy. N.4).
Type LocalJty. Hacienda Magdalena, Viota
(Cundlnamarca).
Three tragments of Internal molds of the
umple 0.256 are referred to thla species.
The section of the whorl has an arched venter
which rounds evenly Into the flattened flank:a,
the latter feebly diverge down to the adumbW­
cal third of the flank where they round rather
abruptly Into the abort aubvertlcal umbillcal
wall.
The omament consists of periodic cons­
trictions that are wide , channel-like, begin tan­
gentlal to the umbilical rim, and near the boun.
dary between the adventral and middle thirds
of the flank they describe an oblique feeble arc
which Is followed on the adventral third by a
shallow adoral concavity that directly cro111ea
the venter. Adorally to the constriction there.la
a rib that becomes very prominent on the ven­
ter but which fades out before reaching the
adunibWcal third of the flank or just extendfJ
Into it.
. Between the two constrictions there are
four visible ribs on venter; they correspond to
the branching of two main ribs, the adapical
one that begins at the umbilical rim, trends
almost radial on the adumbillc al third of the
whorl side, then abruptly bends adapically on
the central third of the flank but on the !ldven­
tral third it trends radially again; at the "point o f
Its first bend a second branch arises and descri­
bes a feebiB concave arc across the flank, and
then across the venter. The second major rib
begins at the same poin t as the first main fu­
rrow, but it trends obliquely describing a feeble
arc on. the central third of the flank, then It
abruptly forms an adoral concavity and rmally
it croaaes the ven ter straight; at the bend, a se­
con d, ali ghtly flexuous secondary rib branch
· Description of holotype. The. conch is in·
�peazs.
OCCURRENCE. Beds above the Capotes
.Member and below the Socoti Member. Upper
Aptian.
Cortezicel'll8 (?) n. sp. 7
Pl. 3 fig. 6; Text-Fig. 3 &
!let specimen C - 513
Only one spectrnen is available. The ada­
plcal portion of the last whorl prese!Ved has
.
flankS that descend abruptly into the umbilicus
and round progressively towan:ls veneer with
the maximum thlcknesa about the mid�le of the
flank, . then they converge toward the narrow
venter forming a low ogive . TowiU'd the adapical
en� of the last whorl, the section tend3 to round
urufonnly.
CENTR AL COLOMBIA BY AMMONITES 29
On the· venter the teat ha periodically
thickened, rounded and llnguilonn flanges
that extend, progmufvely attenuated, to the ·
umbilicus In a falcold manner. The early portion
oi the teat Is covered with tine falcold to bicon­
cave striae with the external adoral concavity
occupying approximately the central and adven­
tral thirds oi the flank, and then project onto
the venter; toward . the adapertural thlrd .of the
lut whorl, tine rounded ribs that are slightly
narrower than the Intercostal spaces become
well defined especially on the central and adven­
tral thirds of the flanks. The specimen Is com- .
pletely septate. ·
NOTE : This species Is characterized by: A) ita
inflated, low ogival whorl section, B)
the tenuous, fine biconcave ribs, and
C) the externally accentuated flanges
which correspond to the adapical ridge
of constrictions on the intemal mold.
M...unment.. D Euw. H W DfEuw"f0 DfH %
c-�13 32 a u 13 211 44
COMMENTS. This species resemblesPseu­
dorbulites convergens (Jaco b) (1907, p.2 9 , pl.2,
fig. 26a, b, c) but differs In the position of the
maximum Inbation which is more toward center
of the whorl ln the Colomb ian species. This spe­
cies also shows great resemblance to the inner
whori:J of Corteziceras cortezi n. sp., however
has a relatively more compressed whorl section
and does not show the biconcave growth striae.
Genus f Pseudosaynell4 ) sp ath , 192 �·
�,....._., _
-
_ -
�·-· -
... ... .,.,.... ""
-
. · . ·
-· -
"" -
-
" ... ' .. ,- .. , . -
··
- · ···
Spath, 1923, p. 66.
- · ··
Pseudosaynell4
··
· --
··
TYPE SPECIES. Ammonites bicuroatus
Michelin, original daslgnntion.
eyeudosg�ne�a ra�fiml!lli n. sp.
.3, Frg; 1; ext- J.g. 4N, 4.0.
P
Holotype. SV.-28 (U.N.C.M.H.N.C.P.Ht.N.14).
Paratype. (U.N.C.M.H.N.C.P.Pt.N.12)
Type loc ality. Lorna San Vicente, Villa de Leiva
(Boyaca).
volu l"e ' discoidal, with sharp venter; the section
of the whorl is lanceolate: the maximum width
is on the flattened peri-umbilical region from
which the flanks slope gently up to the sharp
ven ter- the umbilical rim is blunt, the umbilical
w all is' alm ost vertical, it changes its inclin ation
at depth; when the test is preserved the umbili­
cal rim appears acute.
On a small piece o f test preserved around
the umbilicus occur very rme growth striae. On
the internal mold are low rounded ribs that be­
gin faintly near the umbilical m� trend
radially on the smooth adumblllcal third or the
flank, make an acute bend on part of the central
third and then describe a wide, cre9Centlc adoral
concavity and project sligtly onto the venter.
From the central bend up to the beginning
of the sharp venter the ribs become broad, low
30
and bifurcate; interc$tory bifurcating ribs also
appear. The distance between the branches of
the wne rib is narrow, but the distance be·
tween each two couples of ribs is usually wider
than a single rib. The ribs converge toward the
umbilical rim; they crosa the acute venter as
fiat, low belts.
One quarter of the last preserved whorl
corresponds to the living chamber. Towards the
end of the phragmocone the external ornament
attenuates but it Is strengthened on the living
chamber. The acute venter is beaded by the
crossing of the Battened ribs; on the internal
mold there are wide constrictions which follow
the trend of the ribs.
DIAGNOSIS. This spedes is chlll"&cteri·
zed by: A) its disk-like appeaJence combined
with the sigmoidal bifurcating ribs, and E) the
change In stress of its ornament during growth.
Meumementa. D Euw H W E/Euw'% DfH"'o
SV · 28 87 8 37 177 12
La Ye 24 4 11 1 n
c . 223:1:
•
27 7 7 7 7
COMMENTS. A comparable species was
described by Imlay (1944 , p.278, tex-fig. 2-4,
(i ; but from the Ulustrations ofPse Zl
plates 42,43)
dosaynella mbriata Imlay, the Colombian spe·
eles has more sigmoidal and leas regularly spaced
ribs; furthennore, the holotype of his species
shows a definite change In the ornamentation in
the latter stage of the ontogeny; likewise the su·
ture line of the Colombian species hu L divided
by a massive tnushroom-likc satldle that with
growth renders this lobe bifid, but this lobe is
ln comparison slim, moss..J.ike in Imlay's species
(see fig. 4d, p.279); it must be pointed out that
the diameters are not similar, however, the Co·
lombian species seems to be smaller altogether.
Pseudosaynella walcotti (Hill) [1888, p.
128, pl. 1, fig.1; Scott, 1940, p.995, pl.56, fig.
1,2] Is a more evolute species and seems to have
leas numerous ribs, but these are stronger
throughout than on the Colombian species.
Of the European species described by
Sarasln (1893), PseudosayneUa bicuroata (Mi·
chelln) shows a slm.Uarlty In the type of ribbing,
but the Lower Aptian French species has its
Danks more fiattened; the suture line at firr.t
alght looks similar, however at similar diameter��
the French species has a more massive E/L
saddle. Collet (1�.22, p.17) recorded the French
species from Venezuela, but this record hss
been questioned by Casey (1961, p.172).
OCCURRENCE. Socot.i Member. Upper
Aptian. This species is named In honor of Dr.
Ralph Imlay.
- Family PACHYDISCIDAE
�""�m
�eiiuil w�lceras)
Spa'fll;
19 39
Lewesiceras Spath, 1939, p. 296.
TYPE SPECIES. Ammonites Peramplua J.
F. ETAYO · SERNA
Sowerby, original designation.
M.lJ.l�#t;MQ!_f[l�llmli n . sp .
..Pl.T3";"fig.�. 4D.
Holotype. C·310 (U.N.C.M.H.N.C.P.Ht.N.15).
Paratype. C-310 (U.N.C.M.H.N.C.P.Pt.N.13 ).
Description, largely after C-310. The
conch is moderately evolute; on the penultlma·
te whorl and the adapical portion of the last
whorl the ornament consists of widely spaced
fianges or ridges which begin at the umbilical
rim and trend radially up to a point near center
of the flank from which point they trend pror·
siradiate and crosa the venter forming an acute
chevron-like projection; this ridge is paralleled
adapically by a aha Dow constriction which
reaches the umbilical margin; the second,
deeper but narrower constriction pamllels the
ridge adorally, it fades out near the adumbilical
third of the flank. A leu prominent ridge rima
thiG second constriction.
66
64 Between each two adjacent principal
7 ridges there is a large blunt bullate umbilical
swelling; on the adapical third of the length of
the last preserved whorl this swelling gives rise
to a low rib which dies out about the middle of
· the flank or �ontlnues almost imperceptibly
across the venter. On the adoral half of the
whorl however, this tubercle gives rise to one or
two rounded and prominent n'bc which describe
an are on the flanks and projecb; onto venter in
the same manner as the constriction. One
intercalary rib which does not reach the umblll·
cus may occur.
The intercostal spaces are wider than the
ribs, they are flattened and shallow between the
ribs which arise from the adumbilical tubercles
but narrower and deep near the constrictions.
The section of the whorl is low, ogtval to
renifonn, the umbilical rim rounds abruptly
into the vertical wall ; there are seven prominent
umbillcal tubercles on the last whorl.
DIAGNOSIS. This species is characterized
by: A) the long enduring stage with constric­
tions and intervening umbilical tubercles, B) the
pairs of rlbs i!Iising from the umbilical tubercles
later in the ontogeny. Presumably only the ju·
venlle stages are known as yet.
Mt�uureruenta. D Euw H w D/Euw% D /H "k
C·31 0 211 . - 1 1 9 11 "" 36
C· 310 14 li li 7 36 36
COMMENTS. In th e illustrated species of
LewesicerG 1n the "Treatise" {Wright, 1957,
L-378, fig. 494, la-1b) the principal ribs bonn·
ding the constrictions bear prominent tubercles ·
but•the intercalary ribs appears to begin on the
venter and advanCE toward the umbUlcus with·
out any tubercles, completely the reverse o! tbe
Colombian species. Houaa {1965) ball diacuased
at length the problems wlth respect to the type
specie& of IK.wesiceras.
 ZONATION OF
OCCURRENCE . La
'1\lronian.
This speciE!& is named In honor of the
geologist Carlos tJ1loa, Instituto Nacional de In·
vestlgaciones Geologico·Mlneras.
,,
FamilY
! Genus
'
- [Oc
m er,
fJier#
��H�EI�RATIDAE
.:...: _.,..,•• �- .�.:. :..: _;;>:(. ·'-"-"'""""'"'...,,.,...,..,.......
__
Trochleiceras 'Fallo t and Termier,1923, p.
TYPE SPECIES. Trochleiceras balearense
Dot and Tennier, monotypy.
h�jc\�(7i�h�ce .iUlivW,i n.sp.
I. , _g. , ext· g. f, ftil
� .
Holotype. C-215·1 (U.N.C.M.H.N.C.P.Ht.N.16}.
Paratype. C-215-2 (U.N.C.M.H.N.C.P. Pt. N.14).
Small, relatively evolute conchs (C-215·1,2).
They have a well defined ventral median groove,
paralleled by rounded ridges, In a pulley fashion.
The initial whorls of the phragmocone are
smooth , covered only by sinuous lirae which
trend as an elongated s with stress on the exter·
nal concavity. The lirae cross the venter nonna­
lly to the ventral median line. Bundles of striae
form periodic narrow, low ribs which now and
then are raised.
On the living chamber and adjoining part
of the phragmocone, compact bundles of lirae
form a kind of stem on the inner half of the
whorls, about middle of the flank this stem
splits into five rme, rounded to flattened riblets
that extend to the ventral median line. The
ridges that paralle l the ventral median grove
appear minutely denticulate because of inter·
section with the riblets. One tubercle-like
outgrowth on the right side of the holtype is
interpreted as a adventitious growth to cover an
extraneous object.
DIAGNOSIS. This species is characterized
by: A) Its small conch, B) its rounded subrec­
tangular section, and C) the change in ornament
through growth, showing only sporadically
stressed lirae on the early whorls, and later by
riblets issued in groups of five from an adumbi·
lical principal stem.
Meuurement.. D Euw
C·2 1 &·1 14
COMMENTS. This species can be diffe·
rentiated from the type species of the genus:
Trochleiceras balearense Fallot and Termier
(see Fallot and Tennier, 1923, p.74, pl. 6, figs.
7-9, text-fig. 32; Collignon, Magneticerc.s mag­
neti C., 1950, p.48, p1.6, figs. 1, 1a, 1b text­
�g.4;Wiedmann, 1962, p.323, pl.18, fig. 2, text­
ug. 3J, because the species from the Balearics
d� not show any change in the ornamentation
duf!ng ontogen and only prominent single
"pnmary ribletsr,' occur up to the living cham·
ber.
THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES 31
Frontera Fonnatlon. OCCURRENCE. Lower Albian.
This species is named in honor of Pro­
fessor Manuel Juiivert.
...._
n o /i Sub g�Jl� \...�
-a-ma...o._ft_a_ic_e_ra__,
�_..p s)n. subg.
-= Tru diflv:wo..')
3.
t""ilfof'iruPre!· ·
TYPE SPECIES. Trochleiceras (Jaumaportai·
ceras),ho(fste t.W,n :n. sp.
--:;;r..........,.... .. ....___ DIAGNOSIS. Jaumaportaiceras differs
_
. ..........._ .- �,,
74. from Trochleiceras by the absence of a ventral
sulcus or only a very weak indication of it on
Fa­ the inner whorls, by the presence of ventral
bullae, and by a more lacerate suture llne.
This subgenus is named in honor of
Professor Jauma de Porta.
Troch$ras (Jauma,eortaiceraf!.2.
Jj07fffeJt.ifii. sp .-
PI.10, fig. 4; Text-figs. 4G, 4H, 4L.
Holotype. C-215·5 (U.N.C.M.H.N.C.P.Ht.N17).
Paratype. C-215· (U.N.C.M.H.N.C,P.Pt. N15, 16,
17) .
A) Up to 15 mm D [C-215·3,4]. The
conch is small, flattened, evolute; the section
of the whorl is rounded subquadrate; the umbi·
lical wall is vertical and rounds evenly into the
flattened to slightly convex flanks, which in
tum merge smoothly into the flattened venter.
On the adapical segment of the specimen
studied (mostly decorticated), a shallow broad
groove is present on the internal mold. At this
stage of development, when the test is preser·
ved it appears smooth, covered only by rme
growth striae, occassionally one of them is more
prominent. The surface of the shell has low
broad undulations.
The living chamber occupies half a
whorl, and whose beginning in this case
is marked by the close approximation of the
last two sutures which earlier in the phrag­
mocone are distant from each other. Low
:ounded ribs are developed irregularly on the .
living chamber; they flatten on the venter and
form transversely rounded tubercles on each
ventral margin. Where the shell is preserved
(C-215-4), it shows broad, raised bundles of
lime separated by narrower but well defined
intercostal spaces; theses correspond to the
bulges on the internal mold.
H W DJEuw% D/H %
6 6 6 . 43 43 B) Up to 20 mm D [C-215·51. At the dia·
meter of the holotype, the intemaf mold of the
living chamber shows relatively accentuated ven·
trai bullae on conspicuous rounded ribs; the lat­
ter begin on the external part of the umbilical
wall and trend prorsiradiately on the adumblli·
cal third of the Oank, then they abruptly chan·
ge trend to radially or slightly inclined adapica·
fly, they then describe a narrow concave arc on
the adventral third of the flank; they cross
normal to the ventral median line; a few of the
ribs are feebly depressed on mid-venter; the
principal ribs bifurcate twice: first about the
boundary between the adumbilical and central
32 F. ETA Y O · SERNA

thirds of the flank where a subtriangular thick· This species is named in honor of Profes­
ening is usually fanned, and second on the ad· sor Robert Hoffstetter.
ventral third where no swelling is fanned; the
adoral branch is usually , though not alway£ the
strongest one. The strength of the ribs is not
uniform and the distance between them likewi·
se varies. Adorally to the principal ribs are de­ Cheloniceras Hyatt,1903, footnote p. 101
preBSed bands that follow the trend of the
ribs. TYPE SPECIES. Ammonites comuelianus d'Or·
bigny subsequent designation,
I.C.Z.N., opinion 428, 1956.
�
•

During growth the whorl section changes

from subrenifonn to subquadrate with slightly E
arched venter, likewise the innennost whorls Subgenus { pichelonicen!JCasey, 1954.
are smooth but bear broad flexuous swellings;
in the later stages of growth, and mostly on the Epicheloniceras Casey, 1954, p. 113
living chamber the secondary ribs branch from
primary ones at different levels. There is a TYPE SPECIES. Douvilleiceras tschemyschewi
feeble ventral sulcus on the innennost whorls Sinzow, original designation.
which disappears or is very faint on later stages
of growth; appearance of ventral, rounded �eloniceras (Epichel,gTJicets§)
bullae-fonn tubercles seem to characterize the
beginning of the adult stage.
�aTl!!i
'":�; H
� n. sp .
g. ;Text-Fig. 4W.

The conch appears to consist of in terloc· 1 968 Cheloniceras aff. Ch. juv. ex. aff. comue­
king segments;- Now and then the primary ribs -lianum d'Orb, Riedel, Etayo-Sema 1968,
(?) on the inner whorls may be raised as thin Fig. 3, Table I.
flanges which begin slightly above the umbilical
margin and attain their maximun flaring at the Holotype. VL12·2 (U.N.C.M.H.N,C.P. Ht. N.18)
middle ct' the tlank; parallel and adapical to
them, a wide shall ow constriction develops. One slightly defonned internal mold is at
hand. The adapical portion of the last preserved
DIAGNO SIS. The characteristics of the whorl has prominent low rounded primary ribs
species are the same as for the subgenus. The which begin near the umbilical seam; they bear
variation of the morphologic characteristics is a tiny spinifonn tubercle at the umbilical rim,
significant: the ribs mny vary in strength, like­ and on middle of the flank a very prominent
wise the width and depth of the constrictions spinose tubercle at which point the primary rib
are variable but not their .;hape; on same phrag­ trifurcates, the branches cross the venter, slightly
mocone both raised and low ribs may develop; depressed at the median line and fonn rounded
some ribs are broad and low and arise in pairs shoulders; on the venter the ribs bend slightly
from the periventral margin. forward in a broad a.c; of the three branches
the median one tends to be the most prominent;
Meuuzemente D Euw H W D fEuw% D/H% bet.ween two tuberculate ribs there are two in·
tercalary ribs of which the adapical one begins
C-211)-3 13 II 6 I) 38 38 at the middle or the flank as if bifurcating from
0·210-4 16 6 6 8 37 37 the subsequent tubercle. A succession of closely
0·2111•6 22 10 8 8 411 38 spaced trifurcating ribs follows the preceeding.
C-216� 20 ll 7 7 46 36 Two intercalary ribs reappear about two thirds
C·2l6·7 16 7 6 6 44 37 of length of the last whorl.
C·21fi-8 16 7 6 6 47 40
C·2lli-ll 16 7 6 6 44 37 Of the trifurcating ribs only the central
C·2l !o·10 1 3 7 7 ? 1 1 and adapical branches are depressed on venter,
0·216·12''18 7 8 6 44 37 they are nearly equally strong, and are more
prominent than the other ribs. The intercostal
COMMENTS. Trochleiceras ambanjaben­ spaCPII vary from narrower to as wide as the ribs
se Collignon (1962,
p. 37, 130,
pl. fig. 985) on venter; on the flanks they are deep and
from the Upper Aptian (Gargasian) of Madagas­ channeled. The large lateral spinifonn tubercles
car at equiv&lent diameters (approx. mrn),
9 are pointed adapically, whereas the periumbUI·
lacks the development of the periventral bullae, cal ones are mammiform. Tne section of the
it has a more accentuated, even thou�tb weak, whorl at the apertural end of this specimen is
ventral mediari groove, the section of the whorl inflated renifonn. The conch has 37 ribs on the
is subrectangular rather than subquadrangular venter of the last preserved whorl.
aa in the Colombian sp.e cies, furtherm o re
instead of low broad ribs on the early whorls, DIAGNOSIS. This species is characterized
the Madaga.o;can species has numerous rme rl· by: A) its highly developed lateral tubercles
blets. The Colombian species diffen from the •from which ribs trifurcate, B) its thick renlfonn
type species of the genus by the type of ribbing, intercostal whorl section, and C) the sporadic
more complicated suture line and nearly absent occurrence of intercalary ribs.
ventral groove. " •";,,
Meuuremenu. D Euw H W DfEuw% D!H"k
OCCURRENCE. Lower Albian. VI-u-2 s7 13 a a 31> 40
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
COMMENTS. This species resembles the
inner whoda of Cheloniceraa (E.) debile Casey
(1962, p. 244, pl. 37, r.a, 7, Text·Fig. 85, 86b
and especially ftgums 86a. 86b) but on the
EugUah species the central one of the branching
ribs Is the 1tron ges t, turthennore It has "groups
of tblee to four intermediary ribs at 20 mm
diameter ,. (Casey, ibid..,p. 244) whereu in the
Colombian species the intermediary ribs ue
almost abeaent.
Them Is also a resemblance between the
Colombian specimen and the lectotype of the
type species of the subgenus (Sinzow, 1906, pl.
2 Figs. lla-c; reproduced in Casey, 1962, p.
2S7, Text-Fig. 82a, b, c); on the Russian fonn
the ventral me dian sulcus Is deep and the ven·
tra1 tubercles prominent, opposite to the sltua·
tlon � the Colombian species.
OCCURRENCE. Paja Fonnation (Arcillo·
Utu abigarradu, segment E), and base of Ar·
cllloUtu con nOdulos huecos. Upper Aptian.
This SJ!ecies is named in honor of Mrs. El·
··
vira de Borru, of Villa de Leiva.
. Cheloniceras (Ef!.ic�!JonJ2e�.L camachoi
n. ap.
PI. 2, Fig. 3; Text-Fig. 4T.
1928 Cheloniceras stoliczkanum Gabb,
Buse, p.14 1, pl.S, Fig. 2. Text-Fig.18.
non 1936
' Ch.[eloniceras] Stoliczkanum Gabb,
Basse , p. 88.
'l 1936 Cheloniceras ct. Stoliczkanum Gabb
sp. em. Basse , Breistroffer, p. 156.
1938 Cheloniceras stoliczkanum Gabb, An·
derson , P.. · 176, pars., only reference
to Basse s illustrated specimen.
1964 Cheloniceras stoliczkanum (Gabb ),
Etayo-Sema, pars., p. 118.
1968 Cheloniceras (Epicheloniceras)stolicz­
kanu m sensu Basse (non Gabb ), Eta·
yo-8ema, p.28, Fig. 3, Table I.
1970 Cheloniceras (Epich. [eloniceras V sto·
liczkanum, Wiedmann, p. 484, ex re·
view of Etayo-Sema's 1968 paper.
Holotype. (U.N.C.M.H.N.C.P.Ht.N.19).
Paratype.
.. (U.N.C.M.H.N.C.P.Pt.N.18).
The ad apical portion of the last preserved
whod on the holotype (VI-10-17) shows strong
� ular ribs which bear a tiny mammtfcnn tu·
bercle at the umbilical margin, a conical tuber·
cle abo�t the middle of the flank and a promi·
nent triangular transversely elongated �:reriven­
tral tubercle; on venter the intercostal :iistance
1
1•1 more than twice the width of the ribs but it is
.�.· th.an. two times on the flanks; in gene r� the
�..�&tercostal spaces Qre shallow. The ribs are tnick
. •
1
.
an.d rounded.
33
Between two prominent ribs a nearly
imperceptible Intercalary rib appean, tt Ia
feebly depressed along the ventral median line;
with the tncreuing size the strength of both
ribs and tuberculation rapidly increases, and
approaching the preserved apertural region the
Intercostal spaces are very deep and channel
fonn; at this stage of growth the Intercalary rib
approximates to the periumbUlcal tubercle of
the preceding rib u if arising from �t. ·
The periventral tubercles give the impres­
sion of being slightly bent posteriorly. There
are 18 well defined ribs on the last whorl. The
Jatenl tubercles were long and spinose as shown
by depressions on the umbilical wall of the
holotype and the paratype.
DIAGNOSIS. This species is characterized
by: A) the rapid on to genetic strengthening of Ita
ornament contemporaneously with a slow in·
crease of the umbilical diameter, B) ita subrec­
tangular intercostal whorl section, with greatest
width about the umbilical rim', and C) the regu­
lar occurence of only one intercalary rib appea­
ring near the umbilical rim .
Meuuremenw. D Euw H W DfEuw % DJH %
VI · 10 • 17 47 21 19 28 411· 40
T.I.P. GuaduQ 61 27 ? ? . 44 1
COMMENTS. This species differs fonn
Cheloniceras stoliczkanum (Gabb)(1869) by not
having the very depre!l&ed whorl section during
ita development and the width-length ratio not
remaining the same throughout as on the bolo·
type of the California species (Acad. Nat. Sci.
Phil., no. 4 786). The ventral nodes on Gabb's
species are broadly rounded, lobe-like, but in
the Colombian species their height is greater
than their width (in correlation with the com·
pression of the section?); the periumbilical tu·
bercles are represented by feeble bullae through
the ontogeny on Gabb's species but they are co­
nical and prominent early in the ontogeny of
the Colombian species.
OCCURRENCE. Socota Member. Upper
Aptian. Paja Fonnation (Arcillolitu ablgarradas,
Segment E). Upper Aptian.
This species Is named in honor of geolo­
gist Ricardo Camacho.
1906 Douuilleiceras Stoliczkanum, Gabb, R.
Douville, p. 145 (pars), only pl. 1, Fig. 1,
la.
1925 Douuilleicera.s Amazonarum Burckhardt,
p. 26.
1938 Douvilleiceras stoliczkanum Gabb, An­
derson, p. 176, p�.
Holotype. Specimen figured in Douville's paper;
see synonym.
34 F. ETA Y O · SERNA
Hypotype. (U.N.C.M.H.N.C.P. Hy. N.5).
Type locality. "Vallee des Amazones.. [sic], un·
known.
Descn�· ti on after the only specimen avai·
lable [C-514 . This is a moderately evolute spe­
cies; the oral portion of the penultimate
whorl is characterized by widely separated, rela·
tively thin primary ribs three to four times na·
rrower than tlle intercostal spaces; they bear a
mammtfonn tubercle at the umbilical rim, and
a prominent spine at center of the flank. On the
adapicai portion of the last preserved whorl, the
ribs appear to be extended towards the umbill·
cal seam, with a feeble adoral concave arc; on
the inner flank the ribs are nearly radial but on
the adventral half of the flank and on venter
they swing slightly forward producing a wide
adoral arc; the growth lines are rme raised
threads which sometimes gather in bundles and
resemble intercalary ribs but no true intercalary
ribs are present. All ribs are depreued along the
ventral median line and on both sides of it pro­
minent crest like tubercles are developed. On
the adoral portion of the last whorl the ribs
are only slightly promiradiate. No living cham·
ber is preserved ; up to the anterionnost part of
the preserved conch it is septate. The whorl sec·
tion at this point ia in flated , rentfonn, it has an
Inclined �J:llbilical wall which rounds &mootbly
into the flanks.
�
DIAGNOSIS. This specie s is ch aracterized
by: A) its sparse ribbing with only primary ribs
or sporadic, weak intercalary ribs that disappear
before reaching the umbilical rim, and B) its re·
nifonn whorl &ectlon.
MeUUftlmenta. D Euw H W D/Euw % D/H%
C-614 80 30 20 8 3 IS O 33
COMMENTS.This is a peculiar species not
easily comparable to any described specie&
known to me; it is most aimllar to "Douuilleice·
ras Martini d'Orb." as described by Sinzow
(1907, p. 171, pl. 2, especially fig. 13a, 13b)
but on the Russian species the intercalary ribs
are common and at simllar dlamete:n when
rlbbinll: is attenuated in the Russian species the
o_ppOSlte seema true for the Colombian Wton.
The generic •ignment is questioned because of
the projection of the ribs onto venter on the
Jut whorl.
OC9URRENCE. Socota Member. Upper
Aptian.
� ���
Ch lon f&12.�f!eloni£erus�
�r_o'Sa�QL:n. sp.
PI.2, Fgs. 9·10; Text-Figs. 4R, 48; 11.
Holotype. C·223 (U.N.C.M.H.N.C.P.Ht. N.20).
Paratype. C-223 (U. N. C. M. H. N. C. P. Pt.
N.19, 20, 2 1).
A) Up to 25 mm D [ C-223·11. The umbi·
liens is deep and wide; on· the two imt observa·
· by: A) the ontogenetic development of the no­
ble whorls the sculpture can not be clearly dla·
cemed but at the beglnlng of the third whorl
(4th of the phragmocone?), very prominent
elongated spinose tubercles develop on the mid·
die of the flank (= along the involution line),
they arise from low wide ribs which extend
toward the umbilical rim. On the fourth or last
preserved whorl of this specimen very slight
mammiform tubercles develop at the periumbi·
lical incep tion of the ribs; at the same time on
both sides of the ventral median line, rounded
to acute swellings are developed. Although the
ribs tend to fonn a wide concave arc on the
fianks and to project weakly onto the venter
they also tend to widen on the adumbilical half
of the whorl as if forming an adoral arc1 at the
same time they tend to split longituainally.
Along the ventral median Une the ribs are de·
pressed; sporadically there is an intercalary rib
which differs from the primary ribs only in its
lesser strength.
The whorl section is subcompressed to
subpoligonal: the umbilical wall is inclined, it
merges smoothly into the slightly convex
adumbllieal half of the flank, the adventral half
of tl1e flank converges and rounds into the flatte·
ned venter.
B) Up to 35 nun D [C-223-2). The earliest
part of the last whorl of this specimen is like
· the late part of the preceding specimen. The or­
nament continues developping in a similar. way
_
except that one or two folds (Intercalatory
ribs?) without tubercles appear between two
primary ribs. The periumbilical tubercles conti·
nue inconspicuous but the ventral tubercles
tend to acoontuate; the ribs tend to extend to
the umbilical seam fanning and acute adoral
concavity.
C) Up to 50 mm D [C-223·11 ]. At the
beginning of the last whorl in this specunen the
distance between the centrolateral and periven·
tral tubercles progreuively increases. At this
stage the intercalary ribs are well defined and
characterized by lack of periumbilical or lateral
tubercles, they have only periventral tubercles;
this intercalary rib appears near the preceding
primary and approaches it near the umbUlcal
margin. The last two ribs of this specimen chan·
ge their strong lateral tubercles into tiny spines.
'Ibe Intercostal whorl section il aubrectangular,
it has an oblique umblllcal wall which tends to
become vertical toward the umbilical seam. The
ventral zone is nattened. Near the aperture the
ribs give the lmpression of slightly curved clubs.
D) Up to 7 5 mm D ( C-223]. The oma·
ment is u on the preceding specimen; the living
chamber represents two thl.rda of the last whorl;
the principal morphological features are the
progresaive weakening of the tubercles, especla·
lly the cen troiateral and ventrolateral onea,
which cwaes the ribs to loose their ventral
median depression . 'Ibe adaperturalmoat whorl
section is a slightJ.y depresaed dome.
.
DIAGNOSIS. Thia apeclea is characterized
des whose strength lncreaaes from mammiform
adumbilica! ones to the pertventral comute tu·
!.'lerdes which in intennediate stages of growth
 ZONATION OF THE CRETACEOUS OF CENTRAL C O LOMB I A BY AM M ONITES 35

are very prominent but fade out on the adult of the flank. The umbilicus is deep.
earlier than the lateral and adumbilical, B) its
simple, well separated club-shaped ribs, and C) B) Up to 21 mm (C· 5141. The ribs have a
its subrectangular, dome-like whorl section . tendency to develop a second, faint, radially
elongated tubercle at the periumbilical margin;
Measurement. D Euw li w DfEuw%D{HO/o!"o. R. on the venter they te��··tQ.l\*tpress along the
c��23·t 25 14 9 1'3· 6 6 36 ·J0(1/2) median line and to raise' sl}gJ:ill!: on both sides;
C·223•2 36 1 6 13 18 44 36 16 the ribs are very fine, thread-like to moderately
c-22a-3 33 14 15 19 42 46 19 strong, feebly sigmoidal on the flank and projec·
C-223-4 30 1 2 13 14 40 43 19 . ted as a wide arc on venter. ..
C·223·1i 41 19 16 20 46 39 19
C·223.0 32 14 12 16 40 38 9(1 /2) C) Up to 30 mm D [C-514 ) . The periven·
C·223·7 41 20 16 ? 48 39 19 traiand lateral tubercles accentuate for a short
C·223-8 7 7 7 7 ? 7 9(1/2) distance and then soon fade out. Sporadically
C-223·9 1 1 7 1 ? 7 9(1/2) some intercalary ribs detach from a node on a
C·223·10 43 1 9 16 21 44 39 22 primary at the middle of the flank or from its
C·223·1 1 6 3 25 20 26 47 38 19 periumbilical region. There is a tendency toward
C-223·12 47 21 19 22 44 40 ? an alternation of a secondary rib between two
c-223·13 --33 22 20 25 41 38 10(1/2) primary ones; at the anteriormost observed part
C·223·1 4 1 1 ? ? ? ? 1 0(1 /2) the whorl section is subquadrate.
(·223·15 57 26 21 27 46 37 20
C: ·223·1 6 73 32 25 36 44 34 23 D) Up to 60 mm D [C-5141. Only half of
a specimen is available on which four whorls
can be observed. The umbilicus is deep, funnel
COMMENTS. This species is similar to shaped . On the second internal whorl there are
Cheloniceras (Epich e loniceros) subnodosocos· low ribs which disappear before the umbilical
tatum (Sinzow) [ 1907, p. 175 , t ext- fig_. 4-5, rim; the umbilical wall is subvertical, high, it
pl. 2, figs. 1 8, especially plate 2 fig. 8 j, but
• rounds smoothly into the convex sides. On the
on the Russian species is a secondary rib which third whorl the ribs are rounded imd e x t e n d
arises from the lateral tubercles and a single in· as sharp ridges to the umbilical" seam, they des·
tercalary also occurs, a charateristic that does cribe an adoral ccncavity on the umbilical wall
not occur on the Colombian species; the latter and part of the adumbilical third of the flank
also seems to attain a larger size than Sinzow's but on the remainder of the flank they describe
species. From the preceding table it is clear that an adoral arc; at this stage of development the
the number of ribs is very constant in the Co· intercostal spaces are two to four times as wide
lombian species, contrary to the Russian species as the ribs when no intercalary - ribs are develo·
in which this number is variable. The record of ped.
Sinzow's species in the Villa de Leiva by Biirgl
(1954, p. 16) and quoted by myself ( 1968, p. Intercalary ribs are rare. On the last whorl
28), is now considered doubtful. the ribs are thread-like , acu te and conspicuous;
they begin near the middle of the umbilical wall
OCCURRENCE. Socota Member. Upper and trend rursiradiate to the umbilical rim where
Aptian. they bend abruptly , then they trend nearly stra·
ight or in a feeble forward arc on the central
This species is named in honor of Profe· third of the flank, then they cross the venter .
ssor Carlos E. Acosta, Department of Geocien· straight or bent forward. On both sides of ven·
cias, Universidad Nacional. Bogota. ter the ribs raise in a feeble bullae and the me·
median ventral region appears slightly excava·
Chelonicera� (EpJcheloniceras) jimenqL ted. The intercostal spaces are very wide, cha·
n. sp. nne!-like, three to five times as wide as the ribs.
Pl. 2, Fig. 8; Text-Fig. 4X. The whorl section is nearly rectangular with
rounded periventral margins, flattened ven�r
Holotype . C-5 14 (U.N.C.M.H.N.C.P.Ht.N.21). and moderately oblique umbilical wall.
Paratype. C-514 (U.N.C.M.H.N.C.P.Pt.N.22,23)
DIAGNOSIS. This species is characterized
A) Up to 15 mm D [C-514 ]. The whorl by: A) the transitory development of the lateral
section is renifonn. The sculpture consists of tubercles, B) the numerous fine, acute ribs on
veey faint ribs, slmost plaits, prorsiradiate, they the adult whorls with nearly no intercalary ribs,
describe a wide arc on the venter; the adapical and C) its subquadrate whorl section.
half of the last whorl on this specimen has the
shell preserved ; the ornament visible on it :s low, MeiL:Iurements. D Euw H w D/Euw% D/H0k
belt-like ribs ventrally arched and separated by c . 514 15 6 6 7 40 40
wide intercostal spaces which are covered with c . 614 21 8 8 10 38 38
growth striae ; fine flat ribs may interpose ado· c . 514 30 12 12 15 40 40
rally. The whorl section is a smooth arc. On the c . 614 60 30 21 26 50 35
adoral half of the last whorl of this srecimen,
inconspicuous mammiform prominences begin COI\IMENTS. This species is comparable
to form on the ribs at the middle of the flanks. to "Douvilleiceras Martini d'Orb" as figured by
:rhe ribs begin at the umbilical rim but some Sinzow (1906, p.171, pl.2, fig.13-17) but the
mtercalary ribs begin on the adumbilical third latter differs by the stronger development of the
36 F. ETA YO
tuberculation at equivalent stages of juvenile
development, and because on the Colombian
species at adult sizes the ribbing is regularly fine
with exclusively primary ribbing whereas on the
Russian species primary and secondary ribs al­
ternate and the whorl section is depressed,
oblong and not subquadrate as in the Colombian
species. No other comparable species is known
to me.
OCCURRENCE. Socota Member. Upper
Aptian.
This species is named in honor of the geo­
logist Andres Jimeno, former Director of the
Inventario Minero Nacional, and 1st. Director
of the Institute Nacional de Investigaciones
Geologico-Mineras.
Cheloniceras (Epicheloniceras ) barreroi
n . sp.
Pl.2, Fig.l; Text-Figs. 4P, 4Q.
Holotype. C-514 (U.N.C.M.H.N.C.P.Ht.N.22).
This is a peculiar stubby Cheloniceras
species with the general appearance of a water
wheel. The umbilicus is deep an wide and its
sides vertical on each whorl . The earliest whorls
observed (adoral part of the second whorl) h ave
strong ventrolateral spines, and the umbilical
wall and flank form a continuous broad arc; on
the third whorl a subvertical rather high umbili·
cal wall rounds smoothly in to the convex
flank; on the adapical portion of the fourth
whorl the periventral tubercles appear faint on
flank as broad, slightly raised ribs, shortly after
every other rib develop mammiform tubercles
on the umbilical rim.
On the fifth whorl the adult ornamenta·
tion is well defined and characterized by heavy
primary radial ribs which have a spine at the
umbilical rim and a second heavy and stronger
spine at middle of the flank where the rib bifur­
cates with the adoral branch thinner; on both
sides of the ventral median region all ribs deve­
lop transverse crest-like tubercles and the mid·
venter is depressed.
At the beginning of the last preserved
whorl there is an intercalary rib, between two
branching primaries, which is as prominent as
the primary ribs, but on the central segment of
the whorl the bifurcating ribs follow each othl!r
without any intercalary rib and finally near the
aperture no bifUrt!iltions occur even though one
intercalary rib beginning at level of the midi ate·
ral tubercle may ·be present, but the dominant
ribs are single prominent primary ribs.
The inte.tcostal spaces are twice as wide
as the ribs especially between two successive
primaries and narrower between the branches
of a bifurcating rib. At the end of the last pre­
served whorl its section is very depressed, trans·
versely elongated and polygonal in outline, it
has a vertical u�_bilical wall , flat diverging adum­
bilical half of Whorl, convergent adventral half
of whorl and flattened venter. The ribs stop at
- SERN A
the umbilical rim though when the test is preser·
ved a stressing of the growth lines occurs down
to the umbilical seam.
DIAGNOSIS. This species is characterized
by: A) a very depressed section of the whorl
and B) the development of its ornament which
begins with ribs laterally tuberculate and with
feeble swellings on both sides of the median
ventral line, they subsequently develop mammi­
form umbilical tubercles and bifurcate from the
centrolateral tubercle, finally only tritubercula·
te single ribs are present.
Measurements D Euw H W D/Euw "fo 0 /H %
C·S14 42 19 14 2 4 45 33
COMMENTS. This species resembles Che­
loniceras clansayesense (Jacob} (1905 , p.4.3, pl.
13, figs. 4a, 4b ,4c) but the Colombian species
develops its mature single ribbing earlier in the
ontogeny than the French species.
OCCURRENCE. Socota Member. Upper
Aptian.
Holotype. C-504 (U.N.C.M.H.N.C.P. Ht. N.23).
Four whorls are present on the largest
specimen ; on the first whorl -apparently the
second of the phragmocone- widely spaced
pleats are present, they bear long spines along
the involution line, and disappear about the
umbilical rim; on the third visible whorl small
spine-like tubercles appear at the umbilical rim,
they connect to the ventrolateral ones through
a wide triangular like extension of the rib,
which itself extends on the umbilical wall to
· the umbilical seam; at the same time one or two
intercalary ribs develop as fine pleats between
the primary ribs, it disappears about the umbi·
Iical rim and has ven trolateral spines; at this sta·
ge the umbilical wall is vertical.
On the last preserved whorl there is an
alternation of one or two narrow and acute
intercalary ribs, which may have a tubercle or
l ack it at the middle of the flank, but they
develop tenuous bullae-like tublercles on both
sides of their median ventral depression. Spora·
dically one of the primary ribs bifurcates at the
lateral tubercle with either the adoral or the
adapical- branch of it being the most prominent.
One intercalary rib may rise from the periumbi·
Heal tubercle of a primary rib. In general all ribs
describe a wide adoral concavity from the
umbilical seam to the lateral tubercle, from that
point Jhey slant slightly forward producing a
slight bow at the level of the lateral tubercle, .,
this is especially notable on the intercalary ribs.
lmfnediately adventral to the lateral tubercles
the ribs are sligh tly depressed and then they
fonn an elevated periventral tubercle; the
depressed ventral medhm region between the
periventral tubercles forms a wide shallow
groove.
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
The section of the whorl between the ribs
is suboval,almost as high as wide, but at ribs it
is polygonal with the umbilical wall subvertical
and high.
DIAGNOSIS. This species is characterized
by A) its high whorl section and B) the oma·
ment of its adult whorl -which varies from two
to more intercalary ribs between primaries or
coupl«!S of primaries.
Meuure ment.l D Euw H W DJEuw% D/H %
C·604 4'1 26 22 7 44 38
. COMMENTS .COmpared. to Chelanicerll$
• form, conical spinose and ·nod6Se: for adumbill·
(Epicheloniceros) barreroi n. sp. the angle for·
med by the segments of the ribs between lateral
aJ1d . adventral tubercles is obtuse in the latter
out in. Ch. (E.) pardai it is very steep, almost
ninety degrees ; in the latter species the bifurca·
tion of the ribs is irregular contrary to the for·
·
mer species.
OCCURRENCE. Socota Member. Upper
J\ptlan.
This species is named in honor of geolo­
gist Rafael Pardo.
Cheloni§ te f OCCURRENCE. Socota
Jfonhar W H
ichelonicerizs.L
_q__ n. sp.
Pt 6, fig, 3; Text-fig. 4U.
1938
. Cheloniceras clanseyense Jac., Riedel, p.
22, pl. 5, fig. 4 ,5, pl. 14, fig. 5.
1957.. Cheloniceras clanseyerue Jacob, Bilrgl, p.
1 3 5.
.
. Pl.2, fig. 4; Text-figs. 4Y , 4$ .
1965 Chetoniceros clanseyerue Jacob, Biirgl, in
· . Campbell and Btirgl.
Ilolotype. Specimen figured by Riedel, 1938,
Hypotype . ����l(U�N�.M.H.N.C.P. Hy. N.6)
. ·· .. Only two specimens available (C-223].
The earliest part of the shell observed is distor·
�d by .a malformation. On the adapical part of
�e last preserved whorl the first rib that can be
observed res.embles a low rounded ridge which
attenuates and fades out near the umbilical rim;
it describes.a tenuous,.wide adoral concavity on
the whorlSlde and on the venter they project
formlilg a. wide adoral arc. These ribs have a co·
nical tubercle in the middle of the flank and a
slight. bump on both sides of the ventral median
lb:u�. Th.e ribs are distant from each other and
are s.eparated by flat intercos� spaces. Advan·
·clng adoraly l the next rib has a
ucal mammiform tubercle slight periumbi·
which become
P.tominent on the next (3rd) rib. On subsequent
·
n,bs a. continuous strengthenir.g of the lateral
and yen . trola
teral tubercles is initiated but the
�.rnbdical tubercles scarcely increase in size.
ar �� gro�th lines describe a wide adoral
. ·ie w . c . begrns at the umbilical seam and
· B) Up to 25 mm D (same specimen). The
e:J�; . nds to the periventra
.��tnt they- project onto vente
l margin from which
r; in general their
37
trace follows that of the ribs. The whorl section
· at the preserved apertural end is transversely
subrectangular: the umbilical wall is oblique
and it rounds smoothly Into the convex sides
which themselves round smoothly into a
flattened or broadly arched venter. The umbili·
cal wall seems to become oblique in the adoral
portion of the last whorl. There are 13 ribs on
the last whorl preserved .
DIAGNOSIS. This species is characterized
by: A) its tapering ridge-like ribs slightly exten·
ding onto venter, B) the uneven development of
the size and shapes of the tubercles: mamml·
cal, centrolateral and periventral respectively'
and C) its wide flattened intercostal spaces with
only fine growth striae.
Meuurementa D Euw H W D/E1.1,w% DfH%
C·223 33 14 12 16 43 36
COMMENTS. This species was referred
by Riedel to Jacob's species (1905, p.413, pl.13,
figs. 4a, 4b, 4c; 1906, pl.l, fig. 7a.7b) but on
the latter the ribs bifurcate from the lateral
tubercle, the opposite is true for the Colombian
species.
Member. Upper
Aptian.
This species is named in honor of Dr.
Leonhardt Riedel.
.f!Jelonic!Jr!� C¥EJchelonf!:eras} huertasf..n.
sp.
Holotype. VL-7-Sa-11. (U.N.C.M,H.N.C.P.Ht.
N.4).
Paratype. (U.N.CM.H.N.P. Pt. 24,25).
A) Up to 10 mm D. The second whorl of
the phragmocone is coronate, it has a row of
spiniform tubercles along the periventral margins
(=involution line); from this row of tubercles
the flanks descend into the subvertical umbilical
wall. On the third whorl, ridge like folds extend
from the umbilical seam, describing a tenuous
adoral concavity, up to the periventral tubercles
with the maximum depth of the concavity at
the umbilical rim; however on the adventral
half of the whorl they are slightly convex
adorally or nearly straight. One or two of such
folds however bend on the adumbilical half of
the flank in such a fashion that they appear to
arise from the umbilical rim of the preceding
fold . From the periventral tubercle the ribs
cross the venter; the intercostal spaces are more
than twice as wide as the ribs; they are natte·
ned and they are at times interrupted by
a constriction-like groove. The whorl section
has a sub trapezoidal outline, with a broad roun·
ded venter.
ribs are stronger, they describe a continuous
shallow adoral concavity from the umbilical
seam to the periventral margins from where
38
they project slightly forward on venter, for·
ming a wide adoral arc. At this point the ribs
are differentiated between : A) primary ribs
which develop a tiny bullae like periumbilical
tubercle, that are acute to subrounded on the
flank up to the lateral tubercle, and then they
become rounded, B) intercalary ribs attenuated
in relation to the primary ones; they extend to
the umbilical seam as fine threads but become
well stressed on venter and C) secondary ribs
connecting the lateral tubercles and developed
adapically to the primary ribs. There is no regu·
larity in the occurrence of one type or the
other but at least on this specimen, after two
successive bifurcations there may be several, up
to three, single intercalary ribs.
The primary ribs are more prominent on
the venter and faintly flattened along the
siphonal line . Those ribs which bifurcate are
followed adorally by a constriction-like sulcus.
The intercostal spaces are slightly wider than
the ribs. On the last third of the last whorl of
this specimen all ribs tend to have the same
strength on venter, at the same time the inter­
eostal distance increases. There are 45 ribs on
venter of the last whorl on the specimen studied .
C ) U p to 40 m m D. Although i n th e pre·
viously described stages of growth the ribs are
less strengthened than in the specimen selected
as the holotype, I am considering them conspe·
cific because the general development and the
shape of the ribs is the same. In one specimen
there is a widening of the ribs toward the aper·
ture but this, as judged by sporadically develo­
ped similar spaces at intermediate sizes, is inter·
preted as due to hampering of rib development
during growth.
The last whorl of the largest specimen
examined has about 45 ribs on the venter, they
are sin,gle rounded, closely spaced , separated by
constrictit;li1Jike intercostal spaces; they bear a
: Hny periUmbilical spine, and an spinose centro·
jate�)l.ibercle. from which ribs may bifurqate,
and; sci.n\e'primary ribs at the adapical 'portion of
'"'the whorl have a'tenuous ventrolateral eleva.lon.
The, nb� i;?ild to be equ�y strengthened though
th� interealary ribs,
'
at�nuate more rapidly on
, · ·
�E! flanks. ; · · ' ·
,. , DIAGNOSIS. '!'his species is characterizP-d
by! the development 'of its .. ornament which be ·
gins with A) bifurcatihg primaey ribs and inter·
talary ribs, which B) grade into uniformly roun­
. ded, �losely spaced ribs and i:lituberculate rib·
bing in the adult and C) the section of the whorl,
reniform throughout the ontogeny .
Measurements. D Euw H W D/Euw %
VL • 7 • Sa • 1 1 8 8 8 3 37
VL • 7 · Sa · 1 1 23 10 9 18 43
VL • 7 · Sa · 11 41 51 16 23 36
COMMENTS. This species is similar to Ch.
basseae Breistroffer (1936, p.15 6 : Acanthoceras
(Cheloniceras) n. sp., Basse , 1928, p.138, text·
fig.15, plate 8, fig.1), but in the latter species
the section is almost circular, furthermore con·
F. ETAYO - SERNA
trary to the new Colombian species the ribs are
uniform in size except "jusque ' au voisinage de
l'ombilic a1 bord duquel elles semblent se grou·
per en donnant naisance a un tubercule", Basse
(ibid., p.138).
OCCURRENCE. Paja Fonnation (Arcillo·
litas abigarradas. Segrr.e nt E). Upper Aptian.
This species in named in honor of the pa·
leobotanist Gustavo Huertas, of the Universidad
Nacional de Colombia.
c���s-·�- �-��ecmitesf.�_Y;�!Gi��J
Vec.mites Casey, 1962, part.4, p. 256.
TYPE SPECIES. Vecmites caprotinus Casey,
original designation.
flo.-,
�u���?��'><\#.am�ranoitei�- sg.
TYPE SPECIES. Vectisites (Zambranoit�
}ambran?.f..n. sp.
DIAGNOSIS. Conchs of moderate size; the
early whorls have a depressed section, later
whorls are subquadrate to suboval; early in the
ontogeny centrolateral tubercles occur which
disappear at.a time when ventral elevations deve­
lop on both sides of the depressed median line;
the ribs are more or less strongly projected on
venter and are flared on the adumbilical half of
the whorl. Scaphitoid in general appearance ('?).
COMMENTS. This subgenus differs from
Vectisites (Casey, 1962) because of the strong
extension of the ribs onto the venter, the depre·
ssion of the ribs along the ventral median line
and the development of tubercles on both sides
of venter.
·
Holotype. C-504 (U.N.C.M.H.N.C.P.Ht.N.25)i
Paratype. C-504 (U .N.C.M;H.N.C.P.Pt.N.26,21 �
' t
A) Up to 15 mm D [C-504). The smalle�t.:
specimen at hand has a slightly subellipticatin·'i�
tercostal whorl section, with its maximum widtli:
at the umbilical rim, then it describes a dome·Ii·,
ke curvature; the umbilical wall is rounded-sub�
vertical. The ornament conslsts:or-fiexuous ribs.
which describe a weak adoral concavity ,on tq�·.
umbilical wall, and slant forward to aboUt the
boundary between the adumbilical and central
thirds of the flank, and then they describe a wi­
de adoral concavity on the remaining part of
the flank, and f"m ally they project onto the ven·
0/H% ter in a wide arc.
37
39 Toward the adaperttiral third of the last
39 preserved wh orl in this specimen, an angular
swelling begins to develop at the boandary be·
tween the adumbilical and central thirds of the
flank. The futercostaJ ·llp.BceS are 'Very Wide, as
much as four time!! the width of a single rib. Be·
tween two primary ribs there are two or three
indintinct intercalary riblets,one of which springs
 ZONATION OF THE CRETACEOUS OF
from the centrolateral tubercle of a PrimarY rib
and the other appears in a subdue manner near
the umbilical rim; the primary ribs are wide and
flattened on the venter.
B) Up to 20 mm D [C-504]. With growth
one or two intercalary ribs tend to become well
individualized and at the same time the vertex
of the adoral ventral arc of the primary ribs tend
to be depressed suggesting ventral tubercles.
C) Up to 45 mm D [C-504]. The flattening
or depression of the ribs on venter accentuates
with growth giving the ventral arc a truncated
aspect. The intercalary ribs become as promi·
nent as the primary ones except that they fade
out before reaching the umbilical rim. The
living chamber occupies about two thirds of the
last preserved whorl and It is characterized by
the irregular occurrence of the ribs, which are
well but variably extended onto the venter. The
Intercostal spaces are two or three times as wide
as the ribs. The section of the whorl is ttansver­
sely subrectangular and has rounded comers; the
umbilical wall is oblique. A dorsal shield is pre­
sent.
DIAGNOSIS. This species' is characterized
by: A) the broadly arched and truncated projec·
tion of the ribs on venter, B) the transversely
subrectangular whorl section of the adult, and
C) the discrete development of the ribs.
Me..uements. D Euw H W D{Euw0k D/H %
c . �04 16 6 7 1 37
C . • �04 20 7 7 1 1
0 " �04 46 22 16 20 48
COMMENTS. This species differs from the
type species of Vectisites Casey (1962 = V.ca­
protinus Casey, p . 2 5 6, pl. 40, fig. 9a, b, lOa, b ,
te.xt · fig. 89a), as noted for the subgenus.
·. OCCURRENCE, Socota Member. Upper
Aptian.
· This species Is named in honor of Francisco
Zatnbrano, geologist of the Instituto Nacional
de Investigaciones Geologico-Mineras, Bogota.
Xe
Pl.
'W', fig. 'Zambranoites
ites J cruzi D..J�
Text-Figs. 5A., 5B.
;
Holotype. C-504 (U.N.C.M.H.N.C.P. Ht. N. 26).
The first whorl observed seems to be
first whorl of the phragmocoile and Is smooth;
the second wh orl shows a high subvertical um­
• .
bllic:al wall which changes abruptly into the
lllightly convex aide; very fine, widely separated
pleats. are visible on the test. On the t."lird visible
who.:d those pleats become wide and low and
they have a faint conical node at the line of in­
volution which at that point is about at the
bound uy between the middle and adumbilical
thirds of the flank. At the adapical portion of
the l�t whorl o.f this specimen (4th of the phrag­
mocone) the nbs are clearly visible from the
wnbllical seam; up to the lateral tubercle the
nbs describe a tenuous adoral concavi ty . then
CENTRAL COLOMBI A BY AMMONITES 39
they slightly bend at about the central third of
fiank and rmaUy they describe a wide adoral­
concavity up to the periventral margin and pro­
ject onto venter, describing an acute , almost
chevron-like arc; this bend accentuates with
growth of the whorl.
The ribs become progressively depressed
along the ventral median region in such a
• manner that at the adoralmost portion of the
conch there are two elevated tubercle-like
shoulders on each side of the venter. The first
third of the length of the last preserved whorl
has a faint non-tuberc:ulate intercalary rib
which begins at the periumbilical margin. The
living chamber occupies about two thirds of the
last whorl and is characterized by loss of the in­
tercalary rib and increasing distance between
the primary ribs. The intercostal spaces are wi·
de, one and a half as wide as the ribs if one
intercalary rib is present but from three to four
times when the intercalary rib is missing, this
is especially noticeable on the internal mold.
The umbilicus is graded shaped, deep. The in­
tercostal whorl section at the adaperturalmost
part of this specimen is rounded subquadrangu­
lar.
DIAGNOSIS. This species is charaterized
by : A) its. small , relatively compressed conch,
and B) the rapid development and accentuation
of the adoral arching projection and depression
of the ribs along the midventer.
7 Meuurements D Euw H W D(Euw % D(H0k
7 C·604 2� 10 10 12 40 40
3�
COMMENTS. This species differs from
Vectisites (Zambranoites) zambranoi n. sp. be·
cause of its rapid acquisition of a strong ventral
projection of the ribs.
OCCURR ENCE , Socota Member. Upper
Aptian.
This species is named in honor of Jaime
Cruz, former Subdirector of the Instituto Na·
::ional de Investigaciones Geologico·Mineras,
Bogota.
Vectisites ozambranoite!)._ mateusi n. s�
Pl.6, fig. 2 ':" - ' '
Holotype. VL-11-3 (U. N. C. M. H. N. C. P. Ht.
N.27).
the Paratype. (U. N. C. M. H. N. C. P. Pt. N.28).
Two specimens are available. The conch
has moderately compressed sides and slightly
rounded venter. The adapical portion of the pe·
nultimate whorl bem close pleats on the flank,
ltl umbilical wall is vertical; toward the adapical
region of the last whorl the primary ribs begin
to diffe,rtlll tiat.e, they are flexuous, adorally con­
cave on the external part of the umbilical wall,
adorally arched on the central third of the flank
and finally projected onto the venter. The part
of the ribs on the adumbilical third of flank are
flanged but a tubercle is not developed. On the
ventral median line of the adoral portion of the
40 F.
last whorl the ribs are inconspicuously flattened.
On the last preserved whorl the ribbing is pro­
minent and consists mostly of primary ribs, so­
me of them in couples; intercalary ribs are also
present, they are initiated in a subdued manner
near the adumbilical third of the flank.
DIAGNOSIS. This species is characterized
by: A) the ontogenetically late and inconspi­
cuous (if any) development of the tubercula­
tion, B) its strong flexuous ribbing, and C) its
compressed suboval whorl section.
Meumementl D Eu.w H W bfEu.w0k D/H%
VL-1 1-3 33 12 13 7 36
VL-11·3 30 11 13 13 37
COMMENTS. Compared to the other spe­
cies In the genus Vectisites described from Co­
lombia, this is the only species which combines
a strong flexuoslty of the ribbing with great
attenuation of the tuberculation (?).
OCCURRENCE.Paja Formation (Arcilloli­
tas abigarradas. Segment E.). Upper Aptian.
This species is named in honor of Luis E.
Mateus, geologist of Ministerio de Minas, Bogo­
ta.
V£r;,tes _CZambranoites� cg4£_nqj_ n. sp.
Pl.6, 1g. 4, Text-figs. 5L, P.
Holotype. C-514 (U.N.C.M.H.N.C.P.Ht. N.28)
Only one specimen is available, it has the
shell preserved. The conch is stubby and com­
pressed. The ornament consists of ribs which
usually develop in pairs at the periumbilical
· margin; of these ribs, the adapical one is thic·
ker and rounded, radial, flanging outward on
the adumbilical half of the whorl side, it is be·
veled on the periventral zone and transversely
elevated in a bullae-like fashion on both sides
of a depressed and narrow median ventral re­
gion; futher adorally the ribs tend to be slightly
projected on venter; the adoral or secondary rib
tapers toward the·.'Un1bilical margin, on venter
it is narrower andlower than the primary one
and does not develop periventral tubercles on
the depressed median ventral zone. Sporadica­
lly an isolated primary rib develops.
The intercostal spaces between a prima·
ry and· its paired secondary rib is as wide as the
primary rib, but the intercostal space between
that secondary and the following primary is
wider than the primary rib itself. The ribs in
general are high and rounded and the intercos­
tal spaces are channel-like. Near the aperture
the ventrolateral bullae disappear. The intercos­
tal section of the whorl is low subquadrate with
rounded comers and convex sides. The 1J.mbili­
cal wall is oblique. There are 25 ribs on venter
of the last whorl. The apertural border is plain,
with a slight reflection at the umbilical wall.
DIAGNOSIS. This species is characterized
by: A) its depressed octagonal costal whorl sec­
tion, B) its heavy, rounded ribs, C) the presence
ETA YO · SERNA
of ventral bullate swellings only on the ribs, and
D) the angular periumbilical and centrolateral
bending of the ribs without development of tu­
bercles.
Meumementl D Euw H W DfEuw· % D/H %
C·614 36 16 12 12 44 33
COMMENTS. This species shows some re·
semblance to the type species Vectisites capro­
tinus Casey, (1962. p. 526, pl. 40, fig. 9a, 9b es·
pecially), but on the English species the lateral
tubercles persist longer; on the other side the
Colombian species is more depressed than
39 Casey's species.
43
OCCURRENCE. Socoti Member. Upper
Aptian.
This species is named in honor of Raul
Cadena, Universidad Nacional de Colombia,
Bogota.
D£tisit�s.l,Zg_mfJronQj{esLfl.ug_tteJi n. sp.
Pl. 6, lfg.2; Text-fig. 5!.
Holotype. C-514 (U.N.C.M.H.N.C.P.Ht. N.29)
Paratype. C-514 (U.N.C.M.H.N.C.P.Pt.N.29)
A) (C-514). The earliest whorl observed
(adoral part of the second whorl) has a depres­
sed subelliptical section : a broad arc connects
the center of the flanks, then the whorl rounds
smoothly into the umbilical wall which is incli·
ned. The sculpture at this stage consists of in·
conspicuous folds which describe an adoral
bend on the middle of the flank, and is delimi·
ted by a concavity both adumbilically and ad­
ventrally, these pleats are projected onto the
venter forming a wide adoral arc.
On the adoral portion of third whorl the
ornament is already visible: it consists of
prominent riblets which begin at the imprecise
umbilical rim; from the point to the periphery
the ribs describe one adoral arc at the center of
which a conical tubercle springs; from this
tubercle the ribs describe an adoral concavity
and then extend onto venter describing a
linguiform arc. The intercostal spaces are at
least three times as wide as the ribs, much
greater on venter; they are covered by growth
lines which parallel the ribs.
B) At 15 mm D [C-5 14 ]. At this diame·
ter the w_h orl section increases in height and the
conical nodes occupy a more central position
on the flank; the adventral half of the flank is
compressed, at the same time on the internal
mold the ventral linguiform projection of the
ribs flattens in a visor-like fashion and contem­
poraneously on both sides of the ventral me·
dian line the ribs begin to raise in transverse,
faint but perceptible tubercles. On the adumbi·
lical.half of the flank the ribs flare in a charac·
·
teristic fashion.
C) Up to 30 mm··n ( C-514). On the lar·
gest specimen the lateral nodes begin to fade
out contemporaneously with the development
 ZON ATION OF THE CRETACEOUS OF CENTR A L COLO M BI A BY A M M ONITES
of the ventral n odes . The final
type of ornamen­
tation -as far as i t is known- consists of well
m arke d slightly flexuous, well separated rib
s
which project a bit onto venter where they are
sligh tly d ep resse d along the ventral median area
and sl i gh tly raised on both sides of that area.
The ribs begin at the external part of the um b i­
lical wall w i th a little thi!:kening b ut are not tu·
berculate; there Is usu ally one in tercal ary rib
wh icll begins at the periumbilical rim wh�re it
is incl ined toward the preceding rib . Th e inter·
costal whorl section is transversely rectangular
with rounded comers. The um bili cu s is deep,
slightly inclined.
DIAGNOSIS. This species is ch aracterized
l:ly: A) the early and brief stre n � the �in g of its
lateral tubercles and the ventral hngu1form pro·
j ectio n or the ribs, and B) its compre ssed sub­
qu�drate whorl se c ti on .
Measurements. D Euw H W D/Euw % D/H%
c . 514 16 6 7 8 33 46
C · 514 30 12 12 14 40 47
COMMEN TS. This species differs fro m
Vei:tisites (Zambranoites) zambranoi n. sp. be­
cause on the latter the projection of the ribs on
venter is retained or stressed to a greater diame­
ter. Vectisites (Zambran oites) cruzi n. sp. by
cofrtrast stresses its ornament very early in the
otitogeny and the in tercostal distance is very
. l il��e,
OCCURRENCE. Soco ta Member. Upper
Aptian.
This species is named in honor of Hermann
Duque, p ale on tol o gis t of the Instituto Nacional
?� }nvestigaciones Ge ologico-Mineras, Bogota.
f : �L����C��( a a t
�yp : .�:,�o!.}!_ e�
HJ!pacanthoplites Spath, 1923, p. 64.
(8���:2!�
2 1
TYPE SPECIES. Acanthoceras mille tianum
(d'Orbigny) var. plesiotypica
Fritter , original designation.
raacan thoplites (?)
os_naliciae n. SJ?.: ..
PI03, fig. 1 1 .
Ho)otype. T.P.I. Guaduas (U.N.C.M.H.N.C.P.Ht.
, N.30).
Paratypes. T.P.I. G uaduas (U.N.C.M.H.N.C.P.Pt.
.·
N�30,31).
·.A) Up to 1 5 mm diameter{T.P.I. Guaduas].
At 'the . innermost whorls (D:9mm) the section
� �ubquadrate with rounded periventral mar·
ipDS and venter; the sides are convex, the um b i·
heal willl is subvertical. The ornament consists
o� ribs which begin at the umbilical rim and are
S!lgh tly flexuou s , rounded , with short vertical
s.l� es but flattened top and they progressively
Wl.d.en from the umbilical margin toward the
Penphery; the rib s are sep arated by subequal
.
!n.tercostal spaces; the former cross the venter
1�" a fee ble adoral curvatu re . The ribs are
sligh tly depressed along the ventral median line
41
producing on each side feeble transverse bullae­
like e levati ons : Occasionally a minute conical
tubercle developes on one rib (??). The ribs m ay
also bifurcate on the adumbilical third of the
flank; they tend to become more rounded with
growth.
B) Up to 30 mm D [T.P.I. Guaduas]. Th e
early portion of the last whorl is subquadrate, it
has flattened sides but rounded periven tral and
ventral margins; the u m bil ical wall is subvertical
and rounds smoothly into the flanks; at the
adoral portion of th is specimen the whorl sec·
tion is subrectangular. The ornament consists of
ribs which initiate midway on the umbilical
wall. The ribs are rounded , . well separated ,
sigmoidal; irregularity some of the ribs b i fu rcate
on the adumbilical third of the flank, both
from the adap i cal side of the primary ribs or
from the adoral side of th em ; there are in terca­
lary ribs also . The ribs become sligh tly higher
and wider on the venter. The ribs flare out on
the umbilical rim and are slightly we akened on
the middle of the venter.
C) Up to 50 mm D [T.P.I. Guaduas]. The
ornament is characterized by a conspicuous al­
ternation of ribs which on one side appear as
primary but on the other as secondary. Toward
the aperture the venter is well ro unded , the ribs
tend to be prorsiradiate uniform and elevated
on venter. The ribs have a weakly biconcave pro­
file an d are sligh tly elevated on the venter. There
are 37 ribs on venter of the last preserved whorl.
DIAGNOSIS. This spe cie s is characterized
at mediun and large sizes by : A) its radial to fee·
bly flexu ous ribbing, B) the ten d en cy of the
secondary ribs to branch from the raised peri·
umbilical flanges of the primary ribs and C) the
tenuous periventral bullate swl#llings on ven­
ter.
Measurements. D Euw H W D/Euw "/o D{H%
T.P.I. Guaduu 15 5 7 7 33 46
T.P.I. Guaduaa 30 10 13 12 33 43
T.P.I. Guaduas 32 18 21 18 35 40
COMMENTS. A comparable species was
described by Liss6n (1925, p. 27, p l . 2, figs. 4, 5 )
as "Parahoplites Mille tianum d' Ob . sp.", but
very little can be determined from his illustra­
tion or description except th at the Colombian
and the Peruvian specimens look very similar.
The neotype of Hypacanthoplites plesio·
typicus (Fritte l ) as figured by Casey {1965 , p .
42tl, Text- Fig. 155b, c) has stronger an d
flexuous ribbing o n the flan k s th an on the
Cclom bian species. Furthermore o n the former
the ribs are thic k on venter [on the illustration]
but they are thin in the C olom b ian species.
OCCURRENCE. Soco ta Member. Upper
Ap tian .
This species i s named i n honor of Miss.
Alicia Vanegas Leyva, former librerian of the
Servicio Geologico Nacion al o f Col om bi a.
42 F.
Hypacan thoplites dognaliciae n. sp . ?
Reference. Specimen C· 222.
The early portion of the last whorl is hi·
gher th an wide with maximum inflation on the
adumbilical third of the flank ; the sides are fla·
ttened and slightly convergent towards the wide
flattened venter. The umbilical wall is oblique
and rounds smoothly into the flanks. The early
part of the last whorl has fine acute ribs, which
begin on the middle of the umbilical wall or at
the umbilical rim; these ribs are more or less
falcifonn on flanks.
The primary ribs are characteristically
prominent, flaring out on the adumbilical third
of the flank whereas the secondary ribs may
flank from the adoral side of the primary ribs
on the adumbilical third of flank, or they may
appear independently but they are not empha·
sized adumbilically . The intercostal spaces are
twice as wide as the ribs. On the adventral third
of the whorl and on venter the ribs 1\le flattened.
The ribs are almost imperceptibly elevated on
venter. There are 38 ribs on the .renter of this
specimen.
Measurements D Euw H W D/Euw % DfH0/o
C·228·B 32 11 14 12 34
COMMENTS. This specimen differs from
the holotype mostly because the mature ('?) li­
ving chamber appears at a much smaller diame­
ter, this. occupies two thirds of the last preser·
ved �horl.
OCCURRENCE. Socota Member. Upper
Aptian .
r . -G"���;--��--�a�d�:rham}cer�-¥··'g�·n�i.
"""
'm:E"SPECiEs. Jua;;;r;;;;�··}u-;,;;t�rha-
mi n. g.1 n. sp. L>
n ;
:: Yu,r£nay•O. Kl."ttr� v-- 16.>o•f; .-1!/Ar
. ce those species closely resembling the type
DIAGNOSIS. Evolute conchs reachmg
moderate size . The early depressed whorl sec­
tion rapidly changes into a compressed oval sec·
tion. The early ornament consists of fine
flexuous, closely spaced ribs which cross the
venter uninterruptedly, these ribs progressively
separate from each other and widen on venter,
likewise the ribs differentiate between primary
and intercalary; with growth the primary ribs
flare out on the adumbilical half of the flank,
all ribs are less prominent on the adventral half
of the flank. AU' ribs have a tendency to be
excavated on their adoral face along the ventral
median line, producing on both sides of it more
or less conspicuous elevations. On the living
chamber the depression of the ribs abruptly
ceases and they have a girdle like appearance on
venter f= Colo mbiceras-like stage] . No tubercles
occur throughouWh.e ·ontogeny. . .
COMMENTS. This genus has already been
illustrated as "Du{renoyia " by Casey {1965 ,
p. 4 18, text-fig. 152) : "Dufrenoyia sp. example
showing Colombiceras·like sculpture on final
quarter-whorl, indicative of a link between the
E T A Y O - SER N A
Deshayesitidae and the Acanthohoplitinae,
"Upper Aptian, near Bogota, Colombia (Univ.
California no. C-1325)". I have examined this
specimen and as is shown on the right-hand fi.
gure in Casey's illustration the depression of the
ribs along th� ventral median line is accentuated
adorally before the "Co lombiceras-like" sculp­
ture suddenly develops. In other words, the de­
pression of the ribs ceases before the final living
chamber.
From study of this new genus, it ap·
pears that the "Clavate termination of the ribs
bordering the siphonal band" of the "post·ne­
pionic ventral growth" of Du(renoyiD (Caser,
1964, p.376) and the periventral "elevations '
of Juandurhamiceras are not homologous mor­
phological elements: on the latter the ribs in
early ontogeny cross. the venter uninterrupte·
dly and subsequently they tend to depress
along the ventral median line, but on the for­
mer the reverse is true with the addition that
the ribs are interrupted on venter in an early
stage.
Previous to the present study precise in·
formation on the exact stratigraphic position of
Juandurhamiceras was not available. It is here
presented because together with specimens of
44 the type species I have found a specimen that I
consider conspecific with the specimen illustra­
ted by Casey (op. cit., text-fig.I52). I am ma­
king it the holotype of a new species: Juandur­
hamiceras joepecki n. sp., characterized by the
rapid acquisition of the "Colo mbiceras-like sty·
le of ribbing".
Comments on the specimen illustrated by
Casey, both as to its uncertain stratigraphic
position or its significance as a "link" have been
made by Wiedmann (1966, p. 45) and Wied­
mann and Dieni (1968, p.93), respectively. The
genus Juandurhamiceras differs from Colombi·
ceras (Spath, 1923), here understood to embra­
species, by the lack of any indication of lateral
tubercles, at any stage of �owth. Incidentally
the tendency to proCiuce girdle like ribbings, i
believe, is not per se an indication of phlyoge·
netic linkage.
This genus is named in honor of Emeritus
Professor John (Spanish : Juan) Wyatt Durham,
of the Department of Paleontology, University
of California at Berkely.
...J.BtrodurhQ!!Ji"muuangurhamJ.. n . sp . .
�Pf.4, 1'ig.2; Pl.3, fig.5; Text-figs. 5C, 5J; 12.
Holotype. Santa Ana A.P.G. (U.N.C.M.H.N.C.P.
. Ht. 31).
Paratype. Santa Ana A.P.G. (U.N.C.M.H.N.C.P.
Pt. 32,33,34).
·-.,�. . : •. . . , ..
.
A) Up to 30 mm D (Santa Ana A.P.G.).
.
The early whorls were studied by disecting one
specimen. The protoconch is depressed; the
adapical section of the. .second whorl is wider
than high , with its maximum width located at
center of flank, but the adoral portion of this
 ZONATION OF THE C R E TACEOUS OF CENTRAL COLOMBIA BY AMMONITES 43

G
whorl is subcircular. The third whorl is higher DIA NOSIS. This species is characterized
th an wide. by: A) its moderately large size and evolution,
B) the regular alternation of feebly flexuous,
The earliest ornament observed is on the on
volulion
wen spaced , primary and secondary ribs
second and consists of annular fine middle and late stages of growth, and C) the
rounded ribs which are visible from near
the umbilical margin, they then trend pr�;>rsira· ;�J����rJ: ri:
ss g of �
. in �
� ��
ventral "ele·
- - .-:"i"':.-""·
diate as a very elongated S on the convex·flank ,
and cross the venter unihterruptedly; they are Meuurcneru D Euw H w 0/EUW 0/H No.R
as wide as the intercostal spaces though a cons· % %
trictl on-like intercostal space can be observe�
on this specimen. The ribs rapidly become na­ Sta. Ana · 1
Sta. Ana · 2
30

rrower than the intercostal spaces (adoral por­ Sta. .Alla · 3

tion of the third volution) and at the same tl­ Sta. Ana • 4 26 10 11 8 38 43 20
l]le they broaden on venter; shortly after (H:7 Sta. Ana · r. 26 11 10? 8 44 40'1 21
mm) the ribs become prominent on th e flanks; Sta. Ana • 6 30 14 12 9 47 40 2&
contemporaneously the ribs begin to inflate on Sta. Ana · 7 36 167 1& 10 .42? 42 26
the adumbilical third of the flank and to con· Sta. Ana · 8 29 12 12 9 41 41 22
tractact on the adventral third; this contraction Sta. Ana • 9 37 17 13 1 46 36 24
is followed by the development of a slight de· Sta. Ana ·10 437 1 9 17 12 44 39 20
pression of the ribs on their adoral .side along Sta. Ana ·11 39 19 16 10 49- . 41 26?
the median ventral line, this produces delicate
to accentuated humps on both of venter; sides Sta. Ana ·12
Sta. Ana ·13
38
367
20
17
15
1&
11
10
63
47?
39
42
21
. 24
the ribs have not been observed to be interrup­ Sta. Ana ·1 4 48 24 18 12 50 38 .• 2&
ted . on venter. At this stage of growth the ribs Sta. Ana ·1 5 36 16 13 10 46 37 24
are prominent, separated by wider intercostal Sta. Ana ·16 60 26 18 13 62 36 . 215
apaces and they are straight of feebly flexuous. Sta. Ana ·17 4 1 7 19 16 12 46 36 26
Sta. Ana ·19 337 14 13 9 42 39 22?
Sta. Ana ·20 267 117 11 ? ? 7 . 26?
B) Up to 41 mm D (Santa Ana A.P.G.). Sta. Ana •26 41 21 15 10 51 36 22
43 ·
. •

As in the preceding specimen, the successive Sta. Ana ·27 36 16 13 9 37 21

whorls only cover the external third of the pre­ Sta. Ana ·30 59 30 21 16 61. 35 28
ce ding whorl; the umbilical wall is inclined and Sta. Ana ·31 64 28 21 15 62 39 24
evenly rounds into the flanks; the periventral Sta. Ana ·34 407 18 16 10 45 40 26
rnaigiq is abruptly rounded and the venter is Sta. Ana ·35 31 14 12 9 46 39 22?
rounded to flattened . Sta. Ana ·36 37 17 14 11 46 38 26
Sta. Ana ·37 3 6 ? 18 13 10 46 37 23
Sta. Ana ·38 37 18? 14? 10 49 38 21?
The . ornament consists of primary ribs St.. Ana ·39 327 16 13 9 47 40 19
which appear about the middle of the umbilical Sta. Ana -40 35 17 13 9 48 37 207
wall but may extend, on exterior of shell to the Sta. Ana ·42 49 24 21 11? 49 43 28
umbilical seam. The ribs describe a sligh tly adoral
concavity in the umbilical region, then a con­
laxity on the adumbilical and middle thirds of COMMENTS. Same as for the genus.
th� flank where they are flared, then they are
fe�bly concave adomlly or straight on the O CCURRENCE. Socota Member. Upper
adventral third . of . the flank, there they are Aptian.
'
.flattened and contracted; the ribs cross the
l't!.n ter straight, flat-topped and prominet; along
tbe� periventral margin the ribs have delicate J!MmdYdJami�t{lJISjoepecki.,n. sp
lil:4, fig. 5.
•

�l�vations. The ribs are now differentiated into

cp,unary and secondary ribs, the former genera­ Holotype.U.C.M.P C -1325 (Dlustrated by Ca­
JI:y· flare on ftanks out to the involution line
••

sey, 1965, p . 4 18, text-fig. 152).

��m :\Vhere .they are less prominen t; the secon·
g!!Y ribs anse imp recisely outside the umbilical
Hypotype.Sta. Ana 9 (U.N.C.M.H.N.C.P.lly.7)

f»1� 111Jd generally do not flare out.

��j(,';�� �::S���n:;;nrl��
Only one specimen is available (Sta Ana

nt :!� �� A.P.G. 19). Very flat, slab·like sides, planar ven·

on
a ter and very wide, radial ribs which are like clo­

f:lt: t
��1Ve PMl&Q' or secondary. ribs occasiona·
ve buds; ther are slightly but clearly flanged
the adumbihcal third of the Dank. On t.'te In·
.t'Wl;�lll· Th e in tercostal spaces are almost temal mold the intercostal spaces are one and a
, c �-, I1S wide as the ribs. The periventral half times as wide as the ribs; the ribs are very
. •

,�I�y.a,fions on the ribs attenuate

wide on venter and cross it linearly.
14.o�. end of the conch at this diam
towa.tl
. the
;��g eter. The
n. sp.
iila"'·�
ell 8Jll ber ccuJ?ies three four

'"·
'�lution,
frund
� !bs (3/4) of
on Its venter the n�s broaden
and th� adventral elevatio
luqndurhamic�ras_gj[_qldgj,
PI. 3, fig:-"!3.
l
ns disap­
.

, . owever, particul ar y on the

interna! mol d.
!l_dQ� face of the ribs on venter show a
. ctetllctlon.
. ._,:,;:·..;
Holotype. Sta Ana A.P.G.
C.P.Ht. 32) •
N. 42. (U.N.C.M.N.
 44 F. ETAYO · SERNA
One large specimen is referred to this spe­
cies. The early internal whorls are covered with
fine rounded ribs, convex on the adumbllical
third of the whorl ; some of them, especially
those which begin at the umbilical wall are pro­
minent and flaring (=primary ribs); the secon­
dary ribs start at the periumbilical margin usua·
lly in pails, they are thin. On the last preserved
whorl the ribs are fine, particularly on the in·
temal mold. The primary ribs describe an adoral
concavity on the umbilical wall and part of the
periumbilical third of the flank, then they des·
cribe an adoral bow, sporadically very strong,
approximately on the central third of the flank,
on the remaining third they develop an other
adoral shallow concavity and the profile of the
ribs is consequently biconcave; the ribs are sligh­
tly thicker but rounded on venter where they
are feebly elevated. The whorl section is higher
than wide , It has an inclined umbilical region
which rounds into the flank; the periventral
margins are rounded. The last whorl has 28 ribs
on venter; the living chamber is preserved as half
the length of the last whorl.
DIAGNOSIS. This species is characterized
by: A) the delicately humped periventral mar­
gin of the ribs is transient, B) the flexuous rib·
bing starts early in the ontogeny, and C) the rib·
bing is less prominent.
Me&llll'emenu. D Euw H W D/EUW D/H No.R.
Sta. Ana 42 49 24 21 1 1? 49 43 28
. COMMENTS. This species differs from
Juizndurhamiceras juandurhami n. sp., because
on the latter the whorl section is more contrac­
.ted adventrally, and its ribs have conspicuous
periventral elevations throughout ontogeny.
O CCURRENCE. Socota Member. Upper
Aptian.
This species is named in honor of Mr. Car·
los Giraldo of the Instituto Nacional de Investi·
gaciones Geologico-Mineras, Bogota.
cG·����".":If��-!f��Pfii��� 19�;·\
Acantho hoplites Sinzow, 1907, p.458.
TYPE SPECIES. Acanthohoplites aschiltaensis
(Anthula) , subsequent desig·
nation, Spath, 1921.
A canthol!gp_lites b�oureti[grme. n. sp.
Pl .5, fig. 2; Text-fig. 5U, 5V.
Holotype. C-127 (U.N.C.M.H.N.C.P.Ht. N.33)
Paratype . VL10·10 (U.N.C.M.H.N.C.P.Pt. N.35)
" ?1938 Acanthoplites bigoureti Seusnes, Riedel,
� �
p.45, p,
J, . g.7, pl.14, fig. 24.
_,
19 S8 Ac�rrfohbplites bigoureti sensu Riedel
(non Seus.), Etayo-Sema, p .20, fig.3.
1968 Acanthohoplites bigouretii sensu Riedel
(non Seusnes), Etayo-Sema, p. 28, tab.L
i970 Acanthoh-(oplites) bigoureti, Wiedmann,
p.484, ex review of Etayo-Semas's paper.
Only half an individual is available . On
the earliest observed whorl there appears to be
bullae-like tubercles along the line of involution,
however the preservation does not permit one
to be sure of this, after these bullae there is a
tract on which pleats rather than ribs are deve­
loped. At this stage of growth the whorl section
is depressed; the umbilical wall is subvertical
but there is no well defined umbilical rim. At a
larger stage (on the penultimate whorl) the ribs
differentiate. The prominen t primary ribs begin
at the umbilical seam, they trend across the
umbilical wall and adumbilical half of the flank
describing a slight adoral concavity; one or two
intercalary ribs develop between the primary
ribs, they attenuate before reaching the umbili·
cal seam.
On the last whorl the intercalary ribs are
present in groups of two or three, of these ribs
the adoral one sometimes inclines towards the
immediate adoral primary rib as lf branching
from it. The primary rib develops a prominent
triangular area with a spine; from this tubercle
the ribs bifurcate or trifurcate . The intercostal
spaces are sligh tly wider than the ribs when
shell is preserved but they are one and a half
times as wide as the ribs on the internal mold.
The ribs themselves have steep sides and roun·
ded top, they describe a wide concavity on the
flank and project on venter where they are
slightly wider and flat-topped, the latter .is.
particularly conspicuous on the intemlll mold,>
On the venter the ribs tend to be :equally
developed but occasi onally some are narrower.
At the beginning of the last whorl the section is
subcircular, at its adoral part it becomes ogival,
slightly compressed ad orally .
DIAGNOSIS. This species is characterized
by: A) the accelerated strengthening of its oma·
ment, especially the lateral tubercles, B) the
constancy of the wide intercostal distance, C)
the dominant flaring character of the ribs, and
D) the ogival section of the whorl on ribs.
Mea��Urements D Euw H W D/Euw D{H No. R.
•;, Ofo
C-127 24 11 9 9 46 37 21(1 /2)
COMMENTS. The section of the young
specimen described by Jacob as "Douuilleiceras
bigoureti Seunes sp." (1905, p. 4 15 , pl.3, fig.
Sa, 6b) is transversely elongated, lenticular. The
inner whorls of the Colombian specimen ap­
pears to have more ribs than the specimens refe·
rred to Seunes' species and figured by Pervin·
quiere (1907, p.195, pl.S, fig. 37, 38) both sha·
re however a circular section (intercostal) of the
whorl.
c;'5eunes (1887' p.567) previously 'noticed>
that in this group of taxa the young specimens ·
of related (?) species are similar, the species
being differentiated .. only in the adult stages.
Seunes' figured specimens are larger than my
specimen, however, their primary ribs are out·
 ZONATION OF THE C RETACEOUS OF CENTR AL C O L OMBIA BY A M M C NITES
my species on
standing club-shaped contrary to
which the intercostal spaces are wide and the
nbs only slightly increase their wid th.
OCCURR ENCE. Shales above the Socota
Member and below the Capotes Member. Upper
Aptian. Paja Formatiol). (Arcillolitas abigarra­
das. Segment E). Upper Aptian.
c�nwohon lites seunesitjmn.t n.sp.
. 7,
1g.8; Text-fig. 5Q, 5 R.
.
llolotype. C-127 (U.N.C.M .H.N.C.P . Ht. N.34)
Paratype. C-127 (U.N.C.M.H.N.C.P.Pt. N.36)
A) Up to 24 mm D. This specime!l has
the iimer whorls leached away ; the penultunate
whorl has periodically raised and elevated pri­
mary ribs w..hich bifurcate at the umbilical rim;
between two of the primary ribs there are two
or three secondaries, the fanner appear at the
umbilical seam but the latter usually do not.
The umbili cal wall is subvertical and rounds ra­
ther abru ptly� into the flank. On the third
or last preserved whorl, the primary ribs begin
to . develop triangular tubercles which have a
· small spine on top, from them the ribs bifurcate.
All the ribs cross the venter with similar strength,
they are almost rectiradiate and they cross the
venter nonnal to the median ventral line. When
the. shell is preserved the ribs are as wide as the
intercostal spaces, they have steep sides and
rounded tops. The intercostal whorl section has
flattened sides and broadly rounded venter,
it is higher than wide. Toward the aperture one
intercalary rib usu ally occurs between two
primary ribs lacking tubercles or with only
i
a minute con cal one but without a bifurcation.
B) Up to 48 mm D. In this specimen, the
from the third whorl on, the oma­
· .•
:ho]otype,
Q:tent consists .of one intercalary rib , tapering
· toward the umbilical rim, between prominent
pri:rri ary ribs with subre.ctangular profile ; all
>P.�s cross the venter straight and became slightly
. ?IVi�ened. The ribs are sli ghtly narrower than the
ifitercostaLspaces where the shell is preserved
,ut on the internal m old the spaces are at least
' twice as wide as the ribs; the umbilical wall is
sh:ort and rounded.
'-' : DIAGNOSIS.· This species
Is ch8lacteri­
� ze_� by: _A) . a transitory tuberculate stage (up
,/ttl: the th.irdwhorl), With numerous cloS£:ly spa­
_
}�,-�� rib� and B) the adult stage on which stron­
;flf: J:[a,nn_g rect!radiate primary ribs and less pro­
;ii�J#.ent Intercalary .ribs alternate; all ribs are
�:·��\�Y c:ontrac te d on the adventral third of the
; _��-�menta D Euw H W D{Euw
;_ · < ) .· ;c:.iz7 . 24 10 8 9 r; ;; +4o
x.;i. -�';�.'C-127 48 24 1 7 1 6 &o 3 1'i 20(3/4)
: : .COMM?NTS. This species ;esembles
A aw{Joh�plrtes _se�nesi (Jacob) (Br"istroffer
� Pi7
�
· Dou u1llezceras bigoureti
tr- 4 1 6 , pl. 13, figs. 7a, 7b) , but
9 5• P
atter as s onger ornamen
" tation, the whorl
45
section is more compressed, and the intecalary
ribs are shorter than on the Colombian species.
OCCURRENCE. Unnamed beds above the
Socota Member and below the Capotes Member.
Upper Aptian.
A£an. th�hO-¥J.i1� �lLtasu,nQ.,n. sp.
Pl. 7, fig. 2, Text- 1g. 5Z, �-
Holotype. C-157 (U.N.C.M.H.N.C.P.Ht. N.35)
Only one specjmen is available. The inner·
most observed whorls have the umbilical wall
convex, it merges into the flank without defi·
ning a rim; the :flanks and the ·venter are evenly
convex but the whorl section is wider than high.
At this stage the ornament consists of rounded
ribs, almost as wide as the intercostal spaees,
they are annular to slightly prorsiradiate, they
taper toward the middle of the umbilical wall.
Groups of three (common) to four of these
ribs are separated by slightly more prominent
ones which develop a large, conical to rectangu­
lar tubercle immediately adumbilical to the
involution line; from this tubercle the ribs bi·
furcate : the aaapical branch crosses the venter
straight but the adoral branch projects slightly
adoraly. Tne intermediate rib nearest to the
emphasized primary rib is inclined as if arising
from the latter near the umbilical wall .
At a whorl height of 5 mm (correspon·
ding to the adapicalmost part of the preserved
whorl), the intercostal spaces are wider th an
th e ribs especially o n th e internal mold . A t this
whorl height the venter is broadly 8lChed to
flattened, it has rounded periventral areas. On
the adventral half of the last whorl only one or
two ribs appear between two tuberculate ribs.
On the last quarter of the last whorl the prima­
ry ribs do not bifurcate though the tubercle is
present; the intercostal spaces have become
very wide, channel like, but shallow, the ribs
have si,P.es sloping symmetrically (approxima·
tely 45 ?) and have a subrounded top. The aper­
ture is rectangular with rounded comers and it
is wider than high. There are about 50 ribs on
the last incomplete whorl. Sometimes there is a
�:onstriction-like interspace, adoral to the ribs
with a node.
DIAGNOSIS. This species is characterized
by: A) the decrease in abundance of the ribs from
the inner whorls to the last one , B) the rapid
attenuation of the lateral nodes and disappea­
rance of bifurcation, and C) its transversely sub­
rectangular whorl section.
D!H No.R.
Me>tlllrements.
c - 1 27
D Euw
24 11
H
8
W
9
D{Euw %
46
0/H %
33
COMMENI'S. Acanthohoplites rochi Breis­
Sinzowi"
troffer (194 7, p . 66: "A can thoplites Mille tianum
D'Orb. r.p. var Roch, 1926, p. 290, pl.
18, tigs. 1, 1a, 2, 2a; especially fig. 1), has an
ontogenetic development which Is the opposite
var seunesi to that of the Colombian species: on its early
whorls it has well marked and separated ribs
whose number Increases with growth. From
46 F. ETAYO - SERNA
"Acanthoplites Derognati Roch" (1926, p.
21 8, pl. 18, fig. 4, 4a) the Colombian species
differs mainly in the whorl section which is hig·
her than wide in the French species and in the
varying strength of the ribs on the venter of
Roch's species, they are uniform or only spora­
dically accentuated · on the Colombian species.
OCCURRE NCE. Unnamed beds above
the Socota Member and below the Capotes
Member. Upper Aptian.
. Acanthohoelites odiosu§.n. sp.
Pl. 4, fig. 4;Text-figs ER , 5ti.
Holotype. C- 127 (U.N.C.M.H.N.C.P.Ht.36).
Only two specimens are available but nei­
ther shows the innermost whorls. The best pre­
served specimen has the earliest portion of the
last whorl depressed, the umbilical wall is sub­
vertical to rounded and it passes into the adum­
bllical half of the convex flank, the adventral
third of the whorl is slightly inclined and the
venter is flattened.
The ornament is divided into prominent
primary ribs and less prominent secondary ribs;
the former are of two types: A) prominent
radial ribs which begin at the umbilical seam (?)
and flare out forming an arc on the adumbilicai
third of the whorl and terminate in a small co­
nical tubercle, at this tubercle the rib bifurca­
tes, the adapical branch crosses the venter al­
most straight whereas the adoral branch is fee­
bly arcuate and B) these are thin annular and
parallel ribs between two prominent ribs and
they vary from one to three in number and the
rib _nearest the subsequent adoral primary is
inclin-ed to it as if bifurcating from it.
All ribs converge on the adventral third of
the flank toward the venter as if forming ventro­
lateral shoulders, they are arched but adorally
not flattened. Adorally some ribs bifurcate on
the ad umbilical third of the whorl without deve­
loping a tubercle, or a rib can develop an adven­
tral tubercle without bifurcating, at least on one
flank. These ribs have a tendency to develop a
second tubercle, poorly developed at the umbi­
lical rim; this gives the ribs a definitely polygo­
nal profile.
The adapertural segment of these speci­
mens has a s�.�o);w.uadrate whorl section with
weak periventral bevelings. There are 35 ribs on
venter of three quarters of the last whorl. The
spacing of the ribs increases adaperturally and
the number of ribs between a pair of emphasi­
sed ribs decreases to one by strengthening of
the other. The ribs have symmetrical slopes and
are rounded . The intercostal spaces are shallow,
almost twice as wJ!ie as the ribs on venter, less
wide on the flank; the ribs do not widen on
venter.
.ded tops; on venter and especially when the
DIAG NOSIS. This is a stout species cha­
racterized by : A) its change in whorl section from
depressed to subquadrate, and B) the strength
of its ornament with flaring ribs on the internal
half of the flank.
Measurements. D Euw H W DfEuw % D/H %
c - 1 27 26 11 10 10 42 38
COMMENTS. This species resembles Gar­
gasiceras acutecostatum (Riedel} (1938, p.42,
pl.81 fig. 1-6, pl. 14, fig. 23) but on the latter
the JUVenile has a subquadrate whorl section but
this is transversely subelliptical on the former;
on Riedel's species the flaring of the ribs accen­
tuates with growth but the opposite is true in
A canthohoplites odiosus n. sp. The latter differs
from Acanthohoplites quitasuegno n. S?· by the
less inclined umbilical wall, its section is subrec­
tangular and its ornamentation is weaker at all
comparable diameters.
OCCURRENCE. Unnamed beds above
the Socota Member and below the Capote&
Member. Upper Aptian.
b.f:
Acaatho Ute§ olr,ui,Q6tatym n. sp.
Pl. 7, fig. , Tex-fig. 58, 51\
Holotype. C - 127 (U.N.C.M.H.N.C.P.Ht.N.37).
Paratype. C 127 (U.N.C.M.H.N.C.P.Pt.N.37).
•
The description is based on the holotype.
The part of the phragmocone nearest the pro·
toconch is smooth ; approximately at 2.5 mm
diameter very well marked bullae-like tubercles
develop just adumbilical to the involution line;
these tubercles seem to extend toward the um­
bilicus and at about 3 mm diameter the tuber­
cles are clearly prolongued toward the umbili­
cus forming ribs; at this stage the umbilical wall
is convex but there is no umbilical angle and
the umbilical wall grades smoothly into the
convex flank.
The ribs begin at the umbilical seam, they
run straight on the umbilical wall, they form a
slight concavity on the adumbilical third of the
flank and subsequently follow an arc to the
involution line , from that point to the periven­
tral area they form a concave and finally pro·
ject on venter. As soon as the ribs h ave beco·
me clearly recognizable they appear like fine,
rounded weak adorally biconcave cords separa­
ted by slightly wider intercostal spaces.
The earliest portion of the conch is orna­
mented by primary ribs; shortly after this stage
the ribS" differentiate into primary and interca­
lary, the latter tapering and fading out toward
the umbilical walr; these intercalary ribs incline
toward the preceding primary as if bifurcating
from it near the edge of the umbilical wall.
On the adapical portion of the last whorl
(D: 18mm) the ribs are slightly adorally arched
on• venter; they are thin, relatively high and ha·
ve almost symmetrically sloping sides and roun­
shell is preserved the central part of the ador&!
slope of the ribs widens in a visor·like fashion.
Adorally on the last whorl, some primary ribs
have a slight tendency to flare out on the adum-
 ZON ATION OF THE CRETACEOUS OF C E N TR A L C O L O M BIA BY AMM ONITES 47

bili cal half of the flank. The ribs have a marked tubercles at level of the involution line (adven·
tendency to ap p e ar as primary on one flank and tral third of the flank).
as a secondary on the other. The whorl section
near th e aperture is vertically subelliptical. There Up to 17 mm D · (C·l27� Amore advan­
are 62 ribs ori the venter of the last whorl. ced stage of growth is marked:by a shifting of
the point of ini tiation of th�.,i®ondary ribs to·
. DIAGNOSiS. This species is chara�rized ward the middle oftheo,flank;.s displacement
by : A) its progressively-- steeper umbilical wall occurs progressively ; there may be one or two
during growth, B) its subovate section and C) its intercalary ribs between two primary ribs. The
ornament consisting of fine, numerous, irregu­ aperture is retracted on the adumbilical third
larly alternating primary and secondary n'bs of the whorl.
o
wh se profile varies from fee bly flexuous
to
o
bic ncave .
.

C) Up to 25 mm D (C-127). The orna·

men tation described for the previous stages
Measurement.. D Euw H W D/Euw% . D/H o/o consolidates as alternating priJl!� ribs which
c - 127 24 10 9 9 42 37 begin at the umbilical seam describing a conca·
C - 127 19 7 8 8 37 42 vity , they draw an arc on the central third of
the whorl and draw a faint concavity or follow
NOTE : Width and height give similar values be­ almost straight on the adventral third of the
cause · they are approximate. whorl and then extend onto the venter in a
wide arc; at this point the whorl section is
COMMENT S. This species differs from all vaulted and the umbilical wall is inclined, the
ot:her species here referred to Acanthohoplites
and
ribs are finer than the in tercostal spaces and the
because .of its subovate whorl section tubercles disappear. There are 46 ribs on the
nuinerous fine ribs. last whorl of the largest specimen studied .

OCCURRENCE. Unnamed beds above the DIAGNOSIS. This species i s characterized

Socota Member and below the Capotes Member. by : A) the rapid change of the early tu berculate
Upper Aptian. ornamentation into flexuous ribs, B) its roun­

o�ites elegaTJ,{g,Qll.(f:,n . sp .
ded whorl section and, C) the alternating long
�l.cqnthoh
5 , fi g A;e t-fi g. 5X.
x
primary ribs and short intercalary ribs.

Measurements 0 Euw H W D tEuw ",l, D IH %

Holotype; C-127 . (U.N.C.M.H.N.C.P.Ht. N.38) C-127 12 4 5 5 33 42
Paratype. c:121 (U.N.C.M.H.N.C.P. Pt. N . 38) C-1 27 17 7 6 6 41 35
C-1 27 25 10 10 9 40 40
· ·· A)Up to 12 mm D (C-127). O n this small
specimen the protoconch is visible, it is followed COMMENTS. This species resembles
by the smooth first whorl which has two cons­ Acanthohoplites bigo ti (Seunes) (1887, p. 5 68,
qii�tions, , one at the contact with the proto­ pl.12, figs. 2a, 2b), and although comparison at
con.ch · and a second one half the length of the similar diameters can not be made the retention_
yolution. The beginning of the second whorl is of an accentuated ventral flexion to a larger dia­
Jndlcated by a raised , bullae-like tubercle at the meter is apparen t on the specimen illustrated
Jnyolution line; these tubercles (10?) are widely by Seunes; in general the French species seems
separated from each other and tend to reach to h ave a more compressed section than the Co· ·
'fhe umbUical region as weak pleats; on the lombian sp ecies. Jacob ( 1905, p .4 12) referred
cJ
�tll ir ·whorl these tubercles disappear and only to Seunes species but did not figure it, so his
�Milt,� fol� s are present, some of them slighly comment that at a diameter of 15 m m "certai­
,�,9re pro nu n en t than the o thers , but all seem nes cotes l!!gerement plus accentuees sont bitu·
Jl;) �nd at the umbilical wall ; they are rounded rew!lees" can not be utilized, though such
��g low:, they are prorsiradiate on the visible phenomenon never occurs in the Colombian
·
· 9� the fla,nk; the intercostal spaces are a species ; likewise the French species, as described
Wider thll!l the ribs. by Seunes, has more numerous ribs.

�,'.,c·;;:;�;,q,� .
th e last , whorl (=4th) the ribs are
fi.J!!1'�91Js; they have a convexity on the middle
O�CURRENCE. Unnamed beds above the
Socota Member and below the Capotes Member.
:�d ,p� of the adumbilical thirds of the flank Upper Aptian.
:�#:i :Q.ne; adoral concavity on the adventrJ
'f;lilf(ll· f:he� are projected on the venter where
1tli
. ,Ji!Y<�.I;! shghly thickened .
S
_ft �JUi.n. sp.
.9
Pl. 5, fig.B ; Text-fig. 5W, 5Y.
>;f.;,. ,' {.'fk� �horl
section is rou nd ed , with a Holotype. C-127 (U.N.C.M.H.N.C.P. Ht. N.39)
;!1���· Y�ntral.. 2!0n� (on the ribs) which tends Paratype. C-127 (tt.N.C.M.H.N.C.P.Pt. N.39,40,
·· ' .slightly ftat�ned; the flanks are convex
.

4 1).
�o\1�!;1¥, the. tunbilical wall is subvertical.
'Pl��al
or
ary nb� maY. a proach e!:,her the p A) Up to 1 8 mm D (C-127). The pro to­
tollass . . . .. yn ary as f bifureating
g p m
: . qcc
':�. ;..··'
�d ow1··0nal
! m i
·
conch is followed, after a constriction, by a
·�,ch.es
.·•. . . .. from. . ,. · . one of the secondary smooth whorl. The beginning of the second
a ,Pnm ary. Tow
, �Ill�· Primary . ard the aper­
.nbs develop small rounded
whorl is indicated by the appearance of bullae­
like tubercles at level of the involution line.
48 F. ETA Y O
These tubercles extend rapidly (half a whori Ia·
ter) onto the flank in such a way that the ado·
ral part of the second whorl is covered by raised
sharp ribs that lack any indication of tubercles;
no other indication of tubercles can be obser·
ved subsequently; at this stage the ribs descri·
be an adoral arc on the visible part of the flank;
the umbilical wall is high and rounds smoothly
into the flanks withou t Conning a shoulder.
The intercostal spaces are channel-like and wi­
der than the ribs.
On the adoral half of the third whorl the
previously unifonn ribs begin to differentiate
into prominent primary ribs, well defined
externally to the umbilical wall, and intercalary
ribs. A transition zone occurs where pairs of
primary ribs are separated by an intercalary rib.
At the ad apical part of the fourth · whorl an
alternation of primary and secondary ribs is
clearly established, although sporadically two
successive primary ribs may occur. On the last
whorl of the two specimens, the ribs describe a
feeble adoral convexity on the adumbiUcal
third of the flank and the external part of the
umbilical wall, but on the middle and adventral
thirds of the flank they describe a wide conca­
vity and finally they project in a wide arc on
venter. At that point the intercostal whorl
section is subcircular (H : 6 mm; W: 6 mm), but
the umbilical wall overhangs the previous whorl.
At this stage of the ontogenetic development
the ribs are more widely separated than on the
early �horl.
On a group of small specimens it can be
observed that the curvature described by the
ribs on the flank may be more or less accentua­
ted ; the strength of the early tubercles also
varies. On two specimens dissected to the
protoconch the tubercles on the second whorl
at the involution line are in fact periventral
nodes (=coronate whorl): at that stage the
whorl has a convex-concave section beveled on
each side; the third whorl has a transversely
subelliptical section and the fourth a subcircu­
lar to vaulted section; the overlapping of the
whorls extends only to the periventral !Ilargin
of the preceding whorl.
B) Up' to 50 mm D (C-127). On this spe·
cimen the protoconch and early whorls can be
observed although the initial ornament is weak;
the development of the ornamentation follows
the same pattern described for the smaller spe­
cimens; on the last preserved whorl the whorl
section is higher than wide with the maximun
inflation near the:adumbilical third of the flank.
The ribs are high with steep sides and sharp
tops, the intercostal spaces are at least one and
a half times . � wide as th� �ibs; the ribs are
slightly flexu�h .�e flank and they �extend
onto venter ir( li)l�oad but slightly pointed arc.
On venter all rll5s a're of equal strength and are
not thickened. Sporadically one rib may bifur­
cate . from the adventral third of a preceding
primary but no tubercles develop; the primary
ribs have a faint tendency to flare out periven·
trally.
· �ERNA
C) Up to 125 mm D (C-127). Or. this spe­
cimen the early whorls are observable and they
are similar to those of the previous specimens.
On the last whorl the ornament changes, the
ribs widen and become rounded and the secon­
dary ribs tend to connect more frequently with
the primary ribs near the boundary between
the middle and the adumbilical thirds of the
flank, or sometimes at the umbilical margin.
The primary ribs reach the umbilical seam and
are prominent on the periumbilical rim; they
have a straight to feeble sinuous profile; they
describe a clear concavity on the umbilical rim
and a broad slightly projecting arc on venter.
DIAGNOSIS. This species is characterized
by A) the transitory tuberculate stage of the
early whorls, B) the young adults with acute,
high, feebly falcifonn primary ribs which alter­
nate with one or two intercalary ribs begining
near the middle of the flank, C) the mature
adults have broad primary ribs which become
very prominent at the umbilical rim and bifur­
cate near the middle of the flarik, and D) the
changes in whorl section from rounded to sub­
rectangular.
Measurements D Euw H w D/Euw D/H No. R.
% % +
C·127 l Ei 6 6 6 40 40 42 ?.
C·127 l 6 6 6 6 37 37 50?
C-127 17 7 7 6 41 41 46
C-127 18 7 7 6 39 39 48
C-127 50 23 18 17 46 36 46+
C-127 1 26? 46 50 40 7 7 60 7
COMMENTS. This species resembles to
"Parahoplites obliquu m " Riedel [1938, pl.6,
fig. 11-12; see also under Riedelites herein),
particularly when the inner whorls of the for­
mer are considered ; on Riedel's species the ribs
become conspicuously quadrate in section on
venter but in this new species the ribs are
rounded on venter or develop a visor-like projec­
tion on the ventral median line.
OCCURRENCE. Unnamed beds above
the Socota Member and below the Capotes
Member. Upper Aptian.
This species is named in honor of the late
geologist Luis A. Perez, Universidad Nacional
de Colombia.
_A£aat!JohoDli.te..� (?ll.e.Jltacs:.rati[Qc:mg n.sp.
Pl.7 , fig.4; Text-fig. 58.
Holotype . C-127 (U.N.C.M.H.N.C.P.Ht.N.40).
Only one specimen is available. The conch
is evolute; the nuclear whorls have been dissolved
away; ·on the second whorl the ornamentation
is alrei@y' . visible · ,and characterized by .cq� �
and well • elevated flaring ribs, of which one or
two are' less elevated and appear like intercalary
ribs between primary ribs. On the third whorl
the ribs begin on the .wnbilical wall describing
a feeble concavity, then they fonn an adorai.
arc on the visible part of the flank.
 ZON ATION O F THE C R E T A C E O U S OF C EN T R A L C OL O M D I A BY AMMON ITES 49

The in tercostal spaces are channel-like,
.;, ider than the ribs. On the last whorl the ribs
are ve ry prominent. On the last third of the last
whorl, the primary · ribs develop a blunt tuber·
cle approxi m ately at the boundary between the
middle and the adventral thirds of the flank.
(:;=involution line),and ati.that point the primary
ribs bifurcate. F'rom the tubercle to the peri·
ventral margin the ribs are clearly beveled ; at
the periphery the ribs are raised in to sligh t
hurnps and th e n are depressed at th e ventral
median line . The secondary ribs h ave the same
changes but weakly developed.
The intercostal wh orl section is circular.
The umbilical wall is continuous with the flank
throughout all the stages of growth and there is
no thickening of the ribs on venter. The inter·
costal spaces-tend to widen toward the aperture.
There are. 36 ribs on the venter of the last whorl
of this specime n.
DIAGNOSIS. This species is characterized
by: A) its whorls barely in contact, B) the acute
and flaring ribs, separated by wide, channel like
intercostal spaces, and C) the adventral beveling
and median ventral depression of the ribs.
Measurements. 0 Euw H W 0/Euw % 0/H%
c . 127 30? 13 9 ? ? ?
COMMENTS. At first glance a compara­
ble species is "lloplites Ruspolii Mayer Eymar"
(1893, p. 258, pl. 2, figs. 1 0· 1 1 ) from
land, but.this species has a much more compre·
Somali·
of the third whorl groups of five to six ribs are
separated by a pair of promin-ent primary ribs
the ad apical of which bifurcateS: near the ventral
margin , they in tum are separated by a deep
" constriction-like" space; at this point the in·
tercalary ribs loose their tubercles and the inter­
costal spaces become as wide as the ribs.
The adoral portion of the third whorl h as
a circular intercostal whorl section. Each rib h as
a delicately elongated sigmoidal profile : o n the
umbilical wall the ribs have a faint adoral con­
cavity , then they form a slight convexity on the
adumbilical third of the flank, and finally a wide
concavity on the middle and adventral thirds of
the flank; on the venter the ribs are weakly ado·
rally arch ed .
On the fourth whorl all ribs h ave lost their
tubercles but the prominent primary ribs persist
as do their constriction-like deep intercostal
spaces. One in tercalary rib bifu rcates on th e
ad umbilical third of the flank o n a t least one side.
DIAGNOSIS. This species is ch aracterized
by : A) its circular wh orl section, B) its very wide
umbilicus, and C) its ontogenetic change in or·
namentation from smooth, bullate to ribbed;
the ribs differen tiating in groups of five to six
intercalary ribs, and finally equalization of the
ribbing.
Measurements.
C • 1 27 at
D
15
Euw
6
H
4
W
5 (malformed)

ssed whod•section, higher than wide, a more

�trongly beveled periven tral area with a very COMMENTS. An apparently similar spe�
'narrow median ventral zone, however the type cies from the Madagascan "Gargasian" was figu�
of ribbing appears similar. Collignon (1962, p . red by Collignon (1962, p. 6 1 , fig. 1028) as
�'l.i fig. 1027} figured as "Colo mbiceras sp. aff. "Colombiceras sp. aff. caucasicum Lupp", bu�
caucasicum Lupp." a specimen which slightly he notes that this specimen lacks tubercles: "a
i'e� rnbles the Colombian species but it lacks cotes tres nombreuses, semies, toutes simples,
tile periventral elevation of the ribs and the sans tubercule d'acune sort" ; likewise the Mada·
� �pressed ventral median line.
.-=· ·�::. _.� .
gascar species has a more irregular distribution
.
of the ribs.
OCCURRE NCE. Unnamed beds above
·

the Spcota Member and below the Capotes

M�rn ber.
The new species from Colombia also has
Upper Aptian . some resemblance to the innerwhors of some
of the Upper Aptian species of Macroscaph ites
.•. .· «nthdhoplites L'?l s�:pe'Jli
•· A c n sp Meek ( 1876), especially in the constrictions pa·
:? : . , Pl.· 3 , fig:
...
. .3
; Text-ng. 5o.
[Or'!J� . .
ralleled by two prominent ribs but the ribbing

tt?l?type. G� l27 (U.-N .C.M.H.N.C.P.Ht.N .4 1). in Meek's genus is regularly fine and the wh orls
are slightly more evolute than in my species;
� :)";.- Jialf small . specimen with the proto­
a
the development of the tuberculation in the Co·
Iombian species seems to relate it clearly to the
:cm
: �c� prnsery-:d. At th� beginning of the group of Acanthohoplites.
�ec·o . �d
..• wh orl tiny � ullae-hke tubercles appear
i:ttlpedlately · - adumbiiical to th.e invol ution line;
Jt?�; tl_t ese tu�ercles faint pleats extend t.o the
�
.� .mh,, IUc al .�am; at the incep
tion of the third
OCCURRENCE. Unnamed beds above
the Socota Member and below the Capotes.
W,.pd noticeable ribs .
are present, and some of
�J.llQ1 have_a spine-like tube rcle at
:���: .f()Uo�mg begins.
point
where
Member. Upper Aptian .

'1\'lP:e. as the nbs;

whorl

.
The ribs are �cute
-�!.·!;l.. , j;he mtercostal spaces are at least
twice as r�?�: cc:�o �l� � '";�·;;
"" m a .

;�.- ·:·�-�p()radically . one rib becomes m ore pro­

· ·.·.1•�f':.,.n. ·n'and.fan() t{ter _nb inclines
:-b
Colombiceras Spath , 1923, p. 64.

· lJ.lrca · toward it as if TYPE SPECIES. A m monites crassicostatus

ng ron1 · 1•t b �t· :v•tho
.. ut actually con­
;.<t-;l'�···c.:':•
· .

tihg. N ear the beg

.

d'Orbigny, original design a·

•

mnmg of the ado ral half tion.

50 F. ETAYO - SERNA

.,CQ.IQmQ.ik.er::fJ§. W forg.ro.t, n. sp.

Pl.6, fig.1 3 ; Text-figs. 6A, 6B, 6D.
on the venter and periven tral margins. Their
longitudinal profile is slightly flexuous. The ribs
bifurcate on one side but not on the other. The
1938 Colombiceras aff. tobleri
Jacob, Riedel, last whorl on the largest specimen studied
p.5 1, pl.8, figs.23·24, pl.14, f-7. (C-200-6) has a wide umbilicus and a regular al·
temation of primary and secondary ribs, with
1949 Colombiceras aff. C. tobleri (Jacob), the primaries slightly concave on the adumbili·
Humphrey, p. 151. cal third of the whorl. The umbilical wall is still
short but steep and rounds smoothly into the
? 1954 Colombiceras aff. tobleri (Jacob), BUrgi, flank.
p. 1 7 .
DIAGNOSIS. This species is characterized
1957 Colombiceras aff. tobleri Jacob, Biirgl, by: A) its whorl section that changes from de·
p. 135. pressed to compressed ogival, B) the ribs, sepa·
rated in groups by a constriction and a promi­
1957 Colombiceras nov. spec., Btir�l, pl.8, nent bifurcating rib in the the juvenile, chan­
fig. 2, "specimen HB2327 /1' ; recte ging this pattern into alternating primary and
HB2327/2, X 8/10. secondary ribs in the adult, the former slighly
flexuous but the latter straight and C) the mi­
1964 Colombiceras aff. tobleri Jacob, Etayo· gration of .the point of bifurcation of the
Serna, p. 1 19, pars. primary ribs from the adventral position in
the juvenile to the adumbilical position in the
1965 Colombiceras aff. tobleri Jacob , Biirgl, adult.
in Campbell and Biirgl.
Meuurernenu D Euw H w D/Euw D IH No. R .
Lectotype, Here designated, the specimen figu­ % "k
red by Riedel, 1938, p. 5 1, pl.8, C ·200 · 1 16 7 4? 6? 47 27? 28?
figs. 23-24, pl . 14, fig. 27. C-200·2 23 1 0? 8 8? 43 36 25(3 /4)
Paratype. The second specimen referred to by c-200-3 20 ? 8 7 ? 40 22(3 14 )
Riedel, 1938, p.5 1, M.G.N. Pal. 020. C-200·4 22 11 9 8 60 41 31
Hypotype. (U.N.C.M.H.N.C.P. Hy. N.8,9,10). C-200·5 43 18 15 13 42 35 40?
C-200-6 62 24 18 16 46 35 36?
A) Up to 20 mm D (C-200·1,2,4). The C-200-7 40 17 16 12 42 37 38?
innermost whorl observed (3rd'?) is coronati· C-200·8 38 17 14 12 46 37 26(3/4)
fonn, with its maximum width toward the cen· Lectotype 48 20 17 11 42 36 36
ter of the whorl side; there is a continuous roun·
ding of the intercostal whorl section; the umbi· NOT E : The measurements of D and H were
lical wall is short and vertical . There are groups made on costae ; W was measured iriter-<
of annular well raised ribs that begin at the um­ costal; Riedel's measurements of the
bilical seam, they are sharp on the flank but lectotype were taken from his paper
tend to flatten and widen on venter; adapically (p.5 1 ).
these groups of ribs are separated from each
other by a deep channel-like constriction and COMMENTS. This Colombian species re·
adorally by a prominent, slightly thicker and sembles "Parahoplites tob/eri Jacob" (in Jacob
flanging rib which at the level of the line of in­ and Tobler, 1906, p.l l, pl.l2, figs.4-6), but
volu tion develops an acute node from which unlike the species from Switzerland has juveni·
another rib branches to join a similar tubercle le ribbing (see Jacob's paper, pl.2, figs. 5a,5b)
on the opposite side, these ribs appear always characterizt.>d by its sparseness and irregular spa­
immediately adapical to the constriction; at this cing, with the prominent tuberculate ribs sho·
stage the in tercostal spaces are one and a half ti· wing a tendency to be trifurcated. Colombice­
mes wider th� the ribs. ras (?) {oreroi n. sp. differs from Colombiceras
(?) safmientoi n . sp., by the whorl section chang­
These groups of ribs consist of five, four, ing from rounded to high with flat sides and
three or two ribs, in decreasing order from the rounded venter the fonner, but on the later spe­
adapical to the adoral part of the shell, but the cies the whorl section continues rounded but
pattern seem<; variable . The rib immediately tending to develop an inclined umbilical wall,
adapical to the tuberculate rib is inclined to­ although both species seem to have a similar
ward the latter. At a diame ter of about 20 mm aspect during their juvenile stage.
the whorl section shows a definite tendendy to
increase in hight and correspondingly the peri·
ventral bifurcation of the ribs migrates towards OCCURRENCE. Socota Member. Upper
a subperiumbillcal position . Aptian.
Up to .50 mm D {C-200:7). At about 30
mm diarnerer .an alternation o,f prim ary and se­ This species is named in honor of geolo­
condary ribs becomes dominant; the primaries gist Alberto Forero of the Universidad Nacional
are p r orsi rad i a te approximately to the center de Colombia, Bogota.
of the flank and then slighly rursiradiate to the �-
venter, the second� ribs develop mainly on CQlombj£eras.16w:.mie
1l 1J1.o i n. sp.
the adventral third of the whorl. All ribs widen Pl.6, fig. 5; Text-fig. 6E, 6F.
 Z ON A TI O N OF THE CRETACEOUS OF CEN TRAL CO LOMBIA BY AMMONITES 51

? 1 9 5 7 Colombiceras nov. sp. aff.

tobleri
(Jacob ), Russian species also lacks the regular adult
·
. BUrgi, pl . 8, fig. 6a, 6b, "specimen 666", alternation of primary and secondary ribs of
rect. x 8/ 1 0. the Colombian species. The new species also has
some re�e'T'hlance to ' 'A canthoplitespo treriten­
.H.N.C.P.H t. N.42) se Humphrey" (1949, p. 139, pl. 12, figs. 11· 1 2),
Holo ty pe. C-5 13 (U.N.C.M
paratype. C-5 13 (U.N.C. M.H.N. C.P.Pt.N .42,43) but the Mexican species is more closely costate,
is compressed and has a median yeqtral depres­
. The whorl section ts depressed in �e ear­ sion of the ribs as in Gargasiceras Casey {1954).
ly juvenile it is rounded subquadrangular m the
later w horis. In the juvenile the umbilical wall is OCCURRENCE. Paja Formation (Arcillo­
_the sligb· litas abigarradas. Segment E). Socota Member.
vertical, sh ort, it rounds sl!loothly into
tly convex flank, the penventral margm rou��s Upper Aptian.
In the adult the umblh­
evenlY into the venter.
al wall is incl ined . This species is named is honor of Dr. Al­
c
berto Sanniento A., former Director of the
· on the adult, the ornament consists of Servicio Geologico Nacional (196 1-1964).
strong primary ribs which flare out on
the
flank· they are concave on the umbilical wall and ca;;;:s-�' (Gargasicer�gc:;;:"1954.-]
make' an adoral arc on the adumbilical and
Gargasiceras Casey, 1954, pp. 114.
middle thirds.of the flank, they then cross the
venter straight, but the general appearance
of their outline is as if they were rursiradiate. TYPE SPECIES. Ammonites ga rgasensis d'Or­
bigny, original designation.
on the. adventral fourth of the flank and on the
. venter the ribs are flattened. The primary ribs
begin at middle of the umbilic� wall �r a� the Gar f:icer� in teriectum (Riedel).
umbilical seam; the secondary nbs begm etther PI. , fig. 5, Text-fig. 61. ·

Independently at the periumbilical margin or

. "adjoinin g" the subsequent primary rib ; at the 1938 Acan thoplites in teriectus Riedel, p. 41,
periventral region some of the primary ribs pl. 8, figs. 8-10, pl. 14, fig. 22.
develop .a small to medium size tubercle from
which; usually but not always, a secondary rib 1954 Gargasiceras interiectum Riedel, Casey
branche� . and crosses the venter to join a similar p . 114.
tiJ.bercle on the opposite side; sometimes the
pri�ary �d secondary ribs apparently coalesce ? 1954 Acanthohoplites in teriectus Riedel, Biirgl
giving a very broad section to the venter, the p. 17.
tubercle, however, persists.
? 1955 A canthohoplites in teriectus Riedei,Biirgl
Usually there is only one long secondary p. 1 2 .
rib but s()metimes two are present. With growth
the primary . and secondary ribs regularly alter­ ? 1 9 5 7 A canthohoplites in teriectus Riede� Biirgl
qate . On the flank the Intercostal spaces are p. 135 .
. . deep and wider than the ribs but subequill to
them on venter. 1964 Acanthohoplites in teriectus (Riedel),Eta·
yo Serna, p. 119, pars.
. . DIAGNOSIS. This species is characteri·
zed by: A) its stocky build, B) its rounded sub· 1965 A canthohoplites in teriectus Riedel,Bilrgl
quadrate whorl section, C) its wide umbilicus, in Campbell and BUrgi.
and J?) Its moderate size and regular illtemation
of pnmary and secondary ribs on the adult. Lectotype. Here designated, the specimen illus­
trated by Riedel, 1938, pl. 8, fig.lO�
. Mei.lunmellta. D Euw H w D{Euw 0/H No .R. (S.G.N.Pal. 0310).
% % Hypotypes.ta Ye (U.N.C.M.H.N.C.P.Hy.N.11.
0 · 220 � 1. 3 6 17 12 1 2 47 33 1 7 (2/3) 12).
C � 2.20 - 2 23 11 ' 9 9 ? 48 39 26(3/4) Type locality. Barichara-Sube (Santander).
C - U0 · 3 28? 14 10 11 60? 36? 18(1/2)
C · 220 - 4 197 8 7 7 42? 37? 38?
A) Up to 17 mm D (La YeJ . The earliest
. · . ·· . ,POI\IIMENTS. This species was illustrated nuclear whorls studied have a w1dely rounded
.b(Z !lurgl as Colombicera
s nov. sp. aff. tobleri venter, the whorl section is wider than high and

·. 1e R,. ��o�.>.(1938,
e
. (Biirgl,p.1957, pl.8, fig. 6), Likewise
46, pl. 12, fig. S) recorded
does not show a defined periventral angle; the
ornament consists of well defined flexuous
·· . A.canthoplites aff. abichi Anth." from "Ha- growth striae which are narrower than the inter
cienda Magdalena, al Este de Viota"
, a Iocilli ty spaces between them. This non-ribbed stage is
'\'(here_ . the . same unit as that from which my
followed by the sudden development of a rib
ln1�tenal has been collected crops out "Parah o­ bearing a bullae-like tubercle rLa Ye-2-bis], ap­
P ltes abichi Anthula"
4�b�b, ( 1 899, p. 118, pl. 9, fi gs.
2 _ type speCies of Pro tacanthoplites,
proximately at center of the ftailk, from which
two flattened ribs fork which are slightly depre­
· stiongn\b t97
.. 01l• has irregularly spaced ribs with ssed along the ventral median line; the adaplcal
� rc es from �hich � secon�ary rib
.
·
bi furc tes along
a ,
· ·
· .
the lme of mvolution , the
rib is itself divided longitudinally by a cieft so
as to form two individual riblets.
52
Among the subsequent flared ribs some
of them develop minute tubercles at the level of
the line of involution. On the venter the ribs are
more distinct, due to the fact that their adapi·
cal side is vertical but their adoral side is incli·
ned. The adapical whorl section of the same spe·
cimen, at a diameter of 16mm is almost circular
(H :4 ; W : 4mm) butadorally it has already begun
to fonn a broadly rounded periventral margin.
There is a rather high and steep umbilical wall.
The primary ribs begin at the umbilical
seam as raised lines, they describe an adoral
concavity to the periumbilical margin and from
there to just beyond the middle of the flank the
ribs describe an adoral convexity and then an
adoral concavity on the adventral third of the
flank. The intercalry ribs begin at the boundary
between the adumbilical and the middle thirds
of the flank or near the lateral tubercle.
B) Up to 33 mm D [La Ye-5-bisl. On the
ad apical preserved portio ;'\' he last whorl two
types of ribs are present: one type begins at
middle of the umbilical wall and describes an
adoral concavity to the beginning of the middle
third of the flank, then they describe an adoral
convexity on the middle third, and adventrally
the ribs describe a shallow lidoral concavity.
Some of the ribs bifurcate near the boundary
between the middle and adumbilical thirds of
the flank, or more frequently at the boundary
between the middle and adventral thirds of the
whorl and have a tendency to develop a minute
tubercle there; the other type of ribs begins as
fine threads near the umbilical rim, trend simi·
larly to the others and slowly increase in size
towards venter where all the ribs have a similar
width and are separated by wider intercostal
spaces,
· DIAGNOSIS. This species is characterized
. .
by: ·A) a rather long smooth stage (with cons·
trictions?), B) a longer juvenile stage with nume·
rous sigmoidal ribs, and C) a mature stage with
mostly non tuberculate ribs with an alternation
of primary and intercalary ribs, with a tendency
for dominance of the primary over the interca­
lary ribs adaperturally.
Measurements. D Euw H w D/Euw D{H
% %
La-Ye·2·bls 17 67 7 6 36 417 30(1/2)
La-Ye·3-bls 17 67 ? ? ? 417 ?
La-Ye-6-bls 28 10 10 11 36 3 6 30(1 /2)
La•Ye-5-bls 33 14 13 14 42 39
cm.l.ll\1EKTS. This species differs from
qargasiceras pliJc'J;fp JRiedel) by t�e oval sec­
tions of tile la��h�te� tile fQmler has a
rounded subquadrite one. In G. pulcher (Riedel)
the bifurcation of the ribs occurs regularly near
tile middle of the flank and at similar diameters
the ornament of prominent bifurcating ribs is
still present but the opposite is true for G. inte­
riectum.
OCCURRENCE. Socota Member. Upper
Aptian.
F. ETA Y O - SERNA
Gargasiceras interiectum (Riedel)
(Morph I)
Pl. 6, fig. 8; Text-fig. 6H.
Hypotype. La-Ye-4-bis (U.N.C.M.H.N.C.P.N.13)
A) Up to 13 mm D [La-Ye-0]. The adapi­
cal half of the last whorl appears smooth but it
is covered by fine , low flexuous riblets, they
cross the venter uninterruptedly; the ribbing
becomes prominent rather abruptly on the ado·
ral half of the whorl; the ribs start at the umbi·
lical seam and make a delicate biconcave arc to
tile venter, they flare out on the adumbilical
third of the flank and are sharp; on the adven­
tral third of the flank they are flattened and
on the venter a bit wider and slightly depressed
along the median ventral line. One of the early
ribs has a weak tubercle at level of the involu·
tion line where tile rib trifurcates but on the
other flank no tubercle is developed and the rib
only appears broader. An alternation of long
primary ribs and shorter intercalary ribs begins
almost contemporaneously with the appearence
of the ribbing.
B) Up to 40 mm D [La-Ye-4-bis]. The
whorl section is subrectangular and has broadly
rounded periventral margins and feebly arched
venter; the ornament consists of long primary
ribs which begin near the umbilical rim, these
ribs develop a flange on the ad umbilical third of
the flank, they are high and sharp; the intercalary
ribs begin at the boundary between the ad umbi­
lical and the central thirds of the flank. All the
ribs become flat topped and slightly broader on
the adventral fourth of the flank and venter, they
have vertical sides; the intercostal spaces are
twice as wide as the ribs on the internal mold.
The umbilical wall is short and subvertical.
NOTE : This morph has a well defined altema·
tion of primary and intercalary ribs on
venter and the subrectangular section
seems constant throughout the ontoge­
ny .
Measurements D Euw H W D{Euw D/H No.R
% %
N o ,R. La·Ye-D·bill 13 6 6 6 46 38 1 6 ( 1 {3)
La-Ye·4-bls 40? ? 14 1 4 ? 36? 2 3( 1 /2)
COMMENTS. This species differs from
30( 1/2) Riedel's "A canthoplites in teriectus ", as he illus·
trated it where the inception of tile intercalary
ribs is on the adventral third of the flank. This,
however, could be a a matter of preservation.
However-, the ribs are distinctly more flattened
.in "mQIJ?;NJ.'! tlum on Riedel's specime� . Rie!fe�j: :
remark '(1938, ·pAl): 1'En esta espec1e no hay:
bifurcacion de costillas ni fonnacion de tubercu­
los" as here understood indicates that the tu·
berculate stage is a transient one terminated very
early in the ontogeny as pointed out by Casey
(1954, p. 114); it Jll!ISt be added that the
ven tral depression (bu-t not interruption) of
the ribs along the ventral median line is another
transient characteristic.
 ZON ATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES 53

OCCURRENCE. Socota M�mber�:;� Upper

·
well defined, rounded #"ff;·�--er than the in­
Aptian. . .� tercostal spaces. The pnmaty.�'bs are very pro·
Gargasiceras 'i&{orph (Riedel)1
cher:
I)
minent 11nd h�come well define;: at the external
margin of the umbilical wall.
On the adoral half of the third whorl the
1938 Acanthoplites pulcher Riedel, 1938, number of intercalary ribs decreases from two
p.43, pi.S, figs. 11-14. to one; on the last preserved whorl there are
c alternating primary and secondary ribs. The
?1954 Parahoplites cf. pulcher Riedel, Biitgl, whorl section is subrectangular with slab-like
pl.17. sides and rounded venter, the umbillcal wall is
inclined and merges smoothly into the flank.
1954 G. [a&asiceras] pulcher (Riedel), Casey, On the adoral third of the last preserved whorl
114. the intercalary ribs disappear and the resulting
intercostal distance is five times as wide as the
t
1964 Parahopli es pulcher {Riedel), Etayo-Ser­ ribs. The apertural margin is plain.
na, p. 119.
DIAGNOSIS. This species is characterized
1965 Acanthohoplites pulcher Riedel, Biirgl, by: A) the progressive ontogenetic elimination
in Biirgl and Campbell. of the intercalary ribs, B) the subrectangular
slab-sided and rounded venter whorl section
L6ctotype. Here designated, the specimen illus- and C) the adorally arcuate . trend of the ribs
-
•.. .•._... .
. ..
· trated by Riedel, 1938, pl. 8, figs.
11,12,13,
}'aratype. Riedel's pi.S, fig.14, S.G.N. Pal.0310
·.
on the flanks .
Measurements D Euw H W DfEuw% D/H %
Hypoty.pe. La"Ye-7-bis (U.N.C.M.H.N.C.P. Hy. C·151 29 13 11 10 45 38
N.14). ·

COMMENTS. This species resembles "Pa­

rahoplites abichi Anthula, 1899" rtype species
of Pro tacanthoplites Tobvina, 1970 J, but on
the latter genus as interpreted by Tobvina (En­
glish trans., 1971, p. 338) "Sections of whorls
transverse-oval, tumid or practically subquadra·
te, width invariably exceeding height.", further­
more the primary ribs bifurcate from the lateral
tubercles, character'.stics that are absent from
the Colombian species; the Russian and the Co­
lombian species have in common, however, the
broad umbilicus and the tendency of the ribs
to flatten on venter "to varying degrees" (Tob·
vina, op. cit., p. 338); the Caucasian species has
constrictions that are also absent from this spe­
cies. Colombiceras crassicostatum {d'Orbigny)
(1841, pl.59, fig. 1·5) has only tiny lateral
tubercles and its ribs are wedge shaped on the
flank.
"Hypacanthoplites ? rursiradiatus Hum­
phrey" (1949, p.142, pl. 14, figs. 11, 1 2) l type
"morph" resembles species of Pegnaceras Cantu-Chapa (1963J, is a
fptu_cller. ·tl'l.le(letJ, which WllS des­ species on which "all ribs thicken in transverse­
extem.n • • • anchamen­ ly elongate swellings" (Humphrey, op .cit . , p.
the _latter differs 142) a characteristic not observed on G. juan­
section is oval. wyatti.

One specimen that has some resemblance

to the present species but which has intercalary
ribs to a larger diameter than in the present spe­
cies was figured by R. Douvilllt (1906, p.147,
note, pl. 1, fig. 3) as Pedioceras or a new genus
from an unspecified locality in Colombia.
OCCURRENCE.Upper Aptian.
This species is named in honor of Profes·
sor John Wyatt Durham.
Genus
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
1976 Douvilleiceros mammilal tum (Schlotheim),
Etayo-Serna et al, p. 227 .
1916 Douvilleiceras aff. mammilal tum (Schlo­
theim), Etayo-Sema et al, p.225, tab. 2.
Holo�pe. .(;�8 (U.N.G.M.H.N .C.P.Ht.N .44 )
PiiitYifl.! . (U.N.C.M.H.N.C.P.Pt.44,45).
Two collections (C-13 and P-21B) have
been studied and it is considered tfiat the spe­
cbnens axe conspecific. In my opinion these
specimens overlap in the development of the
omament.
A) Up to 15 mm D [C-218). The earliest
omamen t observed on the adapical portion of
"this specimen consists of widely spaced fold·like
ribs which describe a bend from the umbilical
margin to the middle of the flank, at which
point there is a spine like tubercle , the ribs des·
cribe a feeble adoral concavity and extend onto
venter. A transversely elongated inflation is pre­
sent on both sides of the ventral median zone.
At the adoral end of this specimen (D: 15mm]
a small tubercle is present on the nbs at their
periumbilical margin and the ventral tubercle
beeomesmammiform, but no indication of any
division of the ribs is obsenrable on the ventral
· tubercles. The intercostal growth lines have a
profile similar to that of the ribs. The whorls
Qrt! depres&ed. No clear indication of the ventral
tubercles appear before a diameter of 24mm.
B) Up to about 35 mm D [C-218]. The
omament is stronger but In the previous pattern
_
�e ribs are widelf spaced and a weak annular
iritercalm:y rib, aliriost imperceptible on the in·
:temal mold, occurs midway between the primacy
Jibs� I:rnpressions left ··on - the umbilical wall of
·
.·- .
ttre: subsequent volutions indicate that at this
point the lateral tutlercles represent the base of
long spines, directed adapically. About 30 mm
_ c . 218 . 2
- �di�eter a tiny · tubercle appears on test exter·
!:laJIY to theperiumbilical tubercle but it seeming­
�� �does no t leave any indication on the ln�amal
�Qld at this stage.
C) Up. to 45 mm D [C·13J. A single speci­
>lll en Qn which the Internal whorls can not be
_ d
.!!�died._ The first rib visible on the ad apical por­
· tls»n ofthe ·last whorl arises near the external
·'Part of the umbilical wall and has a minute tu·
: ��rcle at the P riumbilical rim; slightly adven­
! ruvian species the tUbercles tend to be equal
J�IJIIY to the middle of the flank a prominent
\ : � ersely _ elongated node is present on
>b.o� sldes of venter well marked tubercand les are
<P�n.t. The adumbilical and the medial tllber­
i=Jea are connected by the bar-like rib� but bet­
:e
; e,en the lateral and ventral tuber
cles as well as
_ tween the ventral
· .
tubercles the ribs are de-
J���d. .
:;: ::Di. Th e Jibs are
widely separated by in r
::1!1!1;!1! lJ?aces about two
tll.rlb b_ , but an almost tmpercepti
�d a half times as wide
s
X .t-.ry ble interca­
\Tb-i.s J.. IS
Jl�sen,t between �e principal ribs.
\Qlild k, pe
ifui' . 0� the length of thecontin
of sculpture ues on the first
last whorl; the
: c': J· 0 a change is indicate
begin­
d by l:he develop-
55
ment of bipartite ridges on the adventral
margin of the ventral tubercles and by the pro­
gressive inflation of the ribs between the central
and the penumbillcal tubercles, but the latter
are now weak bullae along the periumbilical
margin .
.
A t the adoral end the ribs have six
tubercles on each flank with the centrolateral
one the most prominent. The intercostal whorl
section is smoothly rounded, depressed, reni­
fonn. The umbilical wall is inclined, the ribs
describe a feeble adoral arc on the central third
of the flank with a short weak concavity near
the umbilical shoulder; the ribs cross the venter
straight. In general the ribs appear rectlradiate.
D) Up to 86 mm D [C-218). On the last
whorl of this specimen the ribs begin near the
umbilical seam, they trend adapically on the in·
cllned umbilical wall and cross the flanks and
venter in a slightly arcuate manner. At the ada·
pical portion of this individual the ribs have six
tubercles, of which the umbilical one, the centro­
lateral one and that between the preceding ones
are mammillated whereas the other three are
spirally elongated and more related to each
other than to the tubercles on flank, but
toward the apertural end all tubercles equally
spaced . The median ventral sulcus Is a deep
channel. The intercostal spaces are one and a
half times the width of the ribs, and the ribs are
prominent and flanged. There are 27 ribs on the
last whorl.
DIAGNOSIS. This species is characterized
by: A) its prominent ventral tubercles, B) its dis­
tant flange-like ribs, and C) the profile of the
ribs which appears corrugated .
Measurements. D Euw H w D{Euw% D/H %
c . 218 · 1 16 5 6 9 33 40
31 15 10 15 48 32
c . 13 46 25 14 20 66 31
c . 218 · 1 0 86 46 28 35 52 32
COMMENTS. This species Jiiffers from
the one identified by B�a.'!i.AA$.�q�s_�,!(195 6,
_
�· 442, pl.43, fig. 1·4), ·as 15ouufflezceras momle
(Sowerby). On the Colombian specimen the ribs
have a wider intercostal spacing and there is a
persistent difference between the internal and
external tubercles on the flank whereas in the Pe·
in size, a crowding of l:he ribs occurs at an early
ontogenetic stage. IJ�.monile (J."SQweJ!2Yl �-n'-i
described b y Spadi (1923, p. 72) �d_J;:asey
_ �
(1962, p. 284) differs from the Colombian spe­
cies because on the latter the intercostal distance
is very constant and the tubercles are very well
differentiated.
te
OCCURRENCE. Capotes Member. Lower
Albian.
­
������,��-�!- �.:- �_p_.
. -
Holotype . C · 215 (U.N.C.M.H.N.C.P.Ht.N.45).
Paratype . C 215 (U.N.C.M.H .N.C.P.Pt.N. 46).
•
56 F. ETAYO · SERNA
A) Up to 18 mm D [C-215-21]. To this
diameter the internal whorls have not been ob·
served. The adapicalmost part of the last preser­
ved whorl has a subpolygonal·renifonn intercos·
tal whorl section: a rather abrupt umbilical wan
which rounds smoothly into the flank; the ad·
ventral segment of the flank is broadly rounded,
the venter is narrow with a flat ventral median
band.
The ornament consists aflaw flattened ribs
which begin at the umbilical margin and bear a
rhomboidal prominent tubercle that is spirally
elongated. From this tubercle to the periventral
margin the ribs broaden and become depressed;
along both sides of the ventral median band the
ribs develop radially elongated crest-like t,uber·
cles; on midventer the ribs are depressed, al·
most obliterated but have a tendency to split
transversely. Sporadically an intercalary rib
may develop, it begins at level of the lateral
tubercle and branches from a strong primary ri�
this secondary rib has only ventrolateral tuber·
cles: it is so obscure on venter that the tubercles
appear as if they were independently arising
from the test.
The intercostal space between the primary
ribs is very wide, up to two and a half times the
width of the primary ribs except when an in·
tercalary rib is present; this interval is covere.d
by wen marked growth lines which are adorally
concave on the umbilical wall. Towards the
adora1 half of the last whorl the lateral tubercle
occupies a medial position on the flank and
small conical tubercles appear on the ribs at the
periumbilical margin.
. B) Up to approximately 25 mm D [C-215·
22]. At about a diameter of 20 mm the spirally
elorigated periventral tubercle acquires a mammi·
Dated . shape, and contemporaneously these tu·
bercles develop adumbilically an incipient tuber·
cle that with subsequent growth tends to match
the adventral one in prominence. The spacing
between the upper and lower ventrolateral tu·
bercles varies. The lateral tubercle increases in
size and develops a large circular base. At this
stage the intercostal whorl section has a broadly
flattened to slightly arched venter which rounds
Into the flattened flank; the umbilical wall Is
oblique, the periventral tubercles are slight
bumps on the test and they rarely show on the
internal mold.--en the last rib preserved of this
specimen there is a faint swellin� between the
centro-lateral and the periumbihcal tubercles.
.
C) Up to 95 mm D [C-215·251. Lacking
any in tennediates I am interpreting this speci·
men as conspecific with the previously descri·
bed specimens, based on their apparent simUa·
rity. At this stage of growth the ribs are thin,
high, with rounded top and separated by wide
intercostal spaces. The ribs begin at the exter·
nal part of the oblique umbilical wall as elonga·
ted and flattend lozenge-like subdued elevations.
The ribs are slightly convex adorally on flank
8lld althoul!h a bit wider on venter their width
on the fl8lllts is ve:ry uniform. On the adapical
half of the last preserved whorl there are five
tubercles on each side appearing like symmetri·
cal ripples on top of the ribs, and distributed as
follows: one ventral slightly elevated and rna·
mmillated tubercle, one on center of the ventral
shoulder, another on the rim of the ventral sho·
ulder, a tubercle in the middle of the flank and
one adumbilical tubercle. The lateral tubercles
are equidistant from the one at the middle of the
flank. Toward the aperture there is a progressive
development of another tubercle close to the
ventral median line in such a way that the ini·
tial ventral sulcus almost disappears; although
there is an increase in size of the tubercles from
the umbilical to the ventral, in general the tu­
bercles tend to have a uniform size There are • .
30 ribs on 3/4 ths of the length of the 'last whorl;
the living chamber occupies half of the last whorl
DIAGNOSIS. This species is characterized
by: A) the development of a delicate tubercula·
tion of the ribs which appear sllgh tly undulated
in longitudinal profile, B) the rounded-subqua­
drangular whorl section, and C) the very shallow
and narrow ventral median sulcus in the adult.
Meuurementa. D Euw H W D/Euw % D{H%
C • 21 5 · 21 18 9 ? ? 50 ?
c . 215 . 2 2 26 11 10 12 44 40
c . 21 5 - 2 4 9 5? 46 ? ? ? ?
COMMENTS. The specimen figured by
Collignon (1963, p. 112, fig. 1243) as Douvi­
lleiceras albeme Spath, has a similarly reduced
median ventral sulcus but its tubercles are cle·
arly clavate and its ribs arise in pdirs from
accentuated periumbilical tubercles, the two
latter features are absent from the Colombian
species. The specimen described by Riedel
(1938, p. 32, pl.6, fig. 6,7) as "Douvilleiceras
monile Sow ", and excluded from the scope of
Sowerby's species by Casey (1962 p. 288),
resembles Douvilleiceras abozagloi n. sp. in the
loss of the ventral sulcus and its rounded ven­
ter, but on Riedel's figured specimen the rib·
bing is coarser and the inner lateral tubercles
postulous. The specimen (U.S.G.S.No.23836)
identified by Imlay as "Douvilleiceres aff. D.
monile (Sow.)" from locality Santa Rosita I,
Venezuela (in Rod and Mayne, 1954, p. 267),
from examination of a plaster cast, seems to
belong to a related species; it has wider ribs and
longer and better stressed clavi, further the ven·
tral median sulcus is persistently wider than in
the Colombian species.
Douvilleiceras solitae (d'Orbigny) has a
more persistent and deeper ventral median sul·
cus, the ribs are broader and the ventrolateral
tubercles larger than in D. abozagloi.
OCCURRENCE. Lower Albian.
• This species is nemed in honor of the geo·
logist Jacob Abozaglo M.
\,
_
_
�� �]""���""�Ql10u_�-0 �-���-·��-�. ,_.""'\'}��!�!�2;
.. ...
. ... .
ras
-
Eodouuilleiceras Casey, 1961, p. 191.
 ZONATION OF THE CRETACE O US OF CENTRAL COLOMBIA BY AMMONITES 57

TYPE SPECIES. Douvilleiceras horridum Rie­ Biirf (1961) recorded as occurring

del, original designation. together 'Douvilleiceras mammillatum (Schlo·
theim)" �::! "nouvilleiceras horridumRiedel",
o l s e d&Q,a'if&.l!lb_E_:_.�P·
-Jt� 'fi1g.i fj(:, T�
from Ortega (Depto. Tolima); see synonymy.
, ext�rf
1gs. , .
A!_Pian.
•

OCCURRENCE. Lower

Douvilleiceras horridum Riedel, BUrgi, p.

-"'•

1961 This species is named in honor of the late

�5 ex (U.N.C.M. H.N.C.P.N .1843). geologist Pedro Carvajal •

1964 Douvilleiceras horridum

Riedel, Etayo­ . Eod�uv illejcerp.s · t uendamai. n . sp.
Sema, p.120 pars ex Btirgl 196 1a. ��. 't'ext·i?
gs. 6j{,60,13. · -

197
.
6 Eodouvilleiceras horridum
(Riedel), Eta­
yo-Sema et al, p.225 , tab.2, p.227, ex
(U.N.CM.H.N.C.P.N.18 43).
Holotype. C-170 (U.N.C.M. H.N.C.P. Ht. N.46).
Only one specimen is available; the inner­
mqst whorls are not observable. On the 3rd (?)
�h.orl of the phragmocone prominent spines
develop about the middle of the flank and the
�horl appears coronate ; the ribs they surmount
� Wide and low and start at the umbilical rim;
tOw-ard the adapical portion of the following
whorl, tiny conical tubercles develop at the pe­
riumbilical end of the primary ribs. The umbill­
ea( wall Is rounded, subvertical and merges gra­
dually into the flank. The intercostal spaces are
wide, variable in with, one and a half or t'vo ti­
mes as wide as the ribs.
The intercalary ribs occur at irregular in­
teiVal.s, they h ave only centrolateral and per­
ivel).tnd tubercles, and appear like slight folds of
tile test. On the primary ribs ventrolateral tu­
bercles enl&rge into strong bosses which bear a
parallel,lower and sligh tly indicated clavus. The
whorl section has a rounded, lozenge-like shape.
•. . .
·. . DIAGNOSIS. This species is characterized
�y: A)Jts compressed shape and its lozenge sha­
.. .
Holotype. C-233 (U.N.CM.H.N.C.P.Ht. N.47)
Paratype. C-22 3 (U.N.C.M.H.N.C.P.Pt. N.47,48)
A ) Up to 26 mm D (C-223-17). On the
innermost whorl observed (2nd of the phrag·
mocone) there are long spines along the involu­
r
tion line; short and long s ines alternate in a w
pattern; the impression o the spines is visible
on the dorsal part of the following whorl as de­
pressed furrows. The long spines arise from wi·
de ribs that reach the umbilical rim, but the
intercalary rib corresponding to the short spine
disappears without reaching the umbilical rim.
The third whorl has a high and vertical
umbilical wall which rounds abruptly into the
convex flank; the adapical portion ofthe fourth
whorl has ribs that bear a small conical tubercle
at the umbilical rim. The ribs appear like prorsi·
radiate blunt folds: the portion of the ribs
between the umbilical and the lateral nodes h as
th e shape of an isosceles triangle; from th e
centrolateral tubercle to th e ventral tubercle
the ribs are wide and have subparallel sides,
they tend to cleave longitudinally along their
central axis. In middle of venter the ribs are
wide and depressed. The intercostal spaces are
approximately one and a half as wide as the ribs
on the flank but twice as wide on venter.
At the umbilical rim the tubercles are

ped whorl section, B) the accentuation of the tiny, and elongated in the direction of the ribs;
ventral tubercles which defme a prominent U­ the ventrolateral tubercles have an elliptical
�!lped sulcus, and C) the feeble development section elongated in the direction of coiling but
.of' the intercalary ribs. a subcircular section Is also common. The
�wth lines may be so accentuated as to give
Ma�meu.t.. D Euw H W D/Euw % D{H % the impression o f being Intercalary ribs.
C · 170 "0 211 18 22 110 36
B) Up to 48 mm D (C-223). With increa·
sing diameter there is a tendency for the ribs to
C C? �ENTS. This species differs from increase their wi:ith without a corresponding in·
�odouurllerceras honidum {Riedel), which has a crease In heigh t. An accentuation of the peri­
sJower accentuation of the tuberculations, broa­
� Whorl section and wider and shall ower ven­
ventral tubercles occurs on the living chamber
for two thirds of the length of the last whorl.
sulcus than In E. pedrocarvajali,
finally the Toward the apertural segment. of the last pre­
··�· . n:alary ribs persis
m te served whorl the tubercles attenuate; the ven­
t longer.
trolateral tubercle becomes compressed, bullae·
llke. The last rib is thin, barely visible with only
>'" ·
fi 261; Text-fig
'I'h,e
specimen Illustrated by Casey (1962, lateral and umbilical conical spines aiid without
"E. horridum (Riedel),
u
o:&er . 90)
�ptian, near Bogota, Colombian (Umv.
ventral tubercles. The apertual margin is plain.

CiUv no �� .C-1563)" is more like E. pedro­ DL\GNOSIS. This species is characterized

by: A) Its very depre����ed whorl section, B) the
RiedQJf!1
e 8
than llke Riedel's species. Both
species and �asey's fip1red spP. imen
l9381UlY3�tr
'�es):, !,�!f8Phlc
inconspicuous bullate periumbilical tubercles,
lilclt �
m formation (see R1edel, the long periventral spines and the hom llke
P· Univ. ; llllK1 California, Mus. Paleont., ventral tubercles, C) the dominance from early
In the ontogeny of the primary ribs 'With almost
 58 F . ETA YO · SERNA

no intercalary ribs, and D) the lack of bifur· flank, but this displacement does not have a re­
cation of the primary ribs and their offset longi· gular starting point.
tudinal prome.
At . a height of whorl of 9 mm (D: 11
MeUUftmenta D Euw H w D/Euw D fH No.R. mm), corresponding to the adapical portion of
% % the last preserved whorl, the periventral clavi
C·223·17 26 12 11 1 7 48 42 12 are connected across the venter; at that stage
C·223·18 ao 13 12 20 43 40 13 the ribs describe a wide adoral convexity to the
C•223·19 22 12 8 17 �0 36 12 middle of the flank and from that point to the
C·223·20 2& 12 11 1& 48 44 1 37 ventral margin they describe a deep adoral
C·223·21 48 23 19 27 48 39 16 concavity; the ribs are straight or slightly
concave adorally on the umbilical wall. The ribs
COMMENTS. This species resembles tend to broaden from the adumbilical rim to
Eodouvilleicerus horridum (Riedel) (1938, p.29, venter; the transverse profile of the ribs is roun­
pl.6, figs. 1,2, pl.14, fig. 1 1), but on the latter ded with its adoral side long and oblique, the
at the equivalent diameter the lateral tubercles adapical side is short and steep. On the shell the
are leas developed and the intercostal distance is intercostal spaces are slightly wider than the
greater; in Riedel's species the ventral tubercles ribs. The early portion of the last whorl is sub·
tend to develop an additional tubercle early In quadrate, truncated at venter, slightly wider
the ontogeny (see lower right of figure 2 of toward the adumbilical third of the flank;
. plate 6 of his paper), he noted, however, approaching the aperture however, the whorl
that because of the poor state of preservation section is slightly higher than wide. The siphon
of his specimen, the inital appearance of divi· is so close to venter that is creates a tenuous
· ded external nodes could not be determined. raised line along mid·venter.

Eodouvilleicerus (?) tequendamai differs DIAGNOSIS. This species is characteri­

from Cheloniceras (Epicheloniceras) barreroi n. zed by: A) its bold ornamentation, B) the change
sp., by the ribs bifurcating from the centrolate· in whorl section from subquadrate to subtrape·
ral tubercles and the ventral tubercles changing zoidal; and C) the strongly falcoid ribs on the
from mammllate to transversely elongated early flank, closely spaced on venter.
in the ontogeny of the latter. The juvenile
Eodouvilleiceras differs from Ch. (Epicheloni· Meuuremanta. D Euw H W D /Euw D(H No.R.
ceras) leonhardtriedeli by the weaker ornament % %
of the latter and subrentangular whorl section. C · 22 3x • 10 44 26 19 16 67 43 3&
OCCURRENCE. Socota Member. Upper COMMENTS. Du{renoyirz scalata Casey
Aptian. (1964, p. 386), is a similar species. Casey's.holo·
type (op. cit., pl. 66, fig. 1a, b) has intercostal
Family . �E�AYE�ITIDAE
GenUs
• .
· (Du
. ·-··
n 9if
1 an and ebo�l. 1915
_ _
(Burckhardt MS) Ki· \
. _

•.. · --·----�
__
spaces narrower than those of the Colombian
species but the ribs are wider; on Casey's species
the ribs also do not have a slight adoral conca·
vity.
Dufrenoyirz (Burckhardt MS) Kilian and Reboul,
1915, p.34. OCCURRENCE. Socotai Member. Upper
Aptian.
TYPE SPECIES. Ammonites {urcatus J. de C.
Sowerby, by monotypy. This species is named in honor of Pro­
feasor Gerardo Botero Arango, of the Universi­
..[).yfre@yig pQt..emiJ: . sp. dad Nacional, Medellin .
Pl.1, fig. 6; Tex-figs. 5M:, 5 N.
Du(renoyia codazziana..(Karsten)
Holotype. C-223x-10 . (U.N.C.M.H.N.C.P.Ht. Pl. 1, figs. 1, 7, 8.
N.48).
1858 Ammonites CodiJzzianus Karsten, p.
The hdotype has thej>rotoconch preserved; 108, pl. 3, figs. 4 , 5.
the conch is robust and has six volutions. To
approximately . an umbilical diameter of 5 mm 1881 A T mmonites] Codazzianus , Karsten,
the flanks have broad convex bulges separated GaSb, p. 310.
by narrower depressions; the umbilical wall is
vertical but it rounds smoothly into the sides. 1886 A mmonites Codazzianus, Karsten, pl. 3,
At about the above umbilical diameter a second figs. 4, 5.
and fmer inflation devel6ps by longitudinal
splitting of uie liu-ger bulge just outside the um­ 1913 Par. Tahoplites ]Codazzianus .Karat.,Kilian,
bilical margin . At an umbilical diameter of 8 •
p. 344.
mm the newly appeared swelling (now a rib) de·
taches· from the primary (earlier) rib , and about 1925 A mmonites codiJzzianus Karsten, Burck­
an umbilical diameter of 13 mm the secondary hardt, pl. 10; fig. 10·11.
rib occupies an intermediate location between
the primary ribs, at the same time it has began ? 1937 Colombiceras Codazzianus Karsten, Bo­
to move outwards toward the middle of the tero, p.26, fig. 5 0,51.
 zoNATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES 59

1 94 7 Ammonites Codazzwnus Karsten,Spanish the ribs have an accenb.lated adoral ar c on

version , p.375, pl. 3, figs. 4,5. venter throughout th e on togeny .

1964 D. f ufrenoyUI Jcodazzwna Karsten, Casey, DIAGNOSIS. This species is d i fferentia­

p. '397. ted by: A) its fine, uniformly wide, sickle lil�e
rib s t h ro u g h o u t the ontogeny, B) the early
1964 Deshaye$ ites codazzianus (Karsten),Eta­ ontogenetic attenuation of the periventral no­
yo-Sema, pars., p . 1 20 . des and the broad adoral arc of the ribs on ven­
c ter, and C) the nearly constant suboval whorl
Du{renoyUI codazzwna (Karsten), Etayo· section.
. · 1968
•

Sema, p. 28, fig.3.

1972 DufrenoyiD coddaziana (Karsten) , Etayo·
Sema, p. 43.
UC::to·.· cyp e. Kazsten's fi&1lred specimen under
·• • ·
·
num ber 4 plate 3, 1858.
l>aratype. . Karsten 's figured specimen under
- number 5, pl ate 3, 1858.
J:{ypocype; (U.N:c.M.H.N.C.P.N. 16,17).
'The. whorl section is higher than wide ,
subelliptical; . the u mbili cal wall is subvertical, it
rounds smoothly into the gently convex sides;
the . maxjmum thickness of the whorl is at the
boundiuy between the central and the adumbi·
Uc!J} thirds of the flank, externally to this point
the sides a:re flattened and slightly convergent
on the adventral third of the flank. The ·1enter
is .relatively narro w, truncated. The umbilicus is
Shallow, .abot1t one half the width of the diame·
ter;
The ornament on the early portion of
the last whorl consists of fine, rounded and
untfonnlY wid� ribs, slightly narrower than the
h1tereostal spaces . On the umbilical wall the
ribs ate weak and d escribe a shall ow adoral
i:qticmty, then they trend prorsiradiate on the
.adumbWcal thiid of.the flank ; at the boundary
.between the adumbilical and central thirds of
tlte flank they describe a crescentic adoral
;,Cp��ilvity. At �e periventral margins all the
·J!bS'have slleht pyrifonn swellings, then the ribs
!JWIJig adorai!y describing a wide and prominent
arc r the venter :n!mains narro w. At the bounda·
',ry b!ltween the adumbilical lind central thirds
qf 1:11 � flank the ribs bifurcate, the point of
9l,!Utcation � �in g slightly inflated ; commonly &
•·$9co��IIP' n�· �ppears. without a connection to
, e �Jacent pnmary nb
th
.;:\:?·:;· :�; -· · : .
•
Meuuzements. D Euw H W 0/Euw% D/H o/o
La · Ye • 2 X3rt 14 111 11 40? 43?
COMMENTS. The small specimens are
(pl.l, fig.1,7) comr. arable to ''Ammonites
Codazzianus Karsten' (1858, p. 108;•pt 3, fig. 5
especially ) and to th e photograph of the plaster
replica ot Karsten's original (op. cit., pl. 3),
given by Burckhardt ( 1925, pl. 10, fig. 10-11).
Karsten's figures (op. cit., pl.3, fig. 5a, 5b) are
res tored and composite. [J. Helms , written
communication, 1976) .
Du{renoyia formosa insculpta Casey
(1964, p. 396, pl. 66, fig. 9,9a) is a comparable
tonn but on the latter the ribs are only slightly
concave on the adventral third of the flank and
the primary ribs a:re crowded arou nd the
um bilical rim. Casey (op. cit., p. 397) compared
his subspecies to D. codazzwna Karsten, poin·
ting out that the Colomb ian species has "denser
ribbing. . . and the venter has broader ribs and
sharper margins".
Dufrenoyia mulatoensis Hump hrey (1949,
p. 127 , pl. 9, figs. 11, 12) has less flexuous ribs
an d they are barely adorally arcuate on venter.
Gerhardt's "Hoplites stuebeli" has distinct and
progressively separated ribs o n flanks at a relati·
ve small size.
Duftenoyin wnctorum Biirgl , nom. trans.
PI. 1, flg. 2; Text-fig. 14.
1956 Dufrenoya texana sanctorum BUrgi, p.3,
p.6, p . 12 ; pl .1, fig .2, fig.3; pl.3, fig.3,5.
? 19C4 D. [ufrenoyw] texana sanctorum BUrgi,
Casey, 1964, p. 386.

1964 Dufrenoya texana sanctorum BUrgi , Eta­

: :� ,·• .. On the internal �old , all
-�"!�<a :J:Q�nded top and abrupt adapical side and
the ribs
yo-Serna, p. 121.

1968 Dufrenoyw justinae (Hill), Etayo-Sema

:�-,��hque . bu t short adoral side ;
the ribs are p. 28, fig. 3, tab. 1.
�9re acute and raised on the &dumbilical third
�:�e flank where.a� the� are slightly flat· 1976 Dufrenoyia justinae (Hill), Etayo-Sema
IS�� , oil the rel!lammg third. When the shell
'thf�,�rved et. al, tab. II, p.225.
ibi{l!��mal the nbs look like narrow belts. On
mold the ribs � narrow er than the Lectotype. Here designated. Specimen HB.406/1,
;Uit:e · 0.!iti!.l spaces, especially on venter; the
� �t� �!�es
- �
� l!"
.
are deep along the adap1cal
figured in Biirgl 1956, pl.l, fig. 2.
J
:·-·. : · -�·, .. " : ·::.---- ..
Type locality. Mesa de los Santos (Santander).
Hypotype. (U.N.C.M.H.N.C.P.Hy. N.18,19,20).
)�tdlll
�i�: T()�lr W
a horl hei � t of 36mm the
ribs still
characteris¥ patte rn : acute adumbi
A) Up to 38mm D lC-223x-17]. The
Ilij• fJanm()cjer&tel� � · whorl section is higher than Wide, slightly wider
mclmed adventrally and toward the umbilical third of the flank; the
crescen�c on venter, bu at
; ; />
: tl:l'i& ; �e
�di t about umbilical wall is subvertlcal and rounds abrup·
· ter then profile b
ecomes rounded ; tly into the sides; the p eriventral margin is
60 F. ETAYO - SERNA

subrounded The ornament consists of ncrrow A) Up to a diameter of 31 mm (C-223x-1;

ribs which begin at the umbilical &.eL'll, they
describe an adoral concavity en the umbilical
wall, on the flank they describe 8ll l!doral are
on the internal half and a conei!Vity on the
external half; the rib& tennlnate in prominent
rounded, elongated clavi, the ribs are intercon·
·
La-Ye-1). The adapical section of the whorl is
higher than wide and it has a moderately wide
ventral region, the periventral margins are broa·
dly rounded, the flanks are moderately convex
to their central third, from this point to the
umbilical margin the sides are more tumid, they
round into U1e subvertical umbilical wall.

nected across the venter.

On the internal mold the primary ribs
are represented by elongated awdlings corres·
ponding to flanges on the test, especially on the
adumbilical third of the fl&nk, the ribs contract
about middle of the flank �md from that point
to the ventral margins they flatten in the fashion
of a propellor blade. The intercalary ribs appear
near the adumbilical third of the flank indepen·
dently from the prim8l'Y ribs. With growth the
ribs flatten on the adventral third of the flank
and tend to develop longitudinal grooves.
B) Up to 6 0mm [iC·223x-18]. After the
stage of accentu ated flattening and grooving of
the ribs they abruptly loose their broad shape
on the external half of the flank and become
rounded, they also loose their periven tral clavi
and have a unifonn thickness, they then extend
onto venter in a wide adoral arc.
DIAGNOSIS. This species is characteri·
zed by: A) the early ribs with a propellor-blade
shape, and B) rounded adult ribs.
The earliest ribs observed (La-Ye-l ;H:4
mm) are wide, strap-like; the primary ribs begin
at the umbilical seam and trend straight to the
umbilical rim, from where they are inclined
adorally across the adumbilical fourlh of the
flank, near this point a secondary rib detachs
· from the lldoral side of a primary rib; about
center of the flank the ribs describe a sharp
adoral arc, on the rest of the flank they descri­
be an adoral concavity and finally at the peri·
ventral margin they terminate and develop a
slight spherical bulge.
The width of the ribs is unifonn or
increases very little, and they are twice as wide
as the intercostal spaces. At a whorl height of 8
mm (La·Ye) the wide ribs cross the venter. The
ribs are more prominent on the adumbilical
fourth of the flank where they have acute to
broadly rounded tops, they have their maxi·
mum flattening on the adventral fourth of the
flank (particularly noticeable on the internal
mold) . The periventral tubercles progressively
become more clavate; the adapical side of .�e
ribs is steeper than the adoral side.
·

Meuurementa. D Euw H w D/Euw D/H N o .R.

% %
C • 223x · 1 6 42 14 1 9 13 33 46 ? B) Up to 50 mm (La· Ye-2). The paddle·
C • 223x • 1 7 38 13 16 12 34 40 31 like shape of the ribs become accentuated and
C • 223x · 1 8 68 23 22 1 7 39 38 1 4 (1 ,12) they tend to split longitudinall y ; the intercostal
spaces become very wide especially on venter.
COMMENTS. This species differs from At a D. of 54 mm (La-Ye-3) the ribs quickly
Dutrenoyia hansbuergli n . sp., mostly in the loose their paddle-like shape and become uni·
middle and late stages of growth. Casey (1964, fonnly thin, strongly anteriorly arcuate on ven·
p.386) compared his Dutrenoyia notha to ter and the periventral clavi diminish in strength.
D[ utrenoyia] texana sane to rum Biirgl after a
"topotype" (?) referred to Biirgl's figs. 3a·b, NOTE : This morph has coarser and flatter rib-
plate 2; Casey concluded that the latter "close­ bing tl1an the holotype.
ly resembles the present species at small diame·
ters, but its venter is ribbed at &n earlier stage
and the umbilical wall is higher". My query as l ements
Me8lllr D Euw H w DjEuw D/H No.R
to the topotype locality refers to the exact stra· % %
tigraphic horizon and not to the geographic C·223x·1 31 9 14 11 29 46 36
IU'ea. C·223x•6 36 11 16 12 30 44 30?
C·223x·14 27 f 12 9 26 44 36
La·Ye·1 22 7 10 8 31 4!1 38
Dufrenoyia sanctornm Biirgl La·Ye-2 39 11 18 13 27 46 33·.
Pl. 1, fig. 3; Text-fig.14. La·Ye·3 61 20 28 20 33 47 34

. (Morph I) COM.i\1ENTS, This morph resembles to

Du{reno� durangensis Humphrey (1949, pl27,
pl.9, figs. 3-4), but:at comparable diameters the
1956 Dufrenoya texana var. media Biirgl, p.3, Mexi�an _spe�ies h� strong, outwardly directed.
p �13 , pl.2 , fig. 1, pl.3; fig. 1. : "clavt", 1 ts nbs are;. more flexuous, rmer and se·
parated by wider intercostal spaces. Dufrenoyia
1964 Putrenoya texana �edfa Biirgl, Etayo� . discoidalis Casey ( 1964, p; 393, pl.66, figs. HI;
Serna, p.121. text-fig. 140 g) resembles this mol'Jil., �pecially
in its early ontogeny; the English species differs
Reference specimen. The specimen Bff l/4 by the latter development and attenuation of
illustrated_ by Biirgl, 1956, pl.2, fi((. ) : its ornament, lack of an anterior arc of the ribs
Reference locality. Mesa de Los Santos : c:�-
, . . ., 1· ·• •
on venter and no tendency for its ribs to be wi·
tander). dely separated in the adults atage.
..
 zoNATION O F THE CRETACE OUS OF CENTRAL C O LOMBIA BY AMMONITES 61

�utmno yia hansbuer«Jj�

fig. 4; Text-fig. 14.
COMMENTS. Hill (June 1893) described
. 1, an ammonite species from Texas that he called
Acanthor.era�> iustinae; in November of the sa­
1956 ya texana1 texana
Dufrenop.3,6,12 Burckhardt, me year Cragin (p. 234, pl. 44, figs. 4-5 ) descri·
Burgi, ; [1. , fig. 1; pl.2, fig . 2, 4 ; bed another species from approximately the sa­
. pl. 3, fig. 2, 4 . me region as Hill's SJlecies and called it Hoplites
..,_.
roemeri. Again in 19'01 Hill (pl. 2 1 , fig. 6) illus­
1964 Du{ren oya texana texana Burckhardt , trated his species and gave its stratigraphic posi·
Etayo-Sema, p. 121. tion [p. 141 = "Section No.6,Cow Creek b eds .5] .
A few years later Lassw itz ( 1904, p.4, text-fig.
1965 texana Burckhardt,
Du{renoyia texanaCampbell. 1) illustrated a specimen from Texas that was
. ·Biirgl; in Btirgl and determined as Hoplites furcatus by Kilian. In
1925 Burckhardt (p. 20) did not accept Kilian ' s
. type. The specimen
i:IC}lo Bffl/1 figured by interpretacion and proposed a new specific na·
Biirgl, 1956, pl.l, fig. 1 . me Du{renoya texana for Lassw itz's specimen;
Paratypes. The specimens figured by Btirgl 1956, Burckhardt chose as holotype the specimen
. . .. pl. �l fig. 2 4 . studied by Kilian giving photograph of a plaster
.�
Hypotype. (U,.N.C.M.rt. .N.C.P.Hy.N . 2 3 , 24, 25), replica of the original. Scott (1940, p. 971)
Type . locality:- Mesa de los Santos (Santander). considered all the above nominal species as
synonyms of the first, that is Acanthoceras
·>A) Up to 30mm
D [C-223x-4 ] . The early justinae Hill.
whorl section studied (Hw : 4mm) is subellipti­
cal the. si des are slightly convex on the adum­ For the Mexican specimens referred by
bi1ical third of the flank, the rema1nder is incli­ Burckhardt to "Du{renoya texana" which Scott
ned toward venter and rounds into a moderate­ considered , very different from the Texan
ly wide venter. The u m bilical wall is subvertical, species, Scott proposed the name Du{renoya
it rounds smoothly into the sides. The primary Burckhardti chasing as type the specimen
ribs begin at the umbilical seam, they are prorsi­ fi gured on plate 9, figs. 7-9, of Burckhardt's
ritdlate on the adumbilical third of the flank, paper. Scott (op.cit., p. 1024) l ikewise sugges­
describe a sh allo w adoral concavity on the cen· ted that H ill 's type specimen h ad not ne ecessa­
tial and adventral thirds of the flank; on the in­ rily been collected in situ. Scott (ibid, p. 1023)
ner. h alf of the flank the ribs are narrow, acu te referred to Hill's species as "species thinly
but on the external half they are flattened. Ex­ discoid with nearly flat flanks and fl attened
ternally the intercostal spaces are as wide as the venter that becomes sligh tly rounded on older
ribs, and wider than the ribs on the in ternal mold. wh orls . . . . " .
The ribs end in small rounded tubercles which
� connected across venter in a slightly arched The next reviewer of Hill's species was
bar. Humphrey ( 1 949), who wrote: "Hundreds of

B) Up to 50mm D ( La-Ye-12]. The flexu-

examples of Dufrenoya from various northern
.
·. . Mexico localities, identical with the specimens
ous earlY ribs tend to strrughten ana at the same from the Rio Nazas Valley in Durango figured
time the adventral third of the ribs changes from
Oa:tten e d to round ed; the ribs are feebly arcu ate
by Burckhardt as Dufrenoya texana B urck har
dt
have been studied by the writer . . . . The writer
· on venter and narrower than their intercostal believes there is n o di fference be tween exam·
spaces: ·
pies from the two areas that do not come within
the very considerable range of variation of the
.; C) Up to 1 25 mm D (C-223x-2 3 ] . The species. Dufrenoya burckhardti Scott is here
whqrl section increases in he1ght and th e ri bs considered conspecific with Dufrenoya justinae
accentu ate their anterior arc on venter, on the (I lill ), as are the Colombian forms described and
·nank the ribs resemble flexuous clubs. Du{renoya texana
identified by Riedel as Burck·
hardt:' Humphrey (ibid, p. 122) added : ''Fonn
.-·.· - -

..· ·, , DIAGNOSIS. This species is ch aracterized discoid al ; compressed . ." and l ater he added (p.
�y:. A} a p rolo nged stage during which the ribs
.

123) "This species is characterized especially by

:·4?ross th e yenter approximately normal to th e
v,en tr� median line anQ. only late in the ontogeny
�() they forl11 an ad oral arc, B) the ribs show a
�leaqendency to be sep arated in to prim ary and
U:ttercalary ribs after the early juvenile
,�} stage, and
the change throu gh ontogeny from adventra
.· <Y fl attened to rounded , club-sh
.
aped nbs.
.; MeaSi..tre.tnent. D Euw
··-·. . . . , .
the rel ie f and prominence of the rib bing across
the squared venter".
After having studied several of Hum­
·
phrey's specimens, the Mexican specimens, in
spite of post-burial deformation, appear more
compressed than the Colombian forms; some of
them, however, h ave a pyriform section (v. gr.
H w ·DfEuw D/H No.R. UM 24248; Humphrey's pl . 6, fig. 1, p. 122):
% % the sides are rounded, bulging on the adumbili­

¢ · 21!3�. · ] .·· 31 10 13 1 1 32 42 ? cal half of the flank and slightly contracted on

··
��·�;!: :�� . 36
35
12
12
16 ?.
1 o 12
33
34
46
43
39
1
the adventral side. The Mexican specimens
likewise have flaring ribs on the adumbilical h alf
,c::� 223x ·22 90 ?
' Y&: · Ye:,
:J o 4 2 30 33 46 of the flank and the ribs widen adventrally (v.
.Il 47 13 20 1 6 28 42 37 gr. UM 2424 6 ; Humphrey's pl. 7, fig. 1,2} and
t;a . � Ye . 12 50 18 22 1 6 36 44 33 cross the vente r normal to the median line, the
�

• H\ Ye . 1 a 62 24
.c-: ·:. · · 25 1 9 38 ·U 37 ven ter, however, appears narrow for the size of
62 F. E TA Y O · SERN A
the specimen. In the Mexican specimens studied
by Humphrey an d available to me, there is no
more than a sligh tly rounded widening of the
ribs approaching the periventral tubercles (UM
24248; UM 21 854, Humph rey 's pl. 7, fig. 5).
.Furthermore the ribbing is rather uniform
throughout the ontogeny, withou t bifu rcating
rib s, as Humphrey (op. cit., p. 122) stressed
"Any suggestion of bifurc ation is usually more
apparent than real, . . . ". On the basis of the
above features, I do not find any grounds
to use the specific names " texana" or ''jus tinae "
for the Colombian species.
Apparently unaware of Humphrey's paper
Burgi ( 1956) studied the so-called ''Du{renoya
texana Burckhardt" from Colombia. recognizing
three varieties. CantU-Chapa ( 1963, p. 57) also
referred the Colombian Du{renoy ia ssp. to the
Texan species. Casey (1964 , p. 381) seems to
accept Burckh ard t species without any com­
ments on Scott's and Humphrey 's treatment
of Bu rckhard t' s species. Recently You ng (1972),
has shown that Hill's species is restricted to Te·
xas, (ib. p. 4 ) : "the endemic species Du{renoyia
jusl,inae (Hql. 189 3) I= D. texana (Burckhardt)
= ll. rqemen (Cragin)]".
Du{renoyia ha ns b ue rgli differs from D. san e
toru m BUrgi because of the ad ven tral flattening
of the ribs and the adoral arc of the ribs o n the
venter from early in the ontogeny, however, they
are most early differentiated in middle and late
stages of growth.
·
Genus (jveodeshayesite�lCasey ,1964
(non Prodeshayesites Casey , Dimitrova,
1967' p . 178).
}{e..odesba.yesites. Casey, .1964, p. 289.
TYPE SPECIES. Deshayesites s tu tzeri Riedel,
Original designa ti on .
Neod�shpJ,.esites columbianus , .( !t!edel).
·· -�· pJ� -g·; fig ;
8; pt.· ·10; fig. 7; Text-figs. 6P,
6R, 6T; 15.
1938 Deshayesites colu mbianus Riedel,
1
p . 40 pl.7 h figs. 16-20, pl. 12, fg.12,
pl . l4, Iig. ;�;1.
1954 Deshayesites columbianus Riedel,
·
ImlaYjl) Rod and Mayne, p.266.
non 1954 Deshayesites colombianus Riedel,
Biirgl, p. 16.
1955 Colo m biceras kars teni (Marcou),
BUrgi, 1955, p.15, pl. 4 , fgs. 1 ,2.
1957 Deshayasites colu mbianus Riedel,
Btirgl, p. 1 34, pars.
1957 Colombiceras k-arsteni (marcou),
BUrgi, p. 136, pl. 9, fig.l.
1 964 [Neodeshayesites I colombian us I Rie·
del J, Casey, p . 29 4 .
1964 Co /ombiceras karsteni (M arcou), Eta·
yo-Se m a, p . l 19, pars .
1964 Hypacanthop/ites colombianus ( Rie·
del}, E tayo.Serna, p . 1 20 pars.
? 1965 Hypacan thoplites karsteni (Marcou ) ,
BUrgi in Campbell an d Bi.irgl.
? 1965 Sonneratia colombiana Scott, nurgl ,
,
in Campbell and Biirgl 1965.
non 1968 Neodeshayesites cf. colombianus
[ Riedel }, Etayo·Sema, table 1 .
non 1 9 7 2 Neodeshayesites cf. colombianus
( R iede l ) , E tayo-Serna, p. 4 3.
Lectotype. Here designated, the specimen figu­
red by Riedel, 1938, pl. 7 , fig. 16-17,
pl . 12 , fig. 12.
Hypotype. (U.N.C.M.H.N.C.P.Hy.N.26, 27, 28).
Type locality. Here designated. Apulo (Rafael
Reyes) anticline (see under com·
· ments).
Up to 15 mm D (C-12C·19). The first
whorl of the phragmocone Is depressed ; the
second whorl is sli gh tly higher th an wide, it has
flattened sides that round rather abruptly into a
.steep umbilical wall but merge evenly into a
broadly arched venter. On the adapical half of
the third wh orl there are flat sinuous folds, se�
,
p arated by very narrow intercostal sp aces begi·
ning at the umbilical seam and becoming grea.
tly widened on venter, and strap-like and appea·
ring slightly arcuate; they are marked by "bun·
dies" of growth lines. On the adoral half of the
third whorl the "bundles" slightly elevate on
the umbilical . wall and ad um bil ical third of the
flank and a true ribbing becomes e sta bli s hed.
Up to 50 m m D (C - 1 2-c ) . The whorl sec·
tion increases in height and the flanks sligh tly
converge to the arched venter; the whorl rema­
ins thicker at the periumbilical m argin . The
primary ribs begin at the umbilical seam: they
are concave on the umbilical wall, prorsiradiate
on the adumbilical third of the flank and con·
cave again on the central and adven tral thirds,
crossing the venter straight and flattened like
bars. At the periumbilical region the ribs are rai·
sed in simple bullae (corresponding to pyriform
swellings on the internal mold), and have sligh·
tly flattened crests. The secondary ribs origin&·
te near the adumbili cal fourth of the flan k . In
general the ribs are high and prominent with
their adoral side a little higher than the adapical
one. The intercostal spaces are twice as wide as
the. ribs. The shell is thick.
C) Up to 75 mm D (C-12·C). The ribs ha­
ve symmetrically slopit:tg sides and attain a uni·
form width and height ·except on venter where
they are slightly wider; on the living chamber
and approaching it the ribs tend to lose the
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES 63

adoral concavity on the adven?"al �ird area.

of the Meuurements D Euw H D/E\lW D/H N .R. T.r.
0"
W

appearin g nearly rectiradtate m this %

flank Ill
'l'he �condary ribs tend to .be .less frequent. C·12oC·13 19 19 1 6 37 37 43 E
at ­
The maximum flarin g of the nbs ts the boun
C·l2.C·14 49 19 19?15 39 39 41 D
ary betwe en the centrallmd adumbilical thirds C·12.C·15 50 19 20 1 5 38 40 44 c
d C·l2oC·16 48 18 1 8 13 37 37 45 A
of the flank. C·12.C·17 49 18 19 1 3 37 49 47 A
OJAGNOSIS. Riedel's statement (l93�, C·12oC·18 36 12 1 4 11 34 40 40 F

p. 40) seems to me accurate: "La nueva especte

C·12.C·19 74 31 28 ? 42 38 26* A

esta caracterizada por sus �pstill� pron�I}Ct�das

c-n-c-21 68 27 27 19 39 39 44 A

y dobladas, con intercalacton y btfurca

cton trre· C·12.C·22 57 20 22 17 35 38 41 D
C·l2.C·23 48 19 20 16 39 41 44 c
guta!es". C·12oC·24 65 23 21 1 6 42 38 47 B

.. POPULATIO N ANALYSIS.
Sample C-12· C-12oC·26 46 18 18 1 2 40 40 41 B
from a sinl!le 46
to • C·l2 • C-34, obtained
20 40 F
c-1 C·l2oC·26 17 1 4 43 37

"concretion" was selected as reference popula­

..
C·12.C·27 46 20 20 13 43 43 43 D
tion� From. I!, two .spec�me �s were identi[te� as C·12.C·28 62 23 23 18 37 37 26• F
morpholog�ciilly "tdentlcal f:=closely stmilar] C·1 2.C·29 27 9 12 8 33 44 40 B
+
to the specimens figured by R:tedel, thus I con· C·12oC-30 42 14 18 1 4 ? ? 22*

sider th.em conspecific. C·l2.C·33 37 13 16 ? 35 43 ? A?

C·l2oC·34 39 13 16 6 33 41 40 F

Specimen C-12-C-18 ==Riedel's pl.7, figs. *

16�17. NOTE : ; half a whorl. T.r. = type of ribbing.
Specimen C-12-C-25 =Riedel's pl.7, fig.
18-20. COMMENTS. The similarities of this spe·
cles to the others described by Riedel were t'u·
Using specimens of this sample a series of Uy discussed by him (1938). The specimens
correlation diagrams between mensurable cha· from Apulo attributed by Btirgl (1955, pl. 4,
racteristics were made and the position of the fig. 1, 2) to " Colombiceras karsteni (Marcou)",
two specimens considered as "identical" with which I have been able to study correspond to
Riedel's .figured specimens were marked on the Riedel's species. The assignment of Biirgl's spe­
diagrams. The types (see Text-fig.3B) of ribs are cimens (HB 570/35 ; HB 570/2) to Neodesha­
rePresented by letters. 'The different combina­ yesites has been previously indicated by Casey
ti.ons of ribs .distribution are as follows: (1964, p. 295, footnote).
· Type A : PqP, P2pP. O CCURRENCE. Capotes Member. Lower
Albian.
·

It: �qP, P2pP, PP

·

c :
•

, P2pP, pp.
.

D : PqP, PP. Neodesh¥;e�ites a��lRqre� n, SR�

E : Pf."8", fig .; p1.9, lg�; Text* SV,
_

P2pP. • gs.6S ,
· F : PqP.. 6W, 6Z; 16.

c: · ······· ·· ;. .· • Fr.om
. .
.

th
.e
diagrams (Text-fig.15) it is Holctype. C-219 (U.N.C.M.H.N.C.P.Ht.N. 49).
.. .
:ev:tdent type F is more frequent on
that the Paratypes. (U.N.C.M.H.N .C.P.Pt.N.49, 50, 51).
.
. ·

sp�lmens of small diameter and types B and A

OJ1·.�peciJnens of larger size. Type C seems to be A) Up to 30mm D (C-219-13,14,15 ]. The
deyelC)ped . on the living chamber of larger early whorls have flattened sides and gentle
��lti:lens; The lectotype as figured by Riedel coavex venter. The external umbilical width is
;�eJ!ia to be PPqPP, but it could also be indica· approximately 35o/o of the diameter, it has a
�<1/!11 P�pP pp! in other words1 a . type c. or high inclined wall that rounds evenly into the
·(�-�W
;!ib �gary on. one side but secondary on nbs
With pnrnary and sem1pnmary
..

sides. Before a diameter of 15 mm the flank has

the riblike thickenings on the adumbilical third, the
Qppoln,te); thiS suggests that the largest of following one-fourth of the whorl has only fi.
B..le�.�ls spe ens probably represents the ne growth lines that describe a wide concavi­
·l�.r.em.almoldciJn
ofthe living chamber. ty from the umbilical seam to the periumbili·
cal third of the flank, on the Internal mold
f.l��m�nta D E\lW H w D/Euw D{H N.R.T.r. these are represented by flattened bands, very
o/o % wide on venter.

II
66 26 24 18 38 37 47 A
62 . 23 26 21 37 42 47 B At a diameter of about 15 mm ribs appear
61 25 23 17 41 38 47 c again on the exterior and are also well dermed
70 30 26 17 43 36 47 D on the internal mold : they become prominent
76 32 28 21 4Z 37 45 c at the periumbilical margin, extend with a slight
77 33 27 19 43 35 47 adoral concavity on the adumbilical third of
•. !?tii.4-'7
B
80 33 31 22? 41 flank; on the central third the ribs are adorally

-�···
39 50 D
67 27. 23 17 40 34 47 B convex and then are adoral concave on the
62 26 22 17 42 35 47 B adventral third. On the ventral margins the ribs
63 �1 22 16 40 41 48 A thicken and cross the venter normal to the
57 .22 median line [ C-219·15] ; at this point the
.
24
. .
17 38
;:q;
·· r;t*tz
42 44 c
.

. .
66 21 19 intercostal spaces are shallow and the ribs appear
. . · - .
.
14 38 34 41 c
64 F. E TAY O - SERNA

asflat bands slightly higher on their adoral bor­ Measurements. D Euw H W DfEuw D(H N.R. T.r.
der. OJ. %
C-219·14 26 9 11 9 36 42 24? E?
Near a diameter of 30 mm the ribs are C·219·15 27 11 11 9 40 40 407 A?
well defined, they widen towards venter, the
intercostal spaces are clearly defined on both
sides and venter; the adapical slope of the ribs COMMENTS. Neodeshayesites albertoalva·
becomes shorter but steeper than the adoral rezi n. sp. differs from N. columbianu s (Riedel)
one and in general the ribs have very nanow because during its ontogenetic development,A)
flat top. The ornament Is best described as the intercalary ribs develop independently from
consisting of alternating primary and interca­ the primary ribs earlier than in Riedel species,
lary ribs but several semiprimary or a sequence B) the beveling of the umbilical wall occurs
of three primary ribs may also occur; the earlier and more strongly in N. albertoalvarezi,
secondary ribs detach from the primary ribs at C) the ventral area is relatively narrower than
the umbilical rim but very soon this junction on Riedel's species, D) when the test is preser­
moves outwards to the boundary between the ved the conch appears more inflated and in
central and adumbilical thirds of the flank, but general the adults are smaller than in Riedel's
on the internal mold they seem to begin near species, and E) the prominence of the ribbing
the middle of the flank. The ribs are flared round the umbilicus decreases earlier in my
midway on the flank and on the internal mold species.
small pyriform nodes appear on the umbilical
periphery. From my study of these two _species it is
pertinent to add to the diagnosis of the genus
B) Up to 50 mm D I C-219·6,101. The rib· (see Casey, 1965, footnote p. 289) the follo­
bing is stabilized as an aftemation ot primary wing: the early ontogeny prior to the stages
and intercalary ribs, they tend to h ave a uniform illustrated by Riedel (1938, pl. 7) shows first
width on the flank but they are slightly wider folds that widen toward the venter, these folds
and flatter on venter though maintaining a simi­ split longitudinally during following stage, and
lar elevation. At this stage the primary ribs de­ finally the ribs individualize and their fold like
crease their elevation from the middle of the aspect on venter disappears. The ribs are never
flank toward the venter whereas the secondary interrupted on venter.
ribs become higher. The whorl section is more
inflated toward the adumbilical half of the
flank and the umbilical slope becomes gentler. OCCURRENCE. Capotes Member. Lower
Albian.
C) Up to 70 mm D [C-219·1,5]. The um­
bilicus has widened and its wall is moderately This species is named in honor of geologist
inclined; the flanks are moderately inflated on Alberto Alvarez 0.
the adumbilical third. The ribs are rounded and
appear slightly arcuate on ven ter because of the �
Neodesha esites k fs te n i (M arcou\
slight thickening periventrally and ventrally . Pl. 9, fig. ; pi.io, fig: ; t ext-figs. 6t, 6M,
6N, 17.
DIAGNOSIS. This species is characterized
by: A) the on togenetic development of its oma- . Le ctotype.Th be designated,Etayo-Sema in prep.
ment: early whorls with broad strap-like swe: Hypotypes. (U.N.C.M.H.N.C.P�Hy .N.29, 30,. 31}
llings followed by an almost smooth tract after
which the ribbing begins with a division of a 1858 Ammonites A costae, Karsten, p. 111,
primary rib, the junction soon disappearing pl. 5' fig. 1.
and the intercalary ribs appearing indeJ:enden­
tly near the middle of the flank, B) the pro­ 1875 A mmonites Karsteni, Marcou, p. 165.
gressively gentler gradient of the umbilical wall
through the ontogeny, C) the relatively inflated 1881 A [ mmon ites J acostae, Karst., Gabb,
whorl section and nanow venter, and D) its p. 310.
relatively small adult size.
1886 A m monites Kars ten i Marcou , Karste n,
Measurements. D Euw H W D/Euw D(H N.R, T.r. p. 62, pl. 5, fig. 1.
% %
C·219· 1 64 30 28 1 9 7 7 187 F? non 1928 Colombiceras Karsteni Marcou, Basse,
C·219· 2 50 . 22 20 14 44 40 43 F p . 1 36, fig. 14, pl. 8, fig. 5.
C-219· 3 62 20 20 12 38 38 43 A
C•219· 4 49 21 19 1 5 43 38 37 F ? 1931 Colombiceras Karsteni Marcou, Rhein
C-219· 6 49 20 19 1 5 40 38 60 A in Hubach, p. 58.
C·21 9· 6 47 20 17 1 3 42 .36 47 F
C•219· 7 43 19 15 1 2 44 35 44 F 1933 Colombiceras Karsteni Marcou, Spath,
C·219· 8 387 18 16 1 1 477 397 38 D p.654 ex Karsten 1886,non examples
C-219- 9 34 13 1 3 10 38 38 37 A in the British Museum under Nos. C
C·219·10 39 15 1f> 12 38 38 40 F 4284a, b).
C-219·11 36 14 1 4 12 40 40 35 F
C-219·12 •38 13 1 5? 1 2? 7 ? 1 F? non 1936 Colombiceras Karsteni Marcou, Basse ,
C•219·13 23 8 8 7 36 36 26 :'/ E? p. 88 .
 ZON ATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES 65

Up to 60 mm D [C-501·17,18). The peri·

? . Du{renoya (?) nov. sp. Breistroffer,
1936
p. 156. ventral rounded inflations of the ribs are accen­
tuated on the exterior but very little on the in·
? 1937 Colombiceras Karsteni Buell, Botero, temal mold. The secondary ribs loose their bi·
p. 26, figs. 48 , 49 . furcating nature and change to a more constant
alternating primary-intercalary Pq . The ribs
19 38 Colombicerus Karsteni Marcou, Riedel are moderately flattened on the adventral third
p. 35 . of the flank but continue very prominent near
the umbilical region; the umbilical wall becomes
1938 Ammonites Karsteni Marcou, Riedel, steeper.
p. 42.
D) Up to 80 mm D [C-501·1]. A clear al·
1947 Ammonites Acostae Karst., p. 378, temation of primary and intercalary ribs Pq
. pl.5, figs. on the left upper c:.omer; dominates; the umbilicus is1wid8t;but the whorl
Spanish vemion of Karsten's original section has a proportionally · lower height; the
paper. ribs are prorsocline or weakly concave adorally
on the adumbilical third of the flank, they are
non 1950 Colombiceras karsteni (Mareou� Basse, rectiradiate or feebly convex adorally on the
p . 252. outer two thirds. The ribs are uniformly thick
on the side and venter; the periventral lnfla·
non 1963 C. [olpmbiceras] Karsteni (Marcou), tions have almost disappeared. The ribs are pro­
Cantu Chapa, p.55. minent on the umbilical wall as lf they arose
from the umbilical seam.
·- 1964 Colombicerus karsteni (Marcou), Eta·
yo-Sema, p. 119 , pro part. DIAGNOSIS. This species is characterized
by: A) the ontogenetic trend towards a wider
1965 C.[olombiceras] Karsteni (Marcou), umbilical diameter versus a relatively short whorl
· Casey, p.419, pro Karsten's original height, B) the persistent angular venter that be·
material only, comes rounded late in the ontogeny, C) the
coarseness of its ribbing, and D) the relatively
shallow intercostal spaces.
: · · . tip .to 10 mm D [C-5 01·28). At a whorl
bei!Ptt. of 3 mm, the internal mold is higher Meuurementa D Euw H W DjEuw Dftl N.R. T.t.
than wide and it is divided into segments by % %
Wfde constrictions th at are slightly arcuate on c-501· 1 75 36 25 20 46 32 43 F
, ::�nter; from that point on, flat bands are visible c-501· 2 49 24 20 16 47 39 40 F
·< ()n th:e �ank, they appear as periventral swellings, C-501· 3 60 25 23 17 43 39 38 F'
�; C;_op�mporane()usly on the shell the ribs begin 641 32 23 18 49 37 42 A
t(). .
C·501· 4
at the umbilical rim and are most
· ·
A
·
·
C-501· 5 159 27 20 17 44 36 38
tile�. c-501· 6 53 23 20 liS 42 37 42 F
C·l501· 7 53 26 20 12 415 36 38 F
C-501- 8 47 21 18 13 43 38 39 D
The
mm D [C-501-22,27,28). C·501· 9 38 17 15 12 43 38 38 F
than wide, oval with section c-501·1 0 39 16 16 11 42 37 40 A
-� ·�Iat 18 trutnc.ated on the maximum tltic· C·501·12 42 18 15 12 40 34 40 F'
third of the flank; C·501·13 17 11 10 8 51 ISO + 301 F'1
depression (contraction) of the C·501•14 59 25 22 18 41 36 41
venter. particularly apparent C-501·1 6 51 23 21 115 44 38 37 A
is preserved but not visible on C'·IS01·16 54 24 20 13 44 36 37 F
mold; the venter is flattened and C-501·1 7 62 23 20 14 42 37 34 E
angular margins. The external urn­ C-601·18 55 25 21 14 415 37 34 A
is less thin half the diameter, the C·501·19 43 18 17 11 43 38 37 F
is moderately inclined and rounds C·501·20 45 19 18 14 43 41 32 F7
C-601·21 41 19 16 13 46 38 32 D
C·501-22 41 17 16 11 40 36 31 A
C·501·23 39 17 14 11 42 35 31 E
C-501·24 41 19 17 7 46 40 317 F
C·501·25 37 15 14 9 40 37 31 E
C-1501·26 32 liS 12 9 42 33 32 D
C-601·27 31 14 12 8 45 38 33 F'
C-501·28 29 13 11 8 45 38 32 F
C·501·29 29 12 12 9 41 41 34 F
C-601-30 19 8 7 7 42 36 327 F
C-501·31 151 21 21 17 40 40 35 A
C-501·32 49 22 18 14 44 36 42 F
C·601·33 46 20 17 1 4 42 36 41 A
C-601•34 44 19 18 11 43 36 39 D
c-501·35 37 15 14 11 40 37 42 F
C·501·38 36 15 14 9? 40 39 36 F
C-501-37 34 13 1 4 10 38 41 39 F
 66 F . ETAYO · SERNA
Meuu:rement.B D Euw H W DfEuw D/H N.R; T.r.
% 04
C-601·38 23 9 9 7 39 39 39
C-1101·39 311 14 14 1 0 40 40 37
C·II01-'0 40 16 l li ll 38 36 43
C•601-'1 34 14 13 10 40 37 41
C·IIOl--'2 30 12 12 9 39 39 37
NOTE : F' signifies that there are two interca·
lary ribs between two primary ribs.
COMMENTS. "Ammonites Kars teni Mar·
cou" [1875, p. 1651 (ex Ammonites Acosroe,
Karsten, 1858, p.l li, pl.5, fig.1), was collected
acco"Ung to Karsten in "Tocaima", a locality
where no Cretaceous rocks crop out. Conse­
quently, if Karsten's sample does not represent
float, it must come from the only area where
Middle Cretaceous beds are present, that is
between Apulo (Rafael Reyes) and Anapoima,
where sample C-219 was collected.
Karsten's original description is as follows:
"Testa discoidea, complanata, dono subrotun­
do late umbilicata, argute costata; costis dorsum
clngentibus alternantibus brevioribus aliis com­
pletis undulatis, aliis interjectis in latere evanes·
centibtis vel hie · illis sejunctis; anfractus exte­
riores, interiores ad medium tegentes; ap,ertura
ovata rotundata; septis l"teraliter trilobis '.
·
"Durchmesser . . . . . . . . . . .48 Millimeter
Hohe der letzten Windung 16.5 "
Dicke der letzten Windung 12 " subequal to the ribs on the adapical portion of
Weite des Nabels . . . . . . 14 " the last whorl but appear as a narrow furrow on
However, this description and measure­
ments do not correspond to the figure of the
specimen given by Karsten (ibid. pl. 5, fig. 1a,
1b ). The diameter of the conch corresponds
well on fig.1a, based on the intercostal distance
likewise the height of the last whorl is als�
correct. The width of the umbilicus if measured
parallel to the diameter, and touching opposite
points of the umbilical seam (fig.la.) is 18 mm
on the figure contrary to 14 mm as stated by
Karsten, consequently the conch was drawn
more evolute and rounded than it is in reality
(did the artist try to indicate an oblique umbili­
cal wall?). On Kmten's figure 1b, there is an
error in the perspective: the view was not drawn
looking straight at the specimen but is an obli­
que view, consequently the "aperture" looks
higher, and if my interpretation is correct the
artist tried to draw. the aperture following the
path of a primary rib, consequently the width
(thickness) of the aperture is greater in the
figure than on the original (=18 mm va.14 mm).
The young of Neodeshayesites karsteni
(Marcou) resemble the specimen fi�red by
lliedel (1937, pl. 7, figs.12-13) as a ' transitio­
nal spec1men" between "Deshayesites stutzeri..
and ·�neshayesites nodosus " the adults of the
latter species have very fine or flattened ribbing
and the whorl section appears higher than In N.
karsten�. �rom N. . colum bilzn!'s (Riedel) N:
karstem dtffers by 1ts hemer . nbbing, rounded
throughout on the sides. On Riedel's species the
ribs are high, acute and only adventrally roun·
ded, furthennore on his species . the umbilical
F wall is persistenly steeper.
F
F OCCURRENCE. Lower Albian.
F
F Neodeshayesites karsteni (Marcou)
Pl.lO, fig. 9; Text-fig.17.
(Morph I)
Hypotype. C-501-2 (U. N. C. M. H. N. C. P. Hy.
N.32).
This morph is characterized by its roun­
ded venter very early in the ontogeny; its rib­
bing is attenuated, not flanging at umbilical
rim; on the flanks of the internal mold the ribs
appear as concentric arcs.
OCCURRENCE. Lower Albian.
�deshaye6.ituneylatum,.n. sp.
Pl.lO, ffg. 8, 10; Text-figs. 7 A, 7G, 7H.
Holotype. C-501·0 (U.N.C.M.H.N .C.P .Ht. N .50).
The whorl section is higher than wide,
ogival, with broad venter and flattened flanks;
the umbilical wall is beveled and rounds
smoothly into the sides. The ribs are wide and
low, on both test and internal mold, they are
separated by shallow intercostal spaces that are
the adoral half; the ribs arise in pairs, rarely in
threes, from stressed periumbilical bullae, they
are slightly sinuous and have delicate rounded
periventral thickenings. At a diameter of 55
mm the periumbilical bullae disappear and the
ribs are straplike. Although the ribs are wider
on venter they increase their width slowly.
DIAGNOSIS. This species is characterized
by the change in its ornament from ribs that
arise from accentuated periumbilical bullae
to non bullate girdle like ribs, and by its flatte·
ned ogival section.
Meuu:rementa D Euw H W D(Euw•fo D/H%
C•II01o0 66 28 27 19 43 41
COMMENTS. This species resembles Neo­
deshayesites ro tundus (Riedel) (1938, p. 39,
lam. 7, figs. 14,15; pl.14, fig.20) but the latter
has finer ribbing, more numerous and fmer peri­
umbilical bullae; and the venter is narrower.
Neodeshayesites.contracta (Riedel)_
PL9, fig. 7; Text-figs. 7B, 7D, 7E, 7F.
1938 Deshayesites s tu tzeri var. con tracta Riedel•
p.38, pl. 7' fig. 8, 9 ; pl.14, fig.18.
Hoiotype. Riedel's fii!Ufed specimen.
Hypotype. C-500 (U.N.CM.H.N.c.P.Ht. N.33,34,35).
A) Up to 20 mm D (C-500-1). The whorl
section is higher than wide, the tlanks are sligh·
tly inflated, with their adventral thitd conwrgmg
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES 67

t6ward the venter; the umbilical wall is vertical. Genus [§toyiggwicer� ?�,.::;
the conch observed (H: 4 mm)
Tlie inner part dofventer
haS a . truncate and .subangular ventr� TYPE SPECIES. Ammonites Treffryanus Kars­
m argins. The primary ribs anse from the umba­ ten, 1858, p.150, pl.4, figs.
lical rim where they usuill!Y have a bulla, .�eyal 1a, lb.
describe an adoral concavaty on the umbalic
wall an-d extend pri>rsiradiately to the boundux DIAGNOSIS. High whorled, involute
betw�en the atcentral and the adumbilical thirds conchs ornamented from early iri the ontogeny
of the flank; this point the ribs bifurcate; and with biconcave primary ribs and flexuous inter­
on · the adventral half each branch describes a calary ribs, the former may be raised around the
wide adoral concavity and fin�y cross ven�r umbilical margin but no tubercles are present.
with ·a weak adoralwith arc. At thiS stage the nbs When the shell is preserved all ribs have rounded
appear like sickles long handles. The ribs tops and sides, on the venter they tend to broa­
are broad; rounded to flat topped and have
a den in a visor-like manner but they cross normal
steep adapical side, they are slightly wider than to the median ventral line or with a weak adoral
th� intercostal spaces. On this specimen there arch. The whorl section is high, ogival, with wi·
iJlay be seve!al non bullate ribs or one .rib w�th de venter and inclined umbilical wall. The bor·
8: less ptormnent bulla between two nbs wtth der of aperture is plain with a shallow- sinus at
prominent bullae. the umbilical margin.
·
B)Up to 100 mm D (C-500-2). The peri­ Suture line with E/L large, L symmetrica­
umbilical bifurcations become regularized; lly trifid, L/U and U2 high.
s:ollle intercalary ribs may approach a primaryit
rib near the umbilical rim as bifurcating from COMMENTS. Stoyanowiceras differs from
but the real branching occurs centralat the boundary Colombiceras Spath (1931) by the coronate
between the adumbilical and thirds of early whorls and straight ribs, wedge shaped
the flank; at this point the ribs tend to develop with steep sides, of the latter genus.
a·t;a'iangular swelling. The venter is progressively
more rounded and the umbilical wall becomes Stoyanowiceras tref£B:anus (Karsten)
very inclined; the conch has periodical constric· PI.5, fig.3; pl.4, fi{7Text-Mgs.7M,7N,7Q
f.ions on the living chamber. The living chamber
is aproximately half the length of the whorl. 1858 Am. rmonitesl Treffryanus Karsten,
The ribs are rounded on the umbilical third of p. 101J, pl. 4, ffg. 1.
the flank, .but they are flat, terrace like on the
cimtraland adventral thirds. 1862 Ammonites Treffryanus Karst.,Gabb,
p. 75.
The. general trend of the ribs is to descri­
. tie a wide. adoral arc on most of the flank with 1875 Amm. [onitesl Tre{fryanus Karsten,
deUcate concavities both on the adumbilical Marcou, p. 164.
�d adventral fourths. Likewise the secondary
rib�. may join different primary ribs on each 1883 Hoplites Treffryanus Karst, Uhlig,
fl�k. p. 158.
< ·DIAGNOSIS. This species is characterized non 1883 Hoplites Treffryanus Karst, Uhlig, p.
�ro\}gh its ontogenetic development by: A) its 251, pl. 21, fig. 2 .
!ascic!Jlllte ribbing and increased crowding of
·the nbs around the umbilical margin, B) the 1886 Ammonites Treffryanus Karst., Kars·
liJPid Increase in whorl height, C) the narrow ten, p. 62, pl. 4, fig. 1.
Venter, and D) the numerous fine ribs.
· ·· ··
'! 1898 Hoplites Treffryanus Karst., Gerhardt.
'M'aiunmenq D Euw H w DJEuw D/H No.R. p. 158.
% %
i'���QO • l 20 1899 Parahoplites Treffryanus Karsten,
Anthula, p. 110. ·
6 9 7 33 46 40?
•. c"5oo : 2 4o 115? 17 ? 37 42 52
c�do � 3 63 22? 21 15 41 39 27(lf2)
9·�00 � . 67 25 26 17 37 38 61) non 1899 Parahoplites Treffryanus Karsten,
: l;h500 . 5
'.;,ci�oo �-6
71 29 27 17 40 38 70 Anthula, p. 115, pl. 8, fig. 6.
40
·.>_..:· :· :
+ 77 29 31 20 38 ?
"'
' c-
1907 Am. [monites l Tre{fryanus Karsten,
< COMMENTS.
.
��nvar. !Zont:czcta
·
Although "Deshayesites stut­ Sinzow, p. 485.
Riedel" was based on a juve­ 1907 P. [arahoplites] Treffryanus Karsten,
t�il�.I by Bu�.trg] (1957,
· ;�,.�red
thank · is a valid species. The specimen
fig. 3a, 3b) from Jacob, p. 77, non p. 81.
·.·f'l,UD !mow:n l.ocaljty haspl.a 8,more consistent bi·
., �ating nbbmg pattern. The present morph 1913 Par. [ahoplites] Treffryanus Karsten,
o::;�mbles
. rmple B.some specie s of Burckhardites, for
palu mbes Humphrey (1949, p.1 � 5,
Kilian, p. 344.
>l1. .12, figs. 1,2), but the latter has a greater an- 1921 Am. [monites] treffryanus Karsten,
'':-� .·_.· · · I!Se.dof the whorl heig�t. and the ribbing is not
_. · n-e Spath, footnote p. 317.
< . . sse around th� um
bthcus.
68 F. ETAYO · SERNA
? 1922 Parahoplites Treffryanus Karst., Co­
llet in Eugster, p. 254.
1931. Am. [monites J Treffryanus, Karsten,
Spath, p. 654. . primary ribs; the secondary ribs taper toward
l
? 1936 Deshayesites (?) Treffryanus Krst.,
Breistroffer, p. 155.
1937 A m. [monitesJ Treffryanus Karsten,
Scott, p. 37.
1938 Calombicerus treffryanus Karsten,
Riedel, p. 50.
? 1945 Parahoplites (Parahoplitoides) treffry�
·
(Karsten), Royo y Gomez.
1947 A m [monitesJ Tre{fryanU8 Karst.,
Karsten, SpaniSh version, p.376, pl.4,
top. figures.
1949 Ammonites Treffryanus Karsten,
Humphrey, p. 151.
1949 Ammonites Treffryanus Karsten,
Stoyanow, p. 121.
1952 C. [olombiceras] treffryanus Karsten,
Basse , p. 655.
1963 Colombiceras tre{fryanus (Karsten),
Cant:U·Chapa, p. 55.
1964 Colombiceras Treffryanus (Karsten),
·
Etayo-Sema, p. 120, pars.
fr
? 1964 Burckhardites Tref yanus (Karsten),
Biirgl, in Campbell and Biirgl.
Holotype. The specimen illustrated by Karsten,
1858, p. 109, pl. 4, fig. 1. Lost du­
ring 2nd world war (J. Helms, written
comm. 1976).
Neotype. Here designated Plate 4, fig.l.
Paratype. (U.N.C.M.H.N.C.P.Pt. N. 52, 53).
A) Up to 5 mm D. The whorl section is
transv�rsely subrectangular, with angles roun­
ded; -'�ong the umbilical rim there are broad
undulations that soon become ridges separated
by wide constrictions especially prominent on
the flanks; the ridges have a weak longitudinal
groove, on the shell these ridges are represented
by flexuous dfvided bands.
. B) Up to 21 mm D (Pens. 1, 2, 3, 4). At
a diameter of 14mm the whorl section(Pen.N.2)
is subrectangular, the flanks are flattened, the
venter and periventral margins are rounded, the
umbilical wall is subvertical near the umbilical
seam but externally is more inclined, and rounds
smoothly into"the side.
The ornament consists of rounded
flexuous primary ribs that begin on middle of
the umbilical wall, curve adorally in the lower
part of the whorl, curve adapically on the
middle part, curve slightly adapically on the
upper part and then cross venter straight.
On the periven tral third of the flank and
on venter the ribs are moderately broad, club
shaped; the secondary ribs, usually one but
occasionally two, intercalate between the
the periumbilical third of the flank where they
fade out; sporadically one of the secondary ribs
slants toward the following primary giving the
impression o f bifurcating from it, likewise an
actual bifurcation from the adoral side of a
primary rib may also occur. In general the
primary ribs are more prominent than the
secondary ribs. The intercostal spaces are twice
as wide as the ribs. When the shell is preserved
the tops of the ribs are broadly rounded and on
venter they attentuate.
C) .tJp to 45mm D (Pen 10). With growth
the ribs gain an elegant biconcave outline on the
whorl sides; the primary ribs become broadly
rounded almost flattened on the adventral half
of the flank. Adoral arching of the ribs on ven·
ter does not occur. Some of the intercalary ribs
fork from the adoral slope of the preceding pri­
mary rib on the adumbilical third of the whorl.
Sporadically some primary ribs are more promi·
nent. The whorl section increases its height but
not its width, consequently it is always compres·
sed. The involu tion line is near the boundary
between the adumbilical and central thirds of
the flank.
DIAGNOSIS. As for the genus.
MEASUREMENTS. The specimens stu·
died do not allow accurate measurements.
COMMENTS. The following is Karsten's
original description (1858, p. 109 ) : "A. testa
compressa umbilicata, dorso rotundata, trans­
versim costata; costis rotundatis, flexuosis,
inaequalibus, dorsum cingentibus; altemantibus
incompletis, una suturam unbilicalem attingen­
te alia solitaria vel rarius gemina in latere evanes·
cente, Iibera vel rarius costae anteriori (majori)
conjuncta; anfractibus subinvolutis, exteriore
tres partes interioris involvente; apertura com·
pressa ovato-elliptica: umbilicoparvo; septis la­
teraliter trilobis profounde partitis, margine
crenato-sectis.
Durchmesser 180 Millimeter
Hohe der letzten Windung 53 "
Dicke derselben 50 "
Durchmesser des Nabels 20 "
Karsten's species has been the object of
many comments in paleontological literature :
Uhligh (1883) referred this species to lloplites;
Anthula referred the species to his group of Pa­
rahoplites melchioris, but Sinzow (1907) cha·
llenged the interpretacion "weil wir die Gattung
gar nfcht kennen, zu welcher A m� Treffryanus
Karsten gerechnet wird".
Spath (1921, p.137) considered A. Tref­
fryanus Karsten in the same group as A. cras­
sicostatus d'Orbigny·oecause they " . . . show
the peculiar flattening of the costation . • •" ;
when proposing- his genus Colombiceras (1923,
 ZONATION OF THE CRETACEOUS
p 64 ) Spath did not mention Karsten's species
b�t Jater (1�31 , p.654) wh�n compt�ntif! g on an
'Indian specieS of Colomb1ceras be: considered it
very close to Karsten's species, Riedel (1938, p.
50) referred Karsten's species to Colombiceras.
Stoyanow (1949, ·p; 121) challenged Spath and
Riedel " tendency . . . to connect with
• . . thly into sides.
Colombiceras flat-ribbed species for which the
presence of tuberculate ribs has not been
proved, as Ammonites tre{{ryanus Karsten
( 1 856, p . 109, pl. 4, fig. 1a, 1b) ", and he . .
furthermore indicated that in following Spath
and Riedel's ideas there should be a group
for Karsten's species. Casey (1965, p. 4 20) has
followed Spath's treatment of Colombiceras,
however, his suggestion of a transient coronate
stage is not confirmed by disection; the deve·
Iopmerit of the ribbing in Stoyawiceras is like
that of Neodeshayesites.
Karsten wrote that his species came from
"Tocayma" (sic, p.109), but as previously com·
men ted no Cretaceous rocks crop put in the im·
mediate neighborhood of this locality, and be·
cause no other nearby region than the area co·
vered in the present report (Apulo . La Mesa),
has exposed Lower Cretaceous rocks, this is se·
lected as type locality for Karsten' species. The
specimens studied in this report closely resem­
ble "Ammonites Treffryanus Karsten", none of
of them, however, is as large as the holotype.
"Sonneratia colombiana Scott" the holo·
type (T.C. U. 1108) of which I have examined is
here considered a member of Stoyanowiceras,
contrary to Casey's views (1954, p.llO; 1965,
p. 400, p.516), and its specific status is retained
here because of its more distinct ribbing and re­
gular alternation of primary and secondary ribs,
the ribs are sharper on the inner half of the
flank . and narrower on the external half of the
whorl than in K1li'Sten's species as it is treated
in this study.
OCCURRENCE.Upper Aptian.
_ . :. .Stoyanowiceras tre{{ryanus (Karsten)
"PI: a; ng.s. · - · · · · ·
·
� ·
(Morph I)
Paratype. (U.N.C.M.H.N.C.P. Pt. N.54).
Th� morph is peculiar because its orna·
ment. is stressed very early in the ontogeny,
espe�1ally the flanging of. the ribs on the umbili·
cal nm.
·
Me&.u:rementa o Euw H W D/Euw % D/H "'o
Penldlv. 1 4 18 7 7 8 39
Pen.Uv: 1 3 30 10 1 2 12 33
OCCURRENCE. Upper Aptian.
Stoyanowiceras treffryanus (KaJSten)
P1.3, fig.12.
(Morph II)
Paratype . (U.N.C.M.H.N.C.P. Pt. N.55).
OF CENTRAL COLO M BIA BY AMMONITES 69
A single specimen is available. The whorl
.sectl(ln is higher than F and has a broadly
rounded venter, the sides ate slightly compres­
sed on the adventral third of the flank the cen·
tral and adumbilical thirds are slightly convex;
the umbilical wall is inclined and merges smoo·
The ornament consists of distinct ribs
which begin at middle of the umbilical wall; on
• the early portion of the last whorl the ribs
bifurcate at the margin of the umbilical wall,
they are fine, rounded and slightly biconcave a
bit projected forilard on venter; this type �f
ornamentation persists until the beginning of
the adoral third of the last whorl where the
ribs become strongly biconcave, and strongly
arcuate on venter; bifurcation of the ribs may
occur on one side but not on the opposite. The
intercostal spaces are one and a half times as
wide as the ribs.
NOTE : This morph is chuacterized by : A) the
rapid change in flexuosity of the ribs
that accen tuates with growth, and B)
the narrow well rounded ribs,
Measurements 0 Euw H W D{Euw% DtH% N.R.
PeDSUv. 7 36 11 1 6 1 3 31 43 37
Sto.;xt;WJ!J(J ctras cin tatuf1l.n. sp.
. ..• .,. ,
PJ.7; fig.9. ,._.,,. . , " -· . . .• . .. ,-�- ·-,.�4o,oo • ,-•--,.
Holotype. T.I.P. Guaduaus (U.N.C.M.H.N.C.P.
Ht. N.51).
Only one specimen is available. At the
beginning of the last preserved whorl the sec·
tion is higher than wide, the venter is rounded
and the whorl sides are slightly convex; the
umbilical wall is oblique; toward the adoral end
of the conch the height of the whorl increases
in relation to the width,
The ornament consists of ribs which
begin at the umbilical seam where they describe
an adoral concavity and extend up to a little
beyond the umbilical rim, then they describe an
adoral arc on the adumbilical half of the whorl;
up to this point the ribs are raised and promi­
nent, from here up to the periventral margin
the ribs are slightly concave adorally, they be·
come wide, broadly rounded to flattened and
go straight across the venter.
There is a regular alternation of primary
and intercalary ribs on both flanks, the interca­
lary ribs begin near the umbilical third of the
39 whorl where they slant adapically and approach
40 the preceding primary as if forking from it and
in some instances it seems a true bifurcation.
The intercostal spaces are narrower than the
ribs.
DIAGNOSIS. This species is characterized
by its low and broad heavy ribs.
Measurements D Euw H W D{Euw% DtH %
T.I.P.Guaduaa 79 24 36 26 30 46
70
COMMENTS. This species is closely simi•.
lar to Stoyanowiceras tre{fryanus (Karsten),
but has lower and broader ribs than the latter.
Stoyanowiceras colombiana (Scott) has sharper
and higher ribbing than S. cinta tum , n.sp.
Parahoplites nicholsoni Benavides-Caceres
also resembles S. cintatum n. sp. but the Peruvian
species has a wider umbilicus, its umbilical wall
is vertical and the ribs regularly bifurcate, but
the opposite is true in the Colombian species.
OCCURRENCE. Upper Aptian.
I'Ri:
Genus .f'fdelltes n .g.
. • ) r. f. ,, ;,.o,�r..
(£:r·a,, :<.£�,;"')
.
oJ!IIIil :t?SrcdiiSiii .,.!'
, "'
· · � · · ·· ·
fJvru.:.'' )_.(d):." 'i Reidelites obliq u u m (Riedel) because the latter
TYPE SPECIES. Riedelites esthersemae n. sp.
DIAGNOSIS. Evolute conchs of small to
moderate size; the whorl section is rounded
suboval to subquadrate. No tubercles are pre­
sent. The ribbing be�ins as delicately flexuos
ribs, crowded in the JUVenile but well spaced in
the adults. The ribs are acute adumbilically but
broadly rounded to feebly flattened on the ad­
ventral half of the whorl and on venter.
COMMENT:>. This genus differs from Co·
lombiceras in the lack of tuberculation and the
development of the ribbing throu�tt out the on­
togeny.
This genus is named in honor of Dr.
Leonhardt Riedel,
..Jlj�delites esthersemae n. sp.
Pl. 6, lig. 12; Text-ffgs. 1's, 7T.
Holotype. C·222A (U.N.C.M.H.N.C.P.Ht. N.52)
Patatype. (U. N. C. M. H. N. C. P. Pt .N.56)
A) Up to 38 mm D (T.I.P.G.). The inner
whorls are - slightly depressed (H: 3mm; W:
4mm); the ornament consists of raised rounded
ribs which begin at the umbilical rim, they des­
cribe an adoral bow on the adumbilical third of
the flank, and on the central and adventral
thirds they have an adoral concavity. The ribs
continue across the venter but are slightly wi·
der. The intercostal spaces are rounded and one
and a half times as wide as the ribs. Sporadica­
lly some ribs bifurcate at the umbilical rim.
On the subsequent whorls the whorl section
becomes higher than wide, the umbilical wall is
nearly vertical and merges smoothly into the
flank. The sides are flattened and the venter
is broadly rounded. On the internal mold the
ribs are thin, and high on the adumbilical half
of the flank, they are rounded to slightly flatte­
ned on the adventral half. At this stage the ribs
are slightly sigmoidal, with a regular alternation ·
of primary and secondary ribs. The primanes
begin near the umbilical seam, the secondaries
appear far outside the umbilical rim.
B) Up to 46 mm D (C-222-A). The orna­
mentation of the adapical portion of this speci·
men is as described above, but ad orally the ribs
become well rounded on venter and the ribs that
F. ETAYO · SERN A
on one side appear as primaries even bifurcating
appear as secondaries on the other side. The
umbilical wall becomes inclined. The border of
the aperture is plain .
DIAGNOSIS. This species is characterized
by: A) its stocky aspect, especially in the adult,
and B) its flexuous ribbing.
Measurements D Euw H W D/Euw DfH No.R.
% %
C • 222A 46 19 16 15 42 36 24(2/3)
T.I.P. Guaduas 37 16 13 1 2 36
43 30(3(4)
,,
COMMENTS. This species differs from
never has a stocky appearance and its ribs are
rme throughout its ontogeny.
OCCURRENCE. Socota Member. Upper
Aptian .
1938 Parahoplites cf. o bliquus Riedel, p. 34,
pl . 6, fig. 14; pl. 14, fig. 15.
Holotype. The specimen figured by Riedel, 1938,
EL 6, fig. 14.
!-!ypotype.(U.N.C.M.H.N .C.P.Ht.N. 36, 37).
Type Locality. Utica (Cundin amarca).
A) Up to 23 mm D (C-200-B). The earliest
part of the phragmocone has a subrectangular
whorl section, slightly wider than high. The sides
round into the flattened venter; the umbilical wall
is subvertical but merges smoothly in to the flank.
The ornament consists mostly of fine primary
ribs beginning at the umbilical seam and trending
linearly on the umbilical wall; on the adumbili·
cal half of the flank the ribs are feebly convex
adorally but on the adventral half they describe
a sl ight adoral concavity that is accentuated
on some ribs; the ribs cross the venter straight.
On the internal mold the ribs are high and
rounded. Sporadically there is a rib that appears
to begin near the umbilical margin or on the ad·
ventral third of the flank from the adapical side
of the subsequent rib. On the internal mold the
intercostal spaces are slightly wider than the ribs.
In the innermost whorls some ribs flare out on
the adumbilical half of the flank; on one flank
a rib may appear as a primary but on the other
side it appears as a secondary.
B) Up to 36 mm D (C-200-A). The whorl
section becomes higher than wide (H : 13; W : ll),
and there is a marked tendency for a regulariza­
tion of the alternating primary and secondary
ribs, bu,t this is periodic�Jiy interrupted by cou­
ples of successive primary ribs; the secondary
ribs are better defined toward the adventral
third of the flank but they.,are narrower along
the ventral median line than on the ventral
margin.
 zoN ATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES 71

DIAGNO SIS. This species is characterized
by: A) the increase in height of the whorl sec­
rib
tion B) the irregular accentuation of the
usity during the juvenile stage , and C) the
flex�
tendency of the ribs tow:ard flattening on the
adventral third of the whorl side during the
adult stage, giving the ribs a rursiradiate aspec
. �
Meumementa
200 B
C •
C • 200A
COMMENTS. At first glance these speci·
mens appear similar to the specimen described
by Basse (1928, p. 136, fig. 14; pl. 8, fig. 5) from
·
·
Villa de Leiva and identified as Colombicera.s
·
Lectotype. Here designated the specimen illus·
trated by Riedel, 1938, pl. 6, fig.
11-12.
Hypotype . (U.N.C.M.H.N.C.P.Hy.N. 38, 39,40)
Type Locality. Utica (Cundinamarca).
A) Up to 20mm D (C-199-B,C). The e arliest
part of the last whorl is slightly wider than
high; the sides are slightly convex, they round
smoothly into the arched venter and the subver·
tical umbilical wall.
D Euw H W D/Euw D/H No.R.
% %
Ribs begin on outer half of umbilical wall,
23 9 9 19 39 3 9 ' 38
trend slightly prorsiradiate on ·the adumbilical
36 13 13 11 36 36 29(3/4)
third of the flank and then trend radially to the
ventral margin. The ribs h ave a rather uniform
thickness to the adventral third of the flank
where they broaden and cross the venter with a
delicate adoral arc; the ribs are well defined and
sep arated by intercostal spaces about one and a
half times as wide as the ribs.

ka,:steni (Marcou) , which later was included Sporadically a secondary rib branches
within the genus Pegnace ras (as Penaceras}C antti­ from the adoral slope of a primary rib near the
Chapa, 1963, p. 55), however, I feel the latter umbilical rim. An alternation of intercalary and
genus should be restricted to the Mexican spe­ primary ribs become dominant on the internal
cies ·in the type species of Pegnaceras:the distan· whorls, some of the primary ribs become stron·
b'
ce etween the ribs does not increase with growth , ger. The ribs become progressively wider apart
thefhlckening of the ribs is "transversely elon­ with growth.
gate" (Humphrey , 1949, p. 142), and the early
development of the sculpture unknown, conse· B) Up to 4 0 mm D (C-200·0). At about
qtiently I consider the rursiradiate adult ribs an a diameter of 20 mm the ribs have a regular pat­

undiagnostic character; it is also present on
"Acdnthoplites ?adkinsi'' and A an th plite ?
sandidgei" described by Humphrey from Mexico
but no assigned to Penacera.s by CantU-Chapa.
1\ll&Se's illustrated specimen is poorly preserved
and does not warrant an extensive discussion.
c
tern of alternating uniformly raised primary in·
" o s tercalary ribs, the latter sometimes inclined to·
ward the preceding primary rib as if bifurca­
ting from it at about the adumbilical third of
the flank. On the adoral fourth of the last
whorl the ribs are rme, delicately falciform.

· .· OCCURRENCE. Socota Member. Upper C) Up to 50 mm D (C-5 14-B). The whorl

A,ptian. section is subquadrate with flattened venter and
rounded periventral margins; the umbilical wall

�iefe tes obliquum £Riedel)

is subvettical and rounds smoothly into the sides.
�'-·-·
•.· ·.· · ••
.. M 3; Pl. 6, fig. 7 ; Text-fig. 7P.
L \l.g. The ornament consists of strong ribs as in
(Mozph I). the previously described specimens; the primary
·
' .

ribs begin at the umbilical seam, describe an

. 1938 Parahoplites obliquus Riedel, p.33, pl.6, 11.doral concavity on the umbilical wall and the
figs. 11-13, pl. 12, fig. 9, pl. 14, fig.14. adumbilical third of the flank, on the middle
third the ribs describe a weak adoral convexity ,
1957Parahoplites obliquus Riedel, Btirgl, p.135 on the adventral third they describe an ado­
ral concavity then go straigh t across the venter.
1964 Parahoplites obliquus Riedel, Etayo-Ser-
· . na, pars, p. 118. At this stage the ribs are acute on the
flanks and broadly flattened to rounded on the
1965 Colombiceras obliquum . (Riedel) , Casey, venter but are slightly more flattened on the
.. · .
p; 420. adapical part of the last whorl than on the ado­
ral part.
)968 Colom biceras obliquum (Riedel), Etayo·
·. .· .· .
Bema, p. 34, fig. 7, tab. I. COMMENTS. This mozph is characterized
by: A) its early whorls with weakly sinuous
.

197.0 Colombiceras obliquum (Riedel), Wied· ribs, slightly wider on venter, some of them
, mann, p. 4 84, ex rev. of Etayo-Serna's p a· sporadically emphasized or bifurcating near the
per. umbilical rim, B) at lntennediate stages o f
growth the ribs are progressively more distant
, 1972 Colombicei'as obliquum (Riedel), Etayo· from one to another with primary and interca·
' Serna, p . 45 . lary ribs alternating, and C) throughout the onto·
geny the ribs preserve their delicate flexuous
1976 C�lom biceras obliquum (Riedel), Etayo· oblique trend.
�' ' Serna et al, p. 225, tab. II.
72 F. ETA YO - SERNA
Measurements D Euw H w DfEuw D{H N.R.
"1. % C·225·o
C·199·B 22 10 8 8 45 36 44
C·199·A 20? 87 8? 8? 40? 40? 22(1/2)
C·514·B 60 2 2 16 1 44 32 1 8(1/2)
COMMENTS. The specimens illustrated
by Riedel represent juveniles to young adults
(1938, p.33,34; pl.6, figs. 1 1-13; pl.12, fig.9;
pl.14, fig. 14) agreeing fairly well with the early
whorls of the specimens here studied, on them
the ribs are not yet broadened on the venter,
but distant spacing of the ribs after the crow·
ded stage is visible. This species differs from
Riedelites quebradanegra by its more flexuous
and projected ribs and because its whod section
is not compressed.
OCCURRENCE. Lower San Gil Formation
(Segment B). Upper Aptian.
Riedelites [,ilosum». sp.
Pl.7, fig.1; Text-fig. 7U.
Holotype. C-225·6 (U. N. C.M.H.N.C.P.Ht.N.5 3)
Paratype. (U. N. C. M. H. N. C. P.Pt. N.57,58)
Only internal molds are preserved. A) Up
to H:13 mm (C-225,7 ,8). The whorl section is
subrectangular, approximately as wide as high,
it has rounded periventral margins, broa� ly ar·
ched venter and short, approximately 45 incli·
ned umbilical wall.
The ornament consislb of fine rounded
ribs with some beginning at the umbilical seam;
these describe a slight adoral concavity on . the
u mbilical wall, then trend prorsiradiate descri·
bing a weak adoral convexity near the middle
of the flank and an adoral concavity, more or
less accentuated, on the adventral third, then
they arc adorally on ven ter; the adapical side of
the ribs is steeper than the adoral side; the ribs
have a rounded top.
The intercostal spaces are three times as
wide as the ribs on venter, they narrow on the:
umbilical margin to one and a half times as wide;
Up to H : 20 mm (C-225·6). In this speci- .
men the whorl section is depressed, wider than
high; there is a continuous curvature from one
flank to the other; the umbilical wall is very re·
duced. The ribs are very prominent, they cross
the flanks, varying from weakly flexuous to al· .
most straight; they are slightly wider on venter .
and rounded; one or two of these ribs become
emphasized and bifurcate . near the middle of ·
the flank; no tuberculization is present; the in- ,
tercostal spaces are twice as wide as the ribs and .
channel like on venter.
DIA G �OSIS. This species is characterized
in the adult stage by: A) Its wide subrectangu­
lar whorl section and B) its annular ribs and
deep intercostal spaces.
Mea.surements D H w
? 13 12
C•225·6 ? 20 23
C·226·7 ? 13 13
C·225•8 7 13 13
COMMENTS. This species differs from
the previously described species of Riedelites
because it is the only one with an extremely de·
pressed section and flaring almost annular ribs.
OCCURRENCE. Upper Aptian.
·
k�J:;:]i"'\ p;
Genus
-P:A'HOPI:�Ti���,·�t
Parahoplite'JJ Anffiula,-· ·�"899
·· ·
Parahoplites Anthula, 1899, p. 109.
\lliiiiL rt oaq ttr ll'u )Uiiji'-G!Ejf¥,.. "I 9cW����::-...Lc : �
"
· ,� - .
TYPE SPECIES. Parahoplites melchioris Anthu·
Ia, original designation.
.E..a�hoalites (?} huba£hi n ..!!!.:
Pl. , fig.l.
Holotype. C-127 (U.N.C.M.H.N.C.P.Ht.N.54)
The description is based on the holotype.
The earliest whorl observed is the antepenulti·
mate. The umbilical wall is subvertical, the
umbilical rim is rounded. The ornament consists
of a regular alternation of emph asized primary
ribs which begin at the umbilical seam and flare
out slightly at the umbilical rim ; the ribs appear
prorsiradiate up to the involution line. Between
these ribs there is a tapering intercalary rib that
disappears before reaching the umbilical rim,
this rib is inclined toward the primary prece­
ding it, but does not connect to it. This pattern
may be interrupted by a strong secondary rib
that reaches the umbilical wall but in these ca·
ses the primary ribs are much more prominent.
On the penultimate whorl the alternation of the
ribs is well established with the intercalary ribs
usually but n ot always extending to the middle
of the flank, then they cross the venter' in a wi-
de adoral arc.
A few primary ribs also bifurcate near the
umbilical rim. Beginning at the periventral mar­
gin the ribs curve forward, so appearing wider
on the venter than on flanks, but they are not
flattened . At the beginning of the last whorl
a primary ribs bifurcates near the middle of the
flank. The shell is thick, the ribs are prominent
with symmetrical sides and rounded top on the
flanks, on the venter the adapical side of the
ribs is longer and more inclined than the adoral
side . Adaperturally the ribs have a moderate
biconcave profile that at times is nearly straight.
The ven�r is relatively wide, but the
maximum 'in flation of the whorl Is at the umbi­
lical rim. There is a striking development of the
umbilical wall from the penultimate wh orl to
the apertural part of the last �horl : at the be·
ginning it is subvertical to weakly convex but
it becomes retracted dorsally sloping toward
the venter of the preceding whorl .
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONlTES 73

The central branch of L of the suture has NOTE : This morph is interesting because of the
tendency to become asymmetrical, usually decreasing in number of ribs from the
� oser to the adumbilical branch. inner whorls to the outer ones and
their appearance at different levels.
DIAGNOSIS. This species is characterized
A) the regular alternation of primary and
alaryearly in the ontogeny , B) its rounded
�rcom
ui'
Meuurement D Euw H W D/Euw D{H No.R.
b ·
i
ribs, the fanner stressed adumbil c a­ % '%
c . 1 27
IlY tr
36 14 16 1 4 39 41 39
subrectangular whorl section and C) the change
wall through-
in the inclination of the umbilical COMMENTS. This morph differs from
•

out the onto geny , from subvertical to concave.
MllljUUrBDJBnt.l 0 Euw
c-127 757 26
COl\'IMENTS. This species differs from
Parahoplites melchioris Anthula ( 1899, pl.S,
figs. 4,5) from Russia by the ribs .be�g more
strongly flexuous and the whorl section ts lower.
The same species as illustrated by Sinzow (1907,
p.458, l)L2, fig.l) li�ewise �as more flexuous
ribs thart the Colombian spectes. The topotypes
of Anthula's species illustrated by Casey (1965,
text�figs;146, f.g, h-i) emphasize the subquarlra­
te whorl section of the type species.
Parahoplites melchioris An thula as illustrated
by Glassunova (1953, pl. II) because the whorl
H W DfEuw D{H N.R. section of the Caucassian fornt is m ore circular
"to % and the ribs are more strongly projected ante­
32 ? 357 437 44 riorly on venter, but it lacks the retraction of
the ribs along the ventral median line.
Parahoplites multicostatus Sinzow (1907,
p. 459, pl. 2, fi g. 5-7) has a whorl section infla­
ted on the adumbilical third of the flank and
convergent on the adventral third but on the
Colombian species the whorl section tends to
be slightly compressed; the bifurcation of the
primary ribs ceases earlier in the ontogeny of
the Colombian species than the Caucassian fonn.
O CCURRENCE. Upper Aptian.

O CCURRENCE. Unnamed beds above the Par;.a{l,oiJ,lite$..trWrul.. n. sp.

Socota Member and below the Capotes Member.
Upper Ap tian.
This species .is named in honor of the late
geologist Enrique Hubach director of the Gelo­
1
gical Survey ofColombia trom 1950 to 1957.
Pczrohoplites hubachi n. sp.
P}; 5, fig.6; text-fig. 5F
···
iParatype; (U. N. C. M. H. N. C. P. Pt. N. 59)
· ·- • - The innennost whorls are not clearly ob­
.. .
.
Pi.� , 1ig. 5; Text-fig. 5G.
Holotype . C-127 (U.N.C.M.H.N.C.P.Ht.N. 55).
Paratype . C·l27 (U.N.C.M.H.N.C.P.Pt.N. 60).
The description is based on the holotype,
The protoconch can not be observed; the se­
cond whorl is smooth but under high magnifi·.
cation broad puckers are indicated on the test
near the involution line; at the beginning of the
(Morph I) third whorl rounded ribs are visible, the latter
differentiate into primary and intercalary ribs
at the beginning of the fourth whorl, the pri­
mary ribs appear at the middle of the umbilical
wall whereas the intercalary ribs begin near the

�rvable; The penultimate whorl is ornamented

"!ith.rlbs regularly distributed between primary
�d Intercalary; the fanner begin at the umb ili­
•.c:� seam l1Jld they trend projected and at the
�'W�� time they describe a feeble adoral conca­
��tY on the adum))ilical third of the whorl side,
tqeyalso flange out and bifurcate. Between tw
o of the last whorl (4th?) the ribs develop a slight
,:of ��-!H! primary ribs th ere may. be one or two
lli�rclll ary ribs which begin at different points
P�Proaching the umbilical rim.
'��f• _in general the ribs are high with
raterdb y.roun ded but narrow top; they are sepa­
lntercos� spaces almost one and a
;·'h· lll r· as Wlde as the nbs.
•
umbilical rim ; at this stage the ribs are blu� t1
not very high, they are separated by intercosw
spaces slightly wider than the ribs.
The umbilical wall is rounded and passes
into the slightly convex flank. At the beginning
concavl ty on the urnbilical wall and slight conve·
xity on the central third of the flank, then
describe a concavity on the adventral third and
steep si­
project on venter where they widen in a visor­
like fashion. At this point the whorl section is
as high as wide with the maximum width at the
umbilical margin but there is a constriction of
the flanks along the periphery giving the who rl

�>; '!'he earli�st portion of the last who has rl

a stubby pyriform aspect. On the last whorl the
re:61v if_ attened primary ribs flange out on the adumbilical third
_ _

umbtl lc� wall which rounds in to the of the flank.

J
Yte�at�ned ven ter. Along the ventr
s1des that likewise pass smoo­
'::irle
tnf'' ·

• �i �tyi e
�
._:l�f\vh no ebi�r n .
al
_ . bs are feebJy retrac�d. On the
The suture line (Paratype, N.60 ) has L
weakly asymme trical E{L and L/U2 are sub­
. n of
·• · ;bet the adap1calcati?
'•9owe 0
f
the nbs occurs,
� the two intercalary ribs
symmetrically bifid.
ore
fief.-. - h�e . the
an
u�bllJcal rim bu t the adoral rib
umbthcal seam. All the
DIAGNOSIS. This species is characterized
ribs are by: A) its small size, B) its pyrifonn whorl sec­
· (' _ Y tgh on ven ter.
tion and C) the regular alternation of primary
74 F. ETA Y O
and intercalary ribs, broadening on venter in a
visor-like manner.
Meuurement D Euw H W D/Euw D/H No.R.
% %
C-127 28 10 9 8 43 39 38
C-127 20 9 7 7 45 36 47
COMMENTS. This species resembles "Pa­
rahoplites n u midicus Sornay" from the "Zone
de Clansares. Djebel Harraba, anticlinal de Sidi
Embarka' , Algeria (Somay, 1955, p. 13, pl. 1,
fig. 16, 18) but the African species has a clear
alternation of ribs on the middle third of the
flank that does not occur on the Colombian
species, furthermore the whorl section of Sor­
nay's species is much higher than wide at com­
parable diameters (see pU, fig. 18, op. cit.). Both
species however share in common the constric·
ted periventral margins and the maximum infla­
tion of the whorl at the umbilical rim.
OCCURRENCE. Unnamed beds above
the Socoti Member and below the Capote&
Member. Upper Aptian.
Pqrahoplites (?) inconstans Riedel ?
Pl. 10, fig. 2; Text-fig. 5D.
1938 Parahoplites inconstans Riedel, p.32, pl.6,
figs. 8-10, pl. 12, fig. 11, pl. 14, fig. 13.
1949 Parahoplites inconstans Riedel, Stoyanow,
p. 99.
1957 Parahoplites inconstans Riedel, Biirgl, p.
135. . bose, the umbilical wall is oblique and rounds
1964 Parahoplites incostans Riedel, Etayo-Ser·
na, p. 1 18.
1965 Parahoplites inconstans Riedel, Biirgl, in
Campbell and Biirgl.
Holotype. Specimen described and illustrated
. . by Riedel, 1938, p.32, pl.14, fig.13.
Type locality. Quebrada Negra , al sur de Utica
Hypotype . q�s Jf1��:r�M��.N.C.P.Hy�N��i) .
Only one specimen is available. The whorls
are slightly higher than wide; the umbilical w,all .
is subvertical, it rounds into the flanks; the
intercostal proflle of venter is rounded ; the
flanks are almost parallel to slightly inflated near
their middle. The ribs are strong and rounded,
they begin at the periumbilical margin, they are
almost rectiradiated though on the adventral
half of the flank they are slightly arcuate adora·
lly and cross the venter normal to the median
line where they . are ·'Slightly flattened. On .the
internal mold the ribs show delicate compres­
sed nodes along the umbilical margin. At the
periventral margin the ribs fonn rounded shoul­
ders.
The ornament is composed of alternating
primary and seconday ribs, the latter beginning
slightly adumbilically to the middle of the flank.
The innermost part of the conclr$hows- two iri-
• SERNA
tercalary ribs between primaries but usually
there is only one. The intercostal spaces are one
and a half times as wide as the ribs.
Meuuremente D Euw H W D(Euw D(H No.R.
% %
c - &03-13 28 12 9 8 43 32 21(3/4)
COMMENTS. On Riedel's illustrations,
that which depicts the complete specimen (Rie·
del, pl. 6 , fig. 8) gives the impression the ribs
were delicat&Dy nodose along the ventral margins,
but this is not discernible on the ventral view of
the same specimen (ibid., fig. 9}. His specimen
· is larger than the present specimen. This is the
first time the stratigraphic occurrence of this
species has been documented.
OCCURRENCE. Unnamed beds above
the Socoti Member and below the Capotes
Member. Upper Aptian.
• ,�·-
·_"i_
liG.o_t:l._
p·�- "i-
RA....
TIDABr'
...,
tiTikDIWZ&mtrweumayr and
___
Genus
-
e·
Uhlig, 1881.
- .
TYPE SPECIES. A mmo-nites · pierdenalis von
Buch.
.Eneonocergs Q]galucicw.,n. sp.
Pl. it, fig.4 ; Text=figs. BB,SC.
Holotype. C-501A (U.N.C.M.H.N.C.P.Ht.N.56).
The section of the first whorl is depressed
and much wider th an hightit hm a broadly roun­
ded venter; the second whorl is roughly subglo­
evenly into the convex umbilical third of the
flank; the width of the whorl decreases and the
sides converge toward the slightly arched venter.
On the third whorl there is a marked increase in
the whorl height (H :6mm; Euw:2) and the sec­
tion becomes lanceolate : the flanks are nearly
evenly convergent toward the venter with a
slight compression on the adventral fourth; the
flanks join fonning an acute venter; the maxi·
mum inflation is at the umbilical third of the
flank.
Earch whorl overlaps the preceding one
to its umbilical rim. The ornament on the
fourth and last whorl consists of ribs which
raise as puckers at the umbilical rim and broa­
den adventrally, they have a mammillate
tubercle on the umbilical rim (8 per whorl?);
toward the adapical half of the last whorl the
periventral carinae are serrate and bear out­
wardly directed nodes.
The suture line has E/L wide, subrectan·
gular and ·has an internal branch, the other
saddles are entire, the lobes tend to be bi·
fid, only slightly indented.
D'IAGNOSIS. This species is characterized
by: A} its craterifonn umbilicus, B) its narrow
venter bordered by delicately dentate edges1
and C) the puckers on the · adumbilical third 01
the flank. :; .
 ZONATION OF THE CRETACEOUS OF CEN TRAL COLOMBIA BY AMM ONITES
Meuurementa D Euw H W D/Euw D/H No.R.
'Yo %
C·61 0A 40 3 ? ? 7 ? 8?
COMMENTS. This species resembles En­
gonoceras stolleyi Bohm {1898, pl. 5, fig. 3),
especially the juveniles attributed to that spe­
cies by Hyatt {1903, pl. 23, fig. 7-8, pl. 24, fig.
3, 4), but Bohm's species is more compressed
· and has prominent ribs on the external half of
the flank; the opposite is true In the Colom­
bian species. At equivalent diameters the lobes
of the suture of the Colombian species are more
indented but they have fewer secondary ele­
ments than in Bohm's species.
OCCURRENCE. Middle Albian.
Engono�ras.�P......
Pi.*li, fli -g_
Reference specimen C· 5 10-A.
Only one specimen is available (D:42mm).
The conch is discoidal , compressed and involute;
the umbilical wall is subvertical, it grades into
the side through a rounded rim; the flanks are
slightly convex, gently converging toward the
venter; the maximum width is near the middle;
the venter is narro w, slightly concave, it is bor·
dered by two sharp dentate carinae, the teeth
corresponding on both sides of the venter.
·
.
The ornament on the adapical half of the
. gular fo m ammillate; the tubercles do not co­
· lest
•· .
· .
whorl consists of bundles of lirae which
foi:tn taised and rounded inflations at the adum-
' �iUc.al third of the flank; the ribs have a minute
> c;onical tubercle at the umbilical rim; they trend
Jn·orsiradiate or with a slight adoral bow to the .
:(adventral third of the flank where they form a
;,. crescentic adoral concavity.
., "'"· · · ·
Near the middle of the flank the ribs tri·
On the internal mold flat, low strap­
are impressed, ending at the marginal
in clavi that are as wide as the ribs.
Slngl�, non tuberculate intercalary ribs
occur.
··
The intercostal spaces are shallower
ribs and are indicated by notches on
On the adoral half of the last whorl
. develop very fine bullae near the urn­
.. riin and a conical tubercle near the mid·
the Dank and the carinal teeth become
The suture has subrectangular, bifid
specimen is septate 'up the adoral·
preserved.
·
75
"Parengonoceras elegans Renz.'-' (1970, p.
1036) from La Puya Member, Upper Albian of
Venezuela (Renz, 1970, pl. 3, fig. 1c especially)
has a similar type of ribbing, but its whorl sec­
tion is more lanceolate than in the Colombian
species at comparable diameters.
O CCURRENCE. Middle Albiali:·
•
_Enc fl.,'_o�r;:;;4.
gs.
<?l dwqJ,ei.J!
. �n.
!X.. _.-"-·
' , . ..
• .
.
Pl.
1968 Engonoceras cf. gr. E. gibbosum [Hyatt] ,
Etayo-Serna, p. 37, fig. 9, table I.
Holotype. V. Leiva Est. 33a, bed S. .(U.N.C.M.
H.N.C.P.Ht.N. 57).
·
Paratype . (U.N.C.M.H.N.C.P.Pt.N.61).
A) Up to 101 mm (Paratype. V. L. Est.
3 3a). The whorl section is subellipitical; the
umbilical wall is inclined and merges insensibly
into the flank that has its maximum convexity
slightly adumbilical to the middle; from that
point the flanks converge toward the truncated
venter. �'hen the shell is preserved it is covered
with sinuous lirae and the surface of the shell is
divided into broad folds and narrower, depres­
sed grooves, well dermed mostly· along the
periumbilical third of the flank.
Along the periventral margins there is a
row of tubercles on each margin , they are clava­
te ad apically but adorally they become triangu­
rrespond on both sides but alternate. There is a
rapid increase in the width of the venter from
the adapical to the adoral portions of this
specimen.
B) Up to 150 mm D (Holotype. V. L. Est.
33a). The ornamentations is as described above
but on the living chamber, which is half of the
length of the whorl, a marked change in orna­
ment occurs: the venter widens noticeably and
tends to become rounded, the periventral tu�
bercles fade out but contemporaneously,
rounded nodes begin to develop parallel to the
ventrai margin but with a more internal posi·
tion (or the venter raises?); paired mammillate
tubercles develop at the boundary between the
central third of the flank and the adventral and
adumbilical thirds respectively; these tubercles
surmount raised areas of the test with the adoral­
most one bulging out giving a flaring appearance.
DIAGNOSIS. This species is characterized
by its ontogenetic development. A) smooth ,
broadly undulated conch wit.'I a narrow flatte­
ned venter and alternating periventral clavi in
the young, and B) a broadly rounded venter
and knob-like , periventral internal nodes, and
development of lateral tubercles in the adult.
Measurements D Euw H W D/Euw% D JH %
V.L.E9t.33a 101 16 52 27 16 61
V.L.Est.33a 150 25 6 6 44 17 43
COMMENTS. I previously compared this
species to Engonoceras gibbosum Hyatt (1903,
76 F. ETAYO · SERNA

p .171, pl.22, fig.6-10, pl. 23, fig. 1-6), especia­ COMMENTS. I am following Casey's (1961)
lly to the holotype (ibid., pl.23, fig.1,2), but interpretation of Platiknemiceras. Of the species
the latter has a flattened tectifonn whorl sec· referred to this genus by Casey, Platiknemiceras
tion, and a characteristically persistent deve­ gracile (Douville') ( 19 16, p. 128, pl. 16, fig. 9,
lopment of the nodes along the middle of the text- fig. 42) looks similar, however, it seems
flanks at a diameter at which the Colombian more inflated and has more elongated and pro­
species is smooth. minent crenations on the ventrolateral carinae.

This new species also has a slight resem· J:p-tiknpr:-icera$ f.e· ind. A.
blance to Protengonoceras gabbi (Bohm) in eX"tigs; ·81t;"8K7 w:.L,....
Hyatt ( 1903, p.154), but the latter lacks tuber­
cles throughout the ontogeny and has a distinc­ Reference specimen C • 5 1 OA.
tly concave venter which seems to be rounded
in extreme age (op.cit., p .155). The lobes of the The conch is involute , discoidal, the sides
suture are somewhat similar but they are are almost flat to weakly convex, there is a cons­
more lacerate in . the Colombian species. picuous beveling of the adventral fourth of the
flank.
OCCURRENCE. Upper San Gil Forma­
tion (Bed 8). Middle Albian. The ornament consists of flat; rib-like
bands that begin as bundles of growth striae
This species is named in honor of the late with a slight concavity on the umbilical rim, they
field assistant J. Duarte, of the Geological Sur· trend prorsir&diate up to the adventral third of
vey of Colombia.
.. .....
-�k�� �'!"':"'-�,�-J-t!- .:.;'->· -�·.-· •,
the flank from where they describe a crescentic

( PlatiknemiceradBata11er,
,.,.,._._ ... ,- • ·
adoral concavity to the venter. The intercostal
\.
.

Genus spaces are thread like.

195�.
The venter is slightly concave and bordered
Platiknemiceras Bataller, 1954, p.174. by two sharp carinae. A sligh tly larger specimen
shows the suture, it has the external ele·

bassei menm ordered in an almost vertical echelon. On

TYPE SPECIES. Knemiceras (Platiknemiceras)
Bataller, original desig· the adult (?) a weak median carina along the
nation . ventral line is observable but otherwise the ven­

Pigtikneuriceras colombianq 'ter is flat although the periventral carinae per·

sist. The growth lines cross the venter nonnal
PAP.lL:tEt:J�.
_

to the median line.

Holotype. C-169 (U.N.C.M.H.N.C.P.Ht. N.58) NOTE : This species is characterized by: A) the
subparallel nature of the flanks with
The conch is discoidal, involute. The um· the beveled adventral fourth, apparen·
bilical wall is vertical and has a blunt umbilical tly persistent throughout the ontoge­
rim; the ·adumbilical half of the flank is flat ny, B) its falcate, pucker-like low !'ib·
whereas the adventral half converges evenly to­ bing, and C) the tendency to develop a
ward the narrow, flat to slightly excavated ven· ventral median carina with growth.
ter; the periventral margins are sharp, they have
finely crenate carinae on each side.
·
Meuurementee. D Euw H W D/Euw% D/H %
C - 6 10A 17 3 10 6 18 69
The surface of the shell is covered with .fl.
ne, periodically raised lirae which gather in
bundles producing very low falcate ribs visibl.e COMMENTS. An unfigured species of
only with oblique illumination; they are radial Platiknemiceras (?) was noted by Breistroffer
and straight on the adumbilical and ce�tral (1952, p . 2634) as occurring with Knemiceras
thirds of the flank but on the adventral jhird in the ."Mesoalbien basal du Cundinamarca", the
they describe a shallow adoral concavity . \. characteristic suture described by Breistroffer
agrees in general with that of the present species.
The suture of the type species of Platikne­
There is an alternation of moderately ac· miceras (see Bataller, 1954; Casey, 196l, fig. 1b)
centuated and ofvery·low ribs. On the adventral has shallower accessory lobes than the Colom·
third of the flank feather structure is visible on bian species.
both flanks. On o.ne side on the umpilical
brink there is a tenuous fain t bullate tubercle (?). OCCURRENCE. Middle Albian.

DIAGNOSIS. This species is characterized

by : A) Its fmely crenate ventrolateral carinae, B)
W.ol;flf[if?�l l&:JPE:P�
the trend of the bundles of lirae , radial �n the �
Reference � eeimen. V. L. Est. 33a, bed 8.
central and adumbilical thirds of the flank, ado-
rally concave on the adventral third.
·

Only a fragment of the internal mold of

this species is available. !The conch in discoidal,
Meuuzementa. ' D Euw H W . the whorl section is evenly convex. The surface
c- 189 ? 5 30 14 of the internal mold is smooth; locally and only
 ZONATION OF THE CRETACEOUS OF CENTRAL C O L OMBIA BY AMM ONITES 77

visible with oblique illumination there are
tenuous, rib-like radial pleats.
The venter ls flat to slightly excavated, it
is bounded by two rounded, apparently conti­
nuous carinae that are broadly ondulated.
NOTE : This species is characterized by its bro-
adly wavy carinae and extremely atte-
nuated ornament. ·
very the convex sides. The whorl section is inflated
on the adumbilical third of the flank, it is sllgh­
tly compre!ISed to excavated on the adventral
third, the venter is truncated.
The adapical portion or the last whorl
of this specimen has broad low ribs which arise
from sharp periumbilical tubercles, they trend
• radially up to middle of the flank from which

point they describe a feeble concavity to end at

elevated spinose periventral clavi. At the adapi·
MEASUREMENTS. The specimen has a cal segment where the shell is preserved these
radius of 90 mm but no other measurement can nodes are interconnected by a zig-zag rib.
be made.
·

An intercalary rib begins in ..a subdued

This species is rather simi·
COMMENTS. manner near middle of the flank ·and near the
lar to that illustrated by Casey ( 1962, P,· 355, following adoral primary as if bifurcating from
fig. 1c, ld) as Platiknemiceros sp. from ' Lower it, it terminates in a spinose ventrolateral tuber­
Albian (sandy siltstone at base of Group 5) near cle. The periventral tubercles alternate.
Canta, Peru:' It is possible that this fragment
, from Est. 33a represents lhe adult ofP. colombia­ The living chamber is half the length of
no. the last whorl; no appreciable change in the or­
nament occurs except for the attenuation of
OCCURRENCE. Upper San Gil Formation ribbing and increase in spacing between the pe­
(Bed 8 ). Middle Albian. riumbilical nodes (6 per whorl). Flexuous growth
: t�a�n��· T!f�eitic;;J��!;};i��- lirae are present on the exterior of the shell
where it is preserved.
Knemiceras Bohm, 1898, p. 200.
DIAGNOSIS. This species is characteri·
TYPE SPECIES. A mmonites syriacw von Buch, zed by : A) its moderately inflated, rather pyri­
original designation. form whorl section, ll) the alternation of low
sinuous ribs, the primary ones beginning at sharp
�netW�'af semicostatum periumbilical tubercles, and the secondaries be·
.
oromemw@.�;•.ft_ll•.. fig. 2; ginning near the middle of the flank where the
Text-fig. 8G, 81. primary ribs are slightly inflated.
19l0 Knemiceras attenuatum var. semicosta­ Measurements. D Euw H D/Eu w % DJH %
tum Sommermeier, p. 350, text-fig. 18� V.L.Est. 33 81 17 40 21 50

? 1947 Knemiceras semicostatum Sommermeier,

Knechtel, p. 93, pl. 15, fig. 1, 2, 3, text­ COMMENTS. The present specimen is
fig. 3. very similar to the one referred to the "variety"
(op. cit., p. 351) and illustrated by Sommerme·
1952 [Knemicerasl semicostatum Somm., Bre- ier; the other specimens referred to the same
istroffer, p. :2633. variety by him as well as those illustrated by
Knechtel (194 7, p. 93, pl. 15, figs. 1, 2, 3), have
) 1952 Kn. [emiceras l semicostatum Somm., strm.ger rounded ribs than in the Colombian re­
·
Mahmoud, p. 8'4. presentatives. Knechtel's specimen also has mo­
re numerous ventrolateral tubercles.
?. 1955 Kn. (emiceras ] attenuatum semicosta­
. . fig.
tum Sommermeier, Mahmoud, p. 54,
··
..

27. OCCURRENCE. Upper San Gil Forma­

tion (Bed 8). Middle Albian.
'? 19 64 Knemiceras attenuatum semicostatum
Sornmermeier, Etayo-Sema, p. 121. .. Knemiceros fegnai n. SE·
?196
_
5 Knemiceros semicostatum semicosta-
. J!J;2, ftg.J . iett·tt!t!• so!..��... ..
.
tum Sommermeier, BUrgi, in Campbell Holotype. C-131 (U.N.C.M.H.N.C.P.Ht.N.59).
and BUrgi.
19S .S Knem iceras semicostatu�
Only the holotype is available, it is sep­
Sommermeier tate throughout. The innermost whorls have
Etayo-Sema, p. 37, fig. 9, Tab. I.
· ·

flat, subparallel sides and excavated venter. The

li�potype, V.L.Est. 33a, bed. 8 (U.N.C.M.H.N. subsequent whorls acquire a Ianceolate section
with flat venter; the last stage has subparallel
·
C.P.Hy.N.42). sides which merge smoothly ir.to· the arched
venter; the umbilical wall is subvertical, short
Pe. Thconch
r/�the�� e de�c�ption is based oi! �e �ypoty-
· •

ts t�yolute; the umbilicus d�ep, JS

and rounds into the flanks without a definite
rim. The involution line follows the external
1,
;,;_;�: �
0 que umbthcal wall grades evenly mto
.·
margin of the periumbilical tubercles.
78 F. ETAYO · SERNA

The sculpture consists of fold-like pro·

minent ribs which converge toward a prominent
DIAGNOSIS. Many whorled conchs;
spiny umbilical tubercle ; the ribs are arranged
,
early whorls dep ressed smooth ; juvenile whorls
with transient rounded umbilical tubercles from
as a trident: one adapical rib that reaches the
which ribs may arise ; the venter progressively
umbilical rim but without joining the umbilical
becomes acute and the ribs become accentuate·
tubercle, a very short rib dying withou t reaching
dly falciform on the flanks, well spaced and ar·
the middle of the flank ll!'d in line with the um­
ched adorally on venter.
bilical tubercle, and one adoral rib that extends
toward the umbilical rim; both the adapical and
COMMENTS. As for the type species.
adoral ribs bear a sharp tubercle in the middle
of the flank; all ribs terminate in rounded peri·
This genus is named in honor of Profes·
ventral tubercles. Between these groups of ribs
sor Luis F. Rincon, curator of the Museum of
there is a short intercalary.
the Geological Survey of Colombia.
On the adapertural half of the last whorl
� � �inconi q.
.Ri r!l.i ': .m..
the umbilical tubercle is connected to the ad·
ventral tubercle by a ridge-like rib, whereas the
-pr.; : fig�;ext-figs. sv, sw.
other ribs tend to fade out. There are six peri­
Holotype. C- 169 (U.N.C.M.H.N.C.P.Ht. N. 60)
umbilical tubercles.
Paratype . C-169 (U.N.C.M.H.N.C.P. Pt. N.62)
DIAG NOSIS. This species is characteri·
zed by : A) its ontogenetic change in whorl sec·
The conch is multispiral and evolute ;
tion from sublanceolate to subovate� B) the
the inner whorls were studied on one paratype
changing emphasis of the tubercles, in decreasing
(C-169·2). The protoconch is about 0.05 mm in
order, periumbilical, centro lateral and periven·
tral respecetively, C) its large and robust appea­ transversal diameter, spindleform and the whorl
section is crescen tic� through the first and se­
rance.
cond whorls the wh or1 se ction breomes progressi·
vely more in flated but the crescentic profile still
Meuuremeats. D Euw H W D/Euw% D/H%
persists; on the third whorl the section is almost
48
as high as wide, its maximum width is at the um­
C • l31 177 46 81i 66 27

bilical rim and it has a narrowly arched venter;

on the fourth whorl the flanks flatten, the whorl
COMMENTS. In ornamentation, this has a vaulted section and the arched venter broa·
species is similar to Knemiceras raimondi Liss6n, dens slightly ; on the fifth wh orl the flanks are
as that species has been interpreted by Benavi· planar, they grade evenly in to the rounded urn·
des- Caceres (1956, p.45 3 , pl.48, figs . 1-5, text· ,
bilical w all and the venter is slightly flattened
fig. 39), but the ontogenetic change of the whorl with broadly rounded periven tral margin.
section is retarded in the Colombian species
which attains a greater diameter and thorn-like The first three whorls are apparently
tubercles, unlike the Peruvian species . . smooth (with weak constrictions?), on the
fourth whorl {paratype C-169-1}, conspicuous,
. Knemiceras raimondi tardum Benavides transvenely elongated tubercles develop along
Cace�s (1956, p. 455, pl.48, fig. 6-7, text-fig. the periumbilical rim (six per whorl on speci·
40), i;etains the sublanceolate or . triangular men C-169-3), they are widely spaced ; between
whorksection longer than K. raimondi s.s. but the prominen t tubercles there are one or two
it changes earlier in the ontogeny than the Co· minor tubercles. Both major and minor tuber·
lombian species. cles correspond to the point of origin of the
prominent ribs. On the sixth whorl the ribs
Breistroffer (1952, p. 2634) recorded begin at the umbilical rim and trend almost ra·
from Cundinamarca ''K. [nemiceras]'Raimon ­ dially up . to the periventral m�in from which
dii Liss., tres trituberculise, .--: ." but np iUustra· point they widen on venter describing a weak
tion is available. K. pegnae also rese�bles A m· chevron-like bend on the adapical portion of
monites ebrayi de Loriol (1882, p.7, pl.l, fig.1), this last whorl. However, most ribs seem to
type species of Parengonoceras Spath (1924), start tangent to the umbilical rim, they then
but de Loriol's species h as a higher whorl section describe . a weak adoral arc on the ad umbilical
and a n arrower venter than the Colombian spe· half of the flank and a weak concavity on the
cies, and its ribbing has a purely radial arrange­ adventral h alf. The primary ribs start at the
ment. umbilical rim whereas the intercalary ribs start
sligh tly a9umbilically and in a subdued manner.
OCCURRENCE. Middle Albian.
B) Up to 35 mm. The early development
This species is named in honor of the is as previously described, but on this specimen
field assistant Luis F. Pefia of the Universidad the periumbilical tubercles of the fifth and
Nacional de Colombia. siXth whorls are strongly developed. Near the
beginning or the living chamber the whorl
· Family BRANCOCERATIDAE section tends to become fastigate and the ribs
accentuate their cumture becoming falcoid
!:Oeii\i&r"(�]nc'.fm(ce
.. ..... ...
\., , ·
_
..
r§n. g. :: f'"'.(� ""'"'r�_,. o_. �
� ·-.-.,.,., Y1i;t:'l-'"4....-.•.u.;.o...·.o -....: """ ' -'·--"'"
and well projected on venter, they have a well­
defined chevron-like adoral arch . The living
TYPE SPECIES. Rinconiceras rinconi n. sp.
�

chamber is a third of the last whorl, with

 ZON ATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
sinuous ribs that have a wide concavity on the
adventral half of the flank.
::•;, ,,, ' DIAGNOSIS. This species is charadiri·
zed by : A) the change in its wh orl section
: through the ontogeny from depressed to subfas·
- ti gate , B) the ontogenetic development of'the
ornament from smooth whorls (with constric­
tions?), to whorls covered with rectiradiate ribs,
and finally , with sigmoidal ribs with progressive
stressing of the external adoral concavity, C)
the rounded ribs and the wide intercostal spaces.
Measutementa D Euw H W D/Euw04 D/H % No.R.
C-169·1 28 13 11 9 46 39 25
C·1 69·3 34 17 12 10 50 35 21
COMMENTS. The ontogenetic develop­
ment of this species is characteristic: it does not
develop a keel at any stage, and its inner whorls
• have a widely arched venter, however, in the
latter stages the whorl section is acute ogival.
The absence of acute inner whorls separates it
from Brancoceras Steinmann (1881) and the
same characteristics separate it trom Hysteroce­
ras Hyatt ( 1900) , ll'owever, the presence of
transient tubercles and the chevron-like projec­
tion of the ribs on venter tend to give it a Hys­
te ro ce ras- !ike appeazance (compaze fig. 4 , on
plate 9 o f this paper with Hysteroceras aff.
d 'Orbigny Spath, 1934, p .486, text-fig. 169).
The presence of rounded tubercles and
the tendency to sharpening of venter with the
strong projection of the ribs on venter differen­
tiate Rinconiceras rinconi n. sp. from Eubran­
coceras aegoceratoides (Steinmann), type species
of Eubrancoceras Breistroffer ( 1952).
The Colombian species also resembles
Brancoceras (?) carinatum Collignon ( 1949,
p .95, pl.20, figs.5 , 5a, 5b) type species of
Paroxy tropidoceras Breistroffer (1951, p .267,
nom. nud.) ; the Madagascan species h as a true
carina whereas in the Colombian species the
ven ter is merely sharpened on the adult and the
species is in general more evolute .
OCCURRENCE. Middle Albian.
-· � Family L YELLICERATIDAE
� 9����-::rr;!!!E�!�Is�th:
- ribs on the last whorl .
,� . 9
1 21.
Lyelliceras Spath, 1 9 2 1 , p . 220.
TYPE SPECIES. A. Lyelli Deshayes in Leyme·
rie according to Spath; origi­
nal designation�
Ll!jqicez:g�
fis.
c��P·
�£.� ; ix:::Jis.Ji, s�.
�oloty pe. C-255 (U.N.C.M.H.N.C.P. Ht. N.61)
aratype . C-225 (U.N.C;M.H.N.C.P.Pt. N.63 ,64)
Up to 11 mm D (C-225·1). The fllSt two
, W ho rls are apparently smooth; on the adapical
..• Part
of th e third whorl there are very well mar-
79
ked sigmoid&] folds; shortly a strong acute rib
appears at the periumbical rim, beginning with
a compressed bullate node�and ,exapd� up to a
subtriangular tubercle in the in'i"mtte of the
flank; from �he cen�al � � ercles ��tq the peri­
ven tral margin the nbs wtden an� --flatten and
describe a crescentic shaped concavity to end at
a clavate periven tral tubercle ; at this stage o f
•
growth t h e ribs cross the fastigate venter as seve­
ral riblets that form acute "Vs". Between these
ribs there may be one or two intercalary ribs
which do not reach the umbilical third of the
flank, the latter appear like handle-less sickles,
but theY, are slightly less prominent than the -pri­
mary nbs. There are five primary ribs on the
third whorl. The whorl section is ovate, higher
than wide, with obtusely beveled venter along
which a wavy weak carina develops. In general,
the ribs form an elongated angular S on the
flanks .
Up to 23 mm D {C-225). At the adapical
portion of the fourth whorl, a weak bitubercu­
l ate intercalary rib extends to the periumbilical
rim is still visible but from that point on there
are no in tercalary ribs. The ribs have a more or
less accentuated polygonal profile; they begin
at the middle of the umbilical wall and lead to
the conical periumbilical tubercle forming a n a­
rrow , hi�h and acute prorsiradiate segment; the
periumbtlical and centrolateral tubercles are
connected by slightly wider almost rectiradiate
segment of the rib , and from the latter tubercle
to the periven tral tubercle the ribs are still wider
and are slightly prorsiradiate, the strong ribs
cross the venter almost normal to the ventral
median line or with a light adoral acute arc. The
involution line is immediately above the conical
cen trolateral tubercle. The tubercles along the
ventral median line are clavate and tend to be·
come conical. The whorl section is rounded
subquadrate. There are 16 ribs on the venter of
the last whorl. The intercostal spaces are 2.5 ti­
mes as wide as the ribs and are flat bottomed.
C) Up to 43 mm D (C-255). The medium
size f!)ecimen selected as holotype, has the abo·
ve characteristics with the interrupted rib profi·
le mostly when the cen tral portion of the rib
becomes strongly rursiradiate. The intercostal
space varies from two to four times the width
of the ribs on the internal mold. There are 19
DIAGNOSIS. The species is characterized
by: A) the eazly development of strong, acute
and widely spaced ribs that in longitudinal pro­
file appear as an elongated angular S, B) the
presence of adumbilical and adven tral lateral tu­
be rcles on the ribs, the latter the most promi­
nent.
Measurements D Euw H w D/E•Jw D/H No.R.
C·225·1 11 5 4 ?
%
45
%
36 8(1 /2)
C-225 23 1 0 8 7 8? 43 3 6 16
C·225 43 21 13 12 48 30 19
COMMENTS. This species differs from
Lyelliceras lyelli (Leymerie MS), in d' Orbigny
BO
(1841, pl. 74) as interpreted by Spath ( 1931,
p.3 15), by the more depressed whorl section
and rectiradiate ribs of that species. Breistroffer
(1952, p. 2634) has recorded "Lyelliceras Lye­
IIi Desh. sp." from Boyaca, but his unillustrated
record may not be relied upon; the specimens
from Pen! referred to Lyelliceras 11elli (Leyme­
rie) d'Orbigny, by Benavides · Caceres (1956;
p. 463, pl. 5 1, figs.2,3), differ from the Euro­
pean forms, as observed by Young (1966, p. 18),
because "the ribs . . . are sharper . . .". The Pe­
ruvian taxon though more similar to the Colom­
bian forms than to the European, has coarser
ventral tubercles.
�e{liceras fNifolY.,elli,orme n. sp.
:g-;·'hf."§, . -ti{. BE: " ' "' '·
Holotype. C-255 (U.N.C.M.H.N.C.P.Ht. N.62)
Paratype. (U. N. C. M. H. N. C. P. Pt. N. 65)
A) Up to 20 mm D (C-225). The adapical
part of the second whorl is smooth, the inter­
costal wh orl section has convex sides that tend
to form a fastigate venter, they merge smoothly
into the oblique umbilical wall, the adoral seg·
ment of the second whorl has broad pleat-like
ribs separated by wider in tercostal spaces; on
the third whorl the ornament is characterized
by strong sharp ribs t!tat begin at the umbilical
rim, betweeen them there is a concave intercos·
tal space with an intercalary rib appearing near
the middle of the flank side . A conical centrola·
teral tubercle is present on the primary ribs at
the line of involution. On the fourth whorl
(D:20 mm) the ornament consists of club-sha·
ped convex ribs with three tubercles: one
mammiform along the umbilical rim, a second
more prominent and conical just adventral to
center of flank and a third, clavate and still mo·
re prominent, at the periventral margin . At this
stage all the ribs are equally strong although the
intercalary ribs may have the umbilical tubercle
very reduced or occasionally do reach the um­
bilical rim; the ribs widen on the venter and
cross normal to the median line, they are sur­
mounted by a prominent clavate tubercie . The
intercostal spaces are slightly wider than the
ribs on the flank but narrower than the ribs on
the venter.
B) Up to 37 mm D (C-255). There is no
change in the pattern of ornamentation; the
whorl section is rounded subquadrate , slightly
wider than high. When the shell is preserved the
ribs seem to attenuate. The position of the tu·
bercles on the whorl section migrates slightly
outwards throughout the ontogeny.
DIAGNOSIS. This species is characterized
by: A) the ontogenetic development of the or­
nament during which the intercalary ribs beco·
me as prominent as the primary ribs, B) the
coarse, club-shaped ribs that describe a broad
adoral arc on the flanks, C) the rounded sub·
quadrate whorl section in the adult.
F. E T A YO · SERN A
Meuurements D Euw H W D/Euw D/H No .R.
., %
C·266 20? 8 7 7 40? 36? 9 ( 1 /2 )
C-2 6 6 37 18 13 14 48 36 20
COMMENTS. This species resembles Lye·
l/iceras pseudolyelli (Parana and Bonarelli)
[ 1896, p. 99, pl. 5, fig.2], especially the speci·
men from Escragnolles (France) which Spath
(1931) suggested should represent the species in
the restricted sense, rather than the specimen
referred to the same species by Spath (1931,
p .319, pl. 32, fig. 14a, b; 15a, b). In the Colom·
bian species, however, the ornament is coarser,
the clavi along the ventral median line are very
prominent and the whorl section is quadrangu­
lar. The specimen described by Riedel (1938,
p.54, pl. 9, figs . 5, 6; pl.13, fig. 16) as Lyellice­
ras pseudolyelli Par. 8, Bon., has a compressed
section and its ribs are adorally concave, howe­
ver, it could fall withing the scope of the new
species. It should be noted that on Parana and
Bonarelli's figure 1b-lc of their plate 14 the ju­
venile does not seem to have a fastigate venter
as in the Colombian species.
OCCURRENCE. Middle Albian.
L�el/iceras isaac/eai n . S£:., ..•.
Pert, ng. 1. ·
Holotype. C-255 (U. N.C.M.H.N.C.P.Ht.N.63)
A) Up to 11 mm D (C-255 ). The early
ontogenetic development was studied on several
small specimens whose growth stages overlap,
however, because no intermediate stages be·
tween them and the adult holotype were found ,
they are not considered adequate for specific
determination even though their aspect seems
characteristic; their description is present as a
contribution to the understanding of the genus.
The first and second volutlons following the
p ro to c o n ch have a whorl section which tends
to be circular but has a wide and oblique umbi·
Iical wall; on the adventral fourth the flanks
flatten and converge toward the venter to form
a low but distinct carina; at this stage the only
ornam entation clearly visible is growth lines
that describe a concavity on the umbilical wall
and an elongated adoral arc on middle of the
flank, .<next they change into an acute adoral
concavi ty whose deepest point is in line with
the beginning of the ven tral beveling; externally
to that concavity the growth lines project on
venter forming a sharp angle whose apex is cut
by the median carina.
·
Faint indications of constrictions parallel
to the growth Iirae appear with continued
growth ('?). Bundles of lirae tend to individuali­
ze as a forerunner to the ribbing, they are first
noticed near the umbilical rim.
On the third whorl the section becomes
high and h'iis flat sides. The beginning of the
.
fourth whorl is marked by the sudden develop·
ment of clavate tub@:cles at the periventral
margins. A second tubercle appears at the be·
ginning of the external third of the flank. The
 ZON ATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
distance between the ribs varies from two an d a
half times the width of the ribs.
B) Up to 60 mm D (C-255). The inner
whorls of this specimen resemble those of tile
1atgest or the previously described specimens.
1n the holotype the centr� third of the fl�k
has two tubercles: 1) a moderately developed
ovate tubercle at the boundary of the adumbili·
cal third or the flank 2) a similar but more co­
nical tubercle developed at the boundary with
the adventral third ; there is a third clavate tu·
bercle that grows outwards from the periven
tral margin. The tubercles are equidistant.
·
Along the ventral median Une there is a row o(
clavate tubercles that correspond to the peri·
ventral tubercles but are more raised. .
•.
On the in tern a! mold the ribs seem to be·
gin near the middle of the umbilical wall; the
ribs are sharp adumbilically but become
wider and flattened towards the periventral
·margin ; the ribs cross the venter as a wide band
normal to the median line and have a tenden­
cy to split.
The intercostal spaces are as wide as the
ribs on venter, they become progressively wider
toward the umbilical wall; the umbilical wall is
inclined.
DIAGNOSIS. This species is characterized
by: A) the slow development of the omamenta·
tion, B) the distribution of the tubercles on the
ribs outside the umbilical rim and their increa·
tangular
sing prominence toward venter, C) the subrec·
whorl section with strong inclination
of the umbilical wall.
Meuurements D Euw H W DfEuw D/H No.R.
% %
C·255 60 28 22 20 47 37 26
COMMENTS. This species differs from
"Lyelliceras ulrichi Knechtel" ( 1947, p.99, pl.
23, fig.1a. lb; Benavides Caceres, 1956, p.464,
pl.5 1, fig.4,5,6,7 ,; see next genus), by the ra·
pid development of a coarser, straighter and
uniform ribbing in the Peruvian taxon, the lat·
ter almost lacking any indication of adumbili·
cal lateral tubercles. Lyelliceras pseudolyellifor­
. The first and second whorls have a subelliptical
!"le n. sp. differs by its rounded subquadrate
whorl section and strong arched ribs developed
early in its ontogeny.
OCCURRENCE. Middle Albian.
. . . This species is named in honor. of the 19
century Paleontologist Isaac Lea.
. · '-����--.�.-§1���f��0il_:.J;.yJ'".:aC �;. , ., ·;���:� �?�e
'l''Y"PE SPECIES. Acan thoceras prorsocurvatum
Gerhardt, 1898, p.168, pl.4, r1 �
text-fig.16.
'
�- Uadran gular
. . . .. DIAG NOSIS. Evolute conchs with sub­
whorl section from early in the
•.··. �� geny; three rows of tubercles on the flanks:
; � .n\ltnbUical
bullae, subclavate tubercles abven-
81
tral to prominent ventrolateral clavi ; ventral
median clavi corresponding to the ventrolateral
ones; · ·-- ·· ·-
COMMENTS. The gen us diffe�Jrom Pro­
v
lyellicer!IS Spa� (1930), "{ hi<:Jl is. ·�volu!e
and hash1gh ogtval whorl section �·� JUvemle
and adult stages, and has ventrol aterai and ven·
tral·median clavi from the inception of orna·
mentation.
Species to be included in Ralphimlayites
are : "Aq.<Jntl!Q.cenz�.P.mrso.curuaturn...Gerhardt�·.
'!,�eJliruczs..._ �ri.�hLJ<nechtel". Prolyelliceras
was proposed by Spath (1930, p.65, footnote),
noting that it lacks lateral tubercles and selec­
ting as type species "Prolyelliceras .P,�Pf; Vianum,
nom. nov. = prorsocurvatum (non Gerhardt) R.
Douville. . . , 1907, p. 144, pl. ii) fig. 1,1". The
specimen illustrated by Douville is a fragment
of a juvenile or immature specimen in which
the whorl height is not yet accentuated. Wright
(195 7, L409) illustrated Proly elliceras with
'
Gerhard t s"Acan thoceras prorsocurvatum ", and
described Spath's genus as "with 1 or 2 rows of
ventrol ateral clavi and row of siphonal clavi
subordinate to ribs". However, of the species
with ogival section for the living chamber only
periventral and mid ventral clavi occur on Pro­
lyelliceras peruvianum Spath, sensu Benavides
Caceres, and Prolyelliceras lobatum Riedel
whereas Gerhardt's species has two additional
tubercles on the sides as indicated by Gerh ardt
( 1898, p.169), and substantiated by the speci­
mens here referred to his species; also contrary
to the situation in Lyelliceras Spath, in Ral­
ph imlayites it is the periumbilical bullae rather
than the inner lateral tubercles that tend to
weaken and fade out (see Spath, 1931, p.315).
This genus is named in honor of DrRalph
Imlay of the U. S. Geological Survey.
.. B,alv.himlayites atmloense n. s;e.
Pl.if, ng. 6.
Holotype. C-229 (U.N.C.M.H.N.C.P.Ht. N.64)
Paratype. C-25 3 (U.N.C.M.H.N.C.P. Pt. N.66)
·fhe protoconch is transversely elliptical .
whorl section, whereas the third whorl is
subcircular in its intercostal section. The first
two whorls are smooth, covered only with
growth lines, but they have an undulated
surface; toward the adoral half of the fourth
volution the ornamentation becomes conspi·
cuous; it is represented by strong falcoid ribs
that end periventrally at clavate tubercles.
se ribs are more prominent;
between the primary ribs there may be one or
two intercalary ribs which attenuate toward
the umbilical rim ; o ccassion ally two ribs may
join each other at the u mbilical rim.
At the bee:innin� of the adventral third of
this fourth whorl a tubercle appears, and at the
same time weak clavi-form nodes begin to deve-
 82 F.
lop along the ventral median line. At a diameter
of about 15 mm, the sculpture becomes conso·
lidated, the ribs widen adventraly, but they con­
tinue to be falcoid.
The ribs are flattened on the adventral
third of the flank and cross the venter, although
much attenuated. Some ribs swell periumbilica·
Uy, developihg a tubercle. The intercostal spa­
ces are widt;r than the ribs and the intercalary
ribs are irregularly distributed; the growth striae
..r-foUow the pattern of the ribs, on venter they
project and form a V with its apex pointing
adorally. The suture lines has E divided by
asymmepical saddle.
) PI.l1, fig.8; Text-figs. 8Z, SZ'.
B) Up to 110 m� Q (�-2 5 3 , not collected
in situ). The ornamentconsiSts of strongly fle·
xuous ribs mostly primaries but occasionally
an intercalary rib arises from the adapical slope
of the next primary rib. When the test is preser­
ved, the ribs appear broadly rounded, and
although they are narrower on the internal mold,
they still appear rounded; the flattening of the
ribs between the internal lateral and the medial
tubercles produces sulcus-like depressions on
both sides of venter. The living chamber occupies
2/3 of the last whorL
. DIAGNOSIS. This species is characterized
by: · A) its robust aspect and subquadrate whorl
section, B) the strongly flexuous ribbing in young
adults that is straightened on the living cham·
ber, C) the development of adumbilical clavi on
the early inner whorls and attenuation of the
inner lateral tubercles on the living chamber.
Meuurementa. t> Euw · H w DfEuw D{H No.R.
% %
In situ C-229 11 4 4 4? 36 36 1 2(1 /2)
"
16 6 6 6 40 40 21··
19 8 8 8 42 .42 26
.20 9 8 7 46 40 21
7 13 1 2 13 7 ventrolateral clavus.
ex aitu C-263 21 10 8 ? 48 38 21
c-253 •no 63 37 36 7 ? 'l 6
COMMENTS. This species diffeiS from
Ralphimlayites prorsocuroatum (Gerhardt), be·
cause it has less falcoid ribbing thro�ghout its
ontogeny, and develops adumbilical bullae mo­
re consistently. Breistroffer (1936, p. 156,157)
recorded "un Lyelliceras nov. sp. (gr, Gevreyi
Jacob sp.) asse z voisin du L. prorsocuroatum
Guerh. sp. non R. Douv. sp. =.Prolyelliceros pe­
ruvianum Spath) . . .", apparently from the sa·
me region as the present specimens; subsequen­
tly Breistroffei' (1952, p. 266) suggested the
association of Douvilleiceros solitae d'Orb. and
Prolyelliceras cf. Gevreyi. "Prolyelliceras"gevreyi
(Jacob) (1907�:..p .101 =A m. Lyelli, Leymerie, se·
conde variete', in Picted and Campiche, 1860, p.
198, pl. 24, fig. 7a, 7b), according to Pictet and
Campiche's text and figure, the conch is com­
pressed and high whorled (ib., p. 198, pl.24, fig.
7a), the ventral median clavi do not correspond
to the periventral ones (ib., fig. 7b).
ETA YO · SERNA
At comparable diameteiS both species ha·
ve a well marked adoral concavity of the ribs on
the adventral third of the flank.
OCCURRENCE. Middle Albian.
�. ··. a�����-=·1!,-ocera3�Y.���-1903
.
_
Tegoceras Hyatt, 1903, p. 84.
TYPE SPECIES A mmonites mosensis d'Orbigny,
original designation.
_Tgpce{f!i..,be,navid£,scaceres£ .!1 ·
sp.
? 1956 Lyelliceras pseudolyelli Parona and
Bonarelli, Benavides-Caceres, p.463,
paiS., pl. 52, fig. 3.
non 1956 Lyelliceras pseudolyelli Parona and
Bonarelli, Riedel, 1937-1938, p.54,
pl. 9, figs. 5-6, pl. 13, fig.16, in Be·
navides Caceres, p. 463.
1968 Lyelliceras pseudolyelli sensu Bena­
vides-Caceres (non Par. � Bon.)1 Eta­
yo-Serna, p. 37, fig. 9, Tab. I.
Holotype. V.L.Est. 33a (U.N.C.M.H.N.C.P.Ht.
N. 65).
Only a single specimen is available. The
conch is evolute; on the ·adapical portion of
the last whorl the ribs �egin in a su.bdued man­
ner near the umbilical rim, slightly later they·
tend to develop a weak bullae; just adventrally
to the middle of the flank the ribs develop a
small conical tubercle; from the umbilical rim
to the subcentral tubercle, the ribs are promi­
nent and rectiradiate or weakly convex adonijly;
on the adventral third of the flank the nbJ; des�
cribe a concavity and become flatte�ed,.,:th.ey,
end at a very prominent outwardly directed
Occasionally a rib starts higher on flank
than the otheiS. There are 24 ribs on the last
whorl. On the adoral half of the last whorl the
ribs look like broad folds. The intercostal spa­
ces are slightly broader than the ribs on the in·
temal moid, subequal when the test is preserved.
The whorl section is higher than wide,
and is truncated pyrifonn, with · the maximum
inflation near the boundary between the adum­
bilical· and central thirds of the flank. On the
venter there is a row of low clavi along the ven·
tra! median line.
The ribs• do not oppose each other but al­
ternate from one flank to the other. The umbi·
Jical �wall is short, inclined and rounds into the:
flank. . ..
DIAGNOSIS. Tbis species is characterized
.
by: A) its sinuous, club-like ribs, B) its ven"trally
truncated pyriform whorl section.
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
Me uurementl. D Euw H W D(Euw D/H No.R.
"fo % Pl. 15, fig. 1; Text-figs. BR, 88
v. L. En. 33a 80 32 31 1 40 39 24
" at 1 1 22 1 6 1 1 ? 1938
COMMENTS. This species resembles
-,"Acf1pthocems Seunesi Parona and Bpnare_lli"
).
( 18 97, p. 100, pl. 5, fig. 9 =18goceras , a more
evolute sp.ecles than T. camatteanum (d Orbigny),
but Parona and BonareUi's species has a more
rectangular whorl section than the Colombian
species, furthennore, the former Is more spuse·
ly and strongly tuberculated.
The specimen figured by Benavides.C&ce­
res (1956, pl. 6,.3',
fig. 3) greatly resembles the
Colom bian specunen even in the pattern of the
suture line, but L
is narrower on the Peruvian
species. In my opinion the whorl section of the
Peruvian separates it lrom Lyelliceras pseudol­
yelli Parana and Bonarelli.
OCCURRENCE. Upper San Gil Forma­
tion (Bed 8). Middle Albian.
This species is named in honor of the Pe·
ruvian geologist Dr. V.E. Benavides·Caceres.
TYPE SPECIES. Lyelliceras scheibei Riedel,
1938, p. 55, pl. 9, fig. 7' 8;
pl. 13, fig. 17.
DIAGNOSIS. Homeomorph ofLyelliceras
Spath (1921) in its early development; very eva­
lute, the whorl section changes from reniform
in the early whorls to vaulted in the intermedia· .
y
te stages to subrectangular with flat aullel si­
des in the adult; the living chamber o the adult
tends to become rounded.
The early whorls are smooth, the subse·
quent whorls have straight ribs that appear as
if bifurcating in a V fashion from periumbilical
tubercles; on the living chamber of the adults
the ribs are biconcave .
All primary ribs bear, A) a periumbilical
mammill ate tubercle, B) a clavate to acute pe­
riventral tubercle , C) a ventrolateral clavus, and
D) a conesp onding ventral median clavus. On
the living chamber of large specimens the
tubercles tend to disappear. The suture line has
wide subrectangular, subsymmetrical· saddles, L
is dominantly bifid.
COMMENTS. Codazziceras differs from
Protacanthoceras Spath (1923) because it has
a different ontogenetic development of the or­
namentation, the ribs become thinner and bic·
concave, and the ventral tubercles disappear;
Protacanthoceras Is small and involute when
· compared to Codazziceras.
This genus is named in honor of the 19
.century geographer Agustin Codazzi.
83
Cogazzicergs scheibei (Riedel)
Lyelliceras scheibei Riedel, p. 55,
pl. 9 , figs. 7, 8; pl. 13, fig. 17.
1954 Texanites aff. seiTato nuilf;natus (Red­
tenbacher), Biirgl, p.32 [ex spec. HB-
81A. SGNMP].
1957 Texanites aff.semztomarginatus (Red­
tenbacher), Burgi, p. 139.
non 1957 Lyellicems scheibei Riedel, Biirgl, pl.
10, figs. 3, 3a.
1964 Texanites aff. semztomarginatus (Red­
tenbacher), Etayo-Sema, p. 127.
1964 Lyelliceras scheibei Riedel, Etayo-Ser­
na, p . 124.
non 1965 Lyelliceras scheibei Riedel, BUrgi, in
Campbell and Btirgl.
Holotype. The specimen described and illustra·
ted by Riedel, 1938.
Hypotype. (U.N.C.M.H.N.C.P.Hy.N.43).
Type l.ocality . Cordillera de Virginia, NW de Ia
estacion Virginia, Girardot, Cun·
dlnamarca.
A) Up to H 15 mm (C-134). The initial
four whorls are smooth ; at the beginning of the
fifth whorl, low pleats are visible, they rapidly
become prominent; they begin at the umbilical
seam and develop a thorny tubercle at the umbi­
lical rim, they are nearly mdlal on the flank and
develop a long clavate tubercle at the periven·
tral margin (::;line of involution); adapical to each
of these primary ribs an Intercalary rib develops,
it begins in a subdued manner near the perium­
bilical tubercle but does not reach it, however,
both ribs give the appearance of fanning a V; at
first the intercostal spaces are broad and shallow
on the flank. The ribs become very prominent
on the fifth whorl, however, they are depressed
near the middle of the fi�k.
On the sixth whorl the inclined umbilical
wall rounds into flat parallel sides; tlie latter are
3/4 of the height of the whorl; the adventral
fourth of the whorl Is beveled and limited ad­
ventrally by a sharp clavus; the ribs cross the
venter transversely and bear a siphonal clavus
corresponding to the periventral clavus.
B) Up to approximately 75 mm D (C-134).
The ornamentation of the early whorls is as
previously described; the livin g chamber occu­
pies two thirds of the last volution. At places
where the test is present the ribs appear as if
actually bifurcating in a V fashion from the pe­
riumbilical tubercle which elongates and slants
adorally on the inclined umbilical wall.
Toward the last half of the living chamber
the tuberculation begins to attenuate, especially
the periventral clavi, and the ventral median 11·
ne of clavi disappears, leaving an arched ribbed
84
venter; the ribs become biconcave and split Ion·
gitudinally in several riblets; the angle between
the ribs decreases. .
C) Up to 96 mm D (C-134). On this speci·
men only the ornamentation of the living cham·
ber is observed ;the ribs that bifurcate from the
umbilical tubercle decrease their angle and ap­
proach each other, furthermore, they tend to
divide into several riblets (up to three); in this
specimen the ventral median row of clavi,
though very attenuated is still recognizable.
DIAGNOSIS. This species is characterized
by : A) its large size, B) its prominent ornament
of alternating (or bifurcating) long and short
ribs, C) the arrangement of the tubercles,
spinose periumbilically, clavate ad ventrally and
ventrally, D) the biconcave ribs on the adult
and the loss of the tuberculation.
Measurements D Euw H w D/Euw
% %
C-134 7 ? at 15 1 3 7 7 1 6( 1 /2)
C·134 757 39 24 2 2 527
C•134 96 5"3 34 3 4 56
. COMMENTS. Rit�del (op.cit., p.55) dis·
tinguished his Lyelliceras scheibei from the
· bercle at the beginning of the adventral third
other known species of Lyelliceras by the alter­
nation of long and short ribs; he also compared
his species to A mmonites ospinae Karsten but
the union of the ribs at the periumbilical tu­
bercles and the whorl section, wider than high,
in Karsten's species were considered sufficient
to differentiate both species.
I adhere to Riedel's criteria especially the
latter; Karsten's species has rounded, ball-like,
strong tubercles. Karsten's species apparently
comes from beds overlying beds that yielded
A mmonites toroanus Karsten and Ammonites
mosquerae Karsten of Lower Turonian age.
OCCURRENCE. Lower Coniacian.
£odaz�icer� fina.n. m..
PJ.t3, 1ig, l7; Text-figs. 8M, 8N, 8Q, 88.
1957 A canthoceras (?) ospinae
Biirgl, p.137.
· ribs develop: they begin as if they were elonga·
1957 A canthoceras ospinae (Karsten), Biirgl,
pl.12, fig. 5a, 5b.
·
1964 Acan thoceras cf.ospinae (Karsten), Etayo­
Sema, p. 34.
1964 A canthoceras ? ospinae (Karsten), Etayo­
Serna, p. 124, pars, Burgi's identification
only.
1968 · Acan thoceras ·ospinoe (Karsten), Etayo­
Sema, p.60.
1972 A canthoceras ospinae (Karsten), Etayo­
Sema, p.46.
F. ETAYO · SERN A
Holotype. C-134 (U.N.C,M.H.N.C.P.Ht. N.66)
Paratype. (C-134 (U.N.C.M.H.N.C.P. Pt. N.67)
The first whorl is smooth; the second
whorl is inflated and has a reniform section, the
third whorl has a subelliptical section: the
flanks are subparallel , slightly convex, the ven­
ter is widely arched, the umbilical wall is abrupt
and rounds into the sides without forming a de­
finite rim. At the beginning of the fourth whorl
broad pleat-like ribs develop; they begin to be
well defined at the umbilical rim and on flank
form a faint adoral bow up to the adventral
third of the flank where they describe an adoral
concavity , they end at the perlventral margin at
claviform bulges. Wide constriction-like inter­
costal spaces are parallel to the ribs, but unlike
the latter, they continue weakly across the ven­
ter, forming wide adoral linguiform projections.
Contemporaneously very weak clavi deve­
D/H No.R lop along the ventral median line corresponding
to the ven trolateral clavi. On the fifth whorl the
ornament is well defined : there are coarse ribs
3 2 1 9 ( 1 /2) which begin at the umbilical seam, developing a
35 40?(3/� clavate swelling at the periumbilical rim where
they bifurcate into radial branches in a Y fa­
shion; both branches develop a spiniform tu­
of the flank (= level of the involution line),
from this tubercl� the ribs trend obliquely
adorally on the beveled adventral third of the
whorl, they have a sharp periventral clavus and
cross venter normal to the median line where
they attenuate and develop a clavus. The
intercostal spaces are deep and as wide as the
ribs, especially those separating pairs of ribs.
On the sixth whorl the section is higher
than wide, the umbilical wall is subvertical and
short, rounding rapidly into the fl an k s which on
their inner three quarters of height are flat and
parallel but on the adventral q u arte r are trunca·
ted; on the costal whorl section the venter ap­
pears coronate , but on the intercostal section
the venter is a wide ar�.
·
. On: tl')e seventh whorl the periumbilical
bullae are very prominent and more widely spa·
(Karsten), ced, contemporaneously one or two intercalary:
ted in a subdued manner tangent to the umbili­
cal rim, then they are rectiradiate on the adven-.
tral half of the flank and bear weak extemal la�
teral tubercles, the periventral and ventral me­
dian tubercles are regulary emphasized . On one
paratype (C-134) the intercalarly ribs reach the
umbilical rim and all ribs are much more promi·
nent on the venter.
DIAG NOSIS. This species is characterized
by • the ontogenetic development of its orna­
ment, A) with a rapid appearance of intRrcalapr .
ribs and of bifurcating ribs in a Y-like fashion , ·
B) the areentuated stressing of the periumbilical
bullae with growth, and D) the relatively fine
ribbing.
86 F . E T A Y O - SERNA
rent localities although presumably the same
stratigraphical level : I) a juvenile specimen of
about 1 20 mm diameter (as measured on figure
3, plate 8 of Schliiter's paper), and II) an adult
of about 297 mm diameter (measured on fig. 1 ,
p l . 8, op. cit.). These illustrations were accepted
as representig conspecific specimens by Laube
and Bruder, whose own figure of the species,
one adult specimen, shows a subrectangular sec­
tion comparable to that of Schliiter's specimen_
(op . cit., pi.S, fig. 1).
Juveniles of von Schlotheim's species, in
Schluter's (op. cit. p. 20, pl. 8, fig. 4) sense h a·
ve a high rectangular whorl section and nume­
rous clavi, up to 20 per whorl on the ventral
margins.
In contrast, some South American speci·
mens referred to Mammites, Cor instance that
figured by Benavides Caceres (1956, p. 468, pl.
5 5 , fig. 5·6) have a regular ornament with less
numerous, strong clavi on both sides of the ven·
tral median area, and also have a "subqu adrate"
whorl section; thus, these South American spe·
cies of Mammites have an accelerated develop­
ment of the ornamentation with few prominent
ventral clavi, unlike the type species.
The new species from Colombia also di·
ffers from the Tunisian "Mammites nodosoides
var Afra " Pervinquiere ( 1907, p. 310, text-fig-.-
118, pl. 18, fig. 2, 3) which has radially elonga­
_ted periumbilical tubercles rather than conical.
Pei:'Vinquh�re's species is also compressed. Pervin·
quiere's Mammites nodosoides Schlotheim,"For­
me typique" (ib ., p. 3 10, pl. 18, figs. 1 a, b) h as
very strong, less numerous periumbilical tuber·
cles which dominate in development over the
upper ventrolateral tubercles, the reverse ofthe
condition on the Colombian species. In my
opinion, Pervinquiere's specimen is not to be
referred to M. nodosoides sensu Schlii ter.
OCCURRENCE. La Frontera Formation.
Lower Turonian.
Msunroites �ugQ.UL !ill..
Pl. 13, fig. ; Tex-fig. 8.0.
Holotype. (U.N.C.M.H.N.C.P.Ht.N. 68).
Only the holotype is available. The first
three whorls are smooth , subcylindrical in sec·
tion; on the fourth whorl there are acute to ro· .
unded, subradial, prominent ribs separated by
shall ow, wider intercostal spaces; on the fifth .
whorl a differentiation of the ribs occurs; they
begin as folds at the umbilicitl seam and describe
a concave arc on the umbilical wall ; on the flanks
some are high and acute and other are broad
and rounded; on some of them a sharp subtrian·
gular periumbilical tubercle is present; at this
stage the whorl section.),& _ subquadrangular (H:
5mm; W: 5mm) ; the umbilical wall is oblique
_
and merges into the flanks in a broadly rounded
umbilical rim, here is the maximum whorl width ,
on the adumbilical and central thirds the flanks
are parallel but they have a slight tendency to
converge toward venter; the adven tral third of
the flank is beveled to slightly concave , with a
tubercle at each margin; the ven tral median zo·
ne is shallow and concave ; the intercostal whorl
section is subcylindrical .
On the sixth whorl the ornamentation
changes drastically : no umbilical tubercles, the
ribs become acute folds or accentuated bundles
of growth striae; they describe a shallow, short
adoral concavity on the umbilical wall, then
they form a broad arc on the adumbilical and
central thirds of the flank, then on the adventral
third they form an oblique, crescentic concavity
and extend onto the venter in attenuated form
where they make an adoral arc on the ventral
median zone ; on the periventral margin of the
venter the ribs have a bullate tubercle which Ia·
ter becomes rounded, and on both sides of the
ventral median zone they develop thorny, out­
wardly inclined clavi.
On the adoral half of the last whorl, the
ribs become riblets that bifurcate near the mid·
die of the flank forming a Y, with one branch
bend adapically and the other ad orally , so every
periventral tubercle is connected by secondary
riblets from two different primary ribs, forming
a zig-zag p attern; at this stage the whorl section
is subquadrangular with its maximum width at
the umbilical rim, the venter is marrow, concave.
The growth striae follow the course of the ribs.
DIAGNOSIS. This species is characterized
by its strong ontogene tic changes: A) it is first
smooth and has subcylindrical whorl section, B)
next it becomes trituberculate with strong ribs
(Mammites-Iike stage) , C) it becomes bitubercu­
late, gains a subquadrate, adumbilically bulging
whorl section and develops zig-zag like fine rib·
bing.
Meuurementa D Euw H W D{Euw D/H
% %
c . 263 31 11 13 16 36 41
The measurements are taken where the
preservation (?f the conch permits them.
. COMMENTS. This species is provisionally
referred to Mammites because of the transient
stage du.ring which it resembles .Mam mites; no
indication of constrictions were observed. The
only species known to me that resembles M. fu·
gax, has been referred to Be '\ ueites by Reyment
(Benueites spinosus Reyment, l9 54a, p.156, pl.
3, fig. 2; and' especially 1955, p . 5 8 , pl.13, figs.
24 , and text-fig. 25 a), but in this species the
perillmbilic;al . tubercles develop only late during
growth, contrary to the situation in M. fugax.
Incidentally , Reyment subsequently figu­
red Benueites spinosus as coming from Colom·
bia (La. Frontera, Alban; 1971, pl. 12, figs. 2a·b,
pl. 3, fig. 3a, 36 ) .
I n summary , in B. spinosus the ontogene­
tic change in ornament is from finely costate to
 ZON A TION O F THE CRETACEOUS OF CENTRAL C O L O M BIA
stro ngly costate and umbilically tuberculate,
the opposite is true in Mammites fugax n. sp.
OCCURRENCE. L a Frontera Formation.
Lower Turonian.
�mmi!fUQLt:ulatJ!LU · sp.
: 3;ii
1 1 0 ; Text-fig. 8P.
Holotype. C-310 (U.N.C.M.H.N.C.P.Ht.N.69).
Only the holotype is available. The penul­
timate whorl is of relatively moderate size , its
adap ic al segment is smooth except for ch anne l­
like oblique constrictions preceded adapically
by a rounded flange that bears a prominent
rounded UIJlbilical tubercles; the space between
two constrictions is p olygonal, re niform.
At a point, and more or less suddenly, pa­
ralleling both sides of th e ventral median region,
11. pair of clavi arise on ven ter. The ribs issue
from the um bilical rim and extend radially to
about middle of the flank where a conical-like
tubercle occurs ; from it the ribs trend slightly
prorsiradiate.
On the last preserved segment the section
is lozenge-shaped with maximum wid th at the
umbilical rim; there are single prim ary ribs
which arise from a bull ate umbilical tubercle,
they trend rad ially up to the lateral tubercle
which is adventral to middle of the flank, from
this tubercle the rib slants adorally and develops
another weak, round and feebly clavate tuber­
cle at the periventral margin, and then the rib
goes straight across the venter. Between two pri­
mary ribs with periumbilical tubercles there
may be an intercalary rib ·that disappears near
the umbilical rim ; sporadicallytwo ribs seem to
arise from an umbilical tubercle . At this late sta­
ge the whorl section is lozenge-shaped: the ven·
ter is flat, the flanks diverge to the periumbili·
cal tubercle and then they slant to the umbilical
seam fonnin,g an oblique um bilical wall.
A smaller specimen with the same rib de·
velopment and conspicuous "juvenile constric·
tions" on the inner w horls is referred to the
same species. On both specimens the areas with
test preserved are covered with fine growth
s triae crossed by strigations.
DIAGNOSIS. This species is characterized
by: A) the development of a lozenge-shaped
whorl section in the adult, B) the bullae-like
umbilical tubercles from which usually only
one rib is issued, C) the non attenuation of the
ribs on the venter.
Mea.urementa D Euw H W D/Euw D/H No.R.
% %
, C-310 (Paratype) 20 9 8 9 45 40 18
C·310 (Holotype) ? ? ? 7 ? ? ?
COMMENT.S. The whorl section of the
� a�ul t ofthis species is very peculiar and no si·
, lndar shape is known to me. Mammites revelie-
BY AMM ONITES 87
rianum Coustillier, after the whorl section lllus­
trated by Karren berg (1935, p. 1 32) has a rather
pyriform to quadrangular section.
OCCURRENCE. La Frontera Formation.
� _
Lower Turonian.
. . '..,;
·- ·
· ,. .
Family VASCOCERATIDAE
Genus [F��@sc�if�s;. .] g = 4/c<
TYPE EPECIES. Ammonites toroanus Karsten,
1858, p. 109 , pl. 4, fig. 2.
DIAGNOSIS. Conch of moderate size,
craterumbilicate to angustiumbilicate ; the
w h orl section is subcircular on the innermost
whorls, but it progressively becomes highly
convex or flattened ogival.
The constrictions are bisected by a
prominent rounded rib which extends adorally
on ven ter in a short lingu ifonn arc; the edges of
the constrictions are thickened and rounded as ·
if they were ribs; minor constrictions and rib lets
may develop parallel to the principal ones, but
usually the ornament between constrictions is
dominated by growth striae; the most prom!·
nent observable ornament is in ternal, well im­
pressed on the internal molds.
The s u t u re h as b u sh li k e E/L,
and su bsymme trical , deep, trifid L.
·
COMMENTS. This genus differs from
Thomasites Pervinquiere (1907), which has urn·
b ilical and periventral tubercles; from Neop ty­
chites Kossmat (189 5 ) which has an evenly ar­
ched whorl section, a constricted aperture and
by the absence of constriction-rib type of in ter­
nal ornam ent.
frgnci�r;:oft.g§ §.Uarezi n. sP.o;
Pl 1 3 , fi g . 2 ; Tex t-figs. SX, SY, 8 ii.
Holotype . C-310 (U.N. C.M.H.N.C.P. Ht. N.70)
Paratype . C-3 10 (U.N.C.M.H.N.C.P. Pt. N.69)
A) Up to 6 mm D (C-310). Several speci·
mens ·were dissected but the protoconch was
not observed. The smallest specimen studied
has a moderately globose shell, the whorl sec­
tion is circular to suboblong; the umbilical wall
is subvertical and rounds evenly into the convex
sides, the venter is arched. The surface of the
shell is apprently smooth.
The adapical third of the whorl shows a
ridge-like prominence across the venter and
dying about near middle of the flank; this is
interpreted as fortuituous; adapical to the ridge
and paralleling it there is a narrow shallow sul­
cus; several constrictions without adjacent rid·
ges divide the conch in to segments.
B) Up to 10 mm D (C-'3 10). At this dia·
meter the whorl section becomes compressed,
the s ides are subparallel and the venter Is arched;
the u mbilical wall is vertical and the u mbilical
rirr: is abruptly rounded.
88 F. ETA YO - S E R N A

The adapical end of this conch has a jacent constrictions; the groups of fine acute ribs
prominent rounded rib which starts near the in all cases are developed next to the principal
middle of the flank and prqects adorally as a ribs.
linguifonn arc on venter; adapical to this rib
there is a very weak parallel sulcus, adoral to lt DIAGNOSIS. This species is characterized
th e re is a p arallel c h a n n e l - like c ons­ by the ontogenetic development with : A) chan·
triction. At the end of the first fourth of the ge in the whorl section from subcircular to ogi·
whorl there is a similar more prominent roun· val with subparallel flanks, B) the rounded pri·
ded rib with its adjacent constrictions. mary ribs accompanied by constrictions that
first appear on the ventral region and then ad­
Between the second and third fourths vance toward the umbilical region during growth
of the whorl there is a similar constriction, but C) its narrow umbilicus.
it is preceded adapically by a ridge-like swelling
on the test; the third fourth of the whorl is like· Meuurementa D Euw H w D/Euw% D/H%
wise ended by a prominent rounded rib but "C-310 6 1 3 3 17 l\0
no ridge develops, however, the adapical cons· C·310 10 2 5 4 20 l\0
triction is more accentuated and wider than the C·310 20 3 10 8 15 60
adoral constriction which in addition has a well C·310 26 3 13 11 12 62
marked , thin, rounded rib just adorally.
COMMENTS. The type species A mmoni·
tes toroanus Karsten, from Venezuela (Karsten,
The fragments of test on the last fifth of 1 858, p, 109, pl .4 , fig. 2a, 2b) is a thicker
the whorl show that the growth lines describe species apparently with a larger umbilicus
a feeble and adumbilical concavity and a slight tnowever, KaJ:Sten's original h ave the umbilicus
arc on most of the flank to finally project ado· filled with matrix ; see his pl .4 , fig.2a), and most
rally on the venter like the constriction. There importantly the internal ribs are stronj!er and its
are five pairs of ribs and constrictions per whorl. venter wider than in Franciscoites suarezi.
Between the constrictions the test is smooth
except for the growth lines; the space between
the first four constrictions is equal, the fifth Leanza (1968, p. 192) compared "A mmo­
one has half the previous distance. nites toroanus' ' Karsten to the inner whorls of
his "Lewesiceras ubatense , because of the pre·
"

sence of scattered constrictions; in my view the

C) Up to 20 mm D (C-310). Specimen sli· similarity dee; not indicate affinities.
ghtly compressed. The whorl section has flatte·
ned flanks, the venter tends to be acutely ar· O CCURRENCE. La Frontera Fonnation.
che d. Lower turonian.

The whorl is divided by six prominent

rounded ribs, with a constriction on each side,
Genus j Imlayiceras 1 LeW,..�·,fz3,1w
=
1968.
Cw<JI)

of which the adapical one is wide and shallow
whereas the . adoral one is narro w and deep,
both constrictions coverge and merge at the
umbilical margin where _a channel-like depres·
sion is formed , the rib disappears at the adum·
bilical third of the flank; both the adoral and
adapical constrictions are bordered by rounded
riblets distinct only on the adventral half of
the flank; the intercostal space is covered by
�owth striae only; the spacing between the ribs
1s irregular but there are only six principal ribs
per whorl; a minor pair of constrictions separa·
ted by an acute riblet is visible adoral to each
principal rib.
Viewing the whorl from the adapical end
the segments appear to be bulgirig laterally. In
one p aratype (C-310) there are several of these
acute riblets.
·
Imlayiceras Leanza, 1968, p. 196.
'().f.y ;·.�,)
TYPE SPECIES. Imlayiceras washboumey
Leanza, original designation.
J_
Jmla� iceras (1 ralpQ.imUI.l!L n.
sp. P .r3, fig. , text-fig. au.
l�olotype. C-310 ( U.N.C.M.H.N.C.P.Ht. N.71)
Paratype. · C�310 {U.N.C.M.H.N.C.P.Pt. N.70)
A) C-310. The lnnennost observed whorl
(3rd?) is subelliptical in section (H: l ; W : 1),
with an arched venter; the fourth whorl rapidly
increases its height (H: 3mm, W : 2), with its
flanks evenly converging toward a wide flatte·
ned venter, fonning rounded slightly raised pe·
riventral carinae ; the fifth wh orl has a more ac­
centuated ventral sulcus, the umbilical wall is

subvarticat; it rounds abruptly into the flanks.

D) Up to 25 mm D (C-310). The whorl
section has slightly convex sides and an arched
venter; the width of the main ribs is greater ada·
perturally , but in other respects it has the same
appearance as the previous whorl.
The living chamber occupies the last third
of the whorl. Where the test is preserved the
principal ribs are p aralleled by the fine, attenua­
ted low and rib-like rounded margins of the ad·
The ornament at this stage consists of low
rounded ribs which radiate from the umbilical
rim and are most prominent on the external
half of the flank where they are concave adora·
lly, they disappear approaching the periventral
carinae. The adventral fourth of the flank is
weakly truncated.
B) Up to 25 mm D (C-310). The involu-
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOM�IA
tion of the shell becomes accentuated, at the sa­
me time the strength of the ribs decreases even
though their sinuosity persists ; when the test is
preserved the ribs are indicated by slightly rai·
sed bundles of growth. striae; the adapical slope
pf these.!>undles is steeper than the a.dqral slope;
1
the intercostal spaces are vai'}able m width but,
usually wider than the ribs. Ad orally the ventral
sulcus gets weaker.
•
C) Up to over 35 mm D (C-310). The
whorl section is slightly wider toward the cen·
ter of the whorl; the low, weak ribs arise at the
umbilical edge In couples and are very weak
when they cross the venter; the · periventral
carinae have almost disappeared and the venter is
flat and truncate. The apertural margin is plain,
it follows the falcoid path of the ribs. The living
chamber is about two thirds of the last whorl.
DIAGNOSIS. This species is characterized
, by : A) a juvenile early stage with bicarinate and
sulcate venter, together with prominent falcoid
ribbing; B) a later stage with flat venter and nu­
merous weak, flattened falcoid ribs that arise in
bundles at the umbilical rim and split in twos
on the external third of the flank and cross the
venter producing crenulated edges; C) a narrow
umbilicus throughout the ontogeny.
MeaJ�URmenu D Euw H W D/Euw% D/H %
C-3 1 0 25 3 14 7 12 36
C·310 35 4 20 11 11 57
COMMENTS. Imlayiceras was proposed
by A.F. Leanza (1968, p.l96), and considered
different from Hoplitoides von Koenen (1898)
"restringido al gropo de H. /atesellatus von Koe­
nen" (op.cit., p. 198), largely because of the
presence of shallow but evident constrictions in
the juvenile stage
The early juvenile stage of I. ralphimlayi
at fmt glance resembles that of lmlayiceras
washboumei Leanza, (1968, pl. 4, fig. 3-4) but
the Colombian specimens do not show constric·
tions although the intercostal spaces are slightly
accentuated (not interpreted as constrictions
here); the juveniles also have a narrow umbili­
cus and, although Leanza remarks that his
illustrated juvenile (see op. cit., pl.4, fig.3) has a
proportionately wide umbilicus, I suspect that
this width is an optical effect due to the umbili­
cus being filled with matrix on Leanza's speci­
men (see op. cit., pl.4, fig.7).
. The largest specimen in my collection al­
so resembles another specimen [Paratype USNM
132559] refened by Leanza (op. cit., pl.6, figs.
4 ,5) to his species, but• he describes the orna·
mentation of his species as consisting only of
fine growth striae, whereas on the Colombian
species the striae are in elevated bands and an
incons}:)icuous ribbing is present on the intemal
mold. Furthermore, on the Colombian species,
the ribs cross the venter and produce a railroad
aspect with Irregular crenulated edges. At this
stage the carinae are still prominent in Leanza's
species but on I. ralphimlayi the venter is alrea­
dy truncated and the whorl section is evenly
BY AMMONITES 89
. :II: ·
arched although slightly truncated on the ad·
ventral fourth of the flanks.
The whorl section of the Colombian
species . resembles that of "Hoplitoides munieri
., Pervinquiere (1� 07, . p. 2.1;7·•�·Pl �lD.�·•�- la, �b ,
2a, 2b), but the latter was descnbed as havmg
"Fiancs lisses (ni cotes, ni tubercules) reguliere­
ment mais tres faiblement bombes ,'. Leanza
suggested that this species could also be inclu­
ded in his genus.
In the Mexican Hoplitoides cf• .H. m unie­
ri figured by Kummel and Decker ( 1954, p.317,
pl.33, fig. 1,2, Text-fig.7 ,10) they indicate that,
"Low, indistinct, irre�larly spaced transverse
ribs cross the venter' ; the Mexican �species is
noticeably similar in general shape to my speci­
mens except that it is larger and has a more per­
sistenUy excavated venter; the latter species al­
so was indicated by Leanza as possible member
of Imlayiceras. The internal whorls of Hoplitoi­
des inca Benavides Caceres (1956, p. 475, pl.63,
figs. 6-11), are also similar but on the Peruvian
species the ventrolateral ridges persist longer in
the ontogeny.
OCCURRENCE. La Frontera Formation.
Lower Turonian.
This species is named in honor of Dr.
Ralph Imlay of the U.S. Geological Survey.
§ J Pervinquiere,
Genus esia 1907.
Fagesia Pervinquiere, 1907, p.319.
TYPE SPECIES. Olcostephanus superstes Koss·
mat, original designation.
�
F��a za.LJ i.Jl.s p .
1it13, fi'g. l; P1.14. fig.5;
Text-fig. ST.
195 7 Vascoceras sp. nov., BUrgi, pl.13, figs. Sa,
8b.
Holotype. C-310 (U.N.C.M.H.N.C.P. Ht. N.72)
Paratype. C-310 (U.N.C.M.H.N.C.P.Pt. N.71)
A) Up to 17 mm D (C-310). The inner
whorls were dissected in two specimens, but the
protoconch was not reached. They have a reni­
form whorl section: depressed with the flanks
and the venter form�ng a wide arc,"the umbili·
cal wall is vertical. The umbilical rim is abrupt
but not acute.
Externally the conch h as periodically well­
spaced flanges that begin at the umbilical rim
with a bullae or knob-like swelling and trend
toward the venter with a slight adoral concavi­
ty on ·the flanks; they project forward on the
venter forming an acute arc. Adapical to this
ridge there is a wide shallow constriction that
reaches the umbilical rim; adoral to the ridge
there is another constriction which however, is
narrower than the preceding one and which is
paralleled adorally by a moderate ridge which
90 F. ETAYO · S E R N A

dies out near the middle of the flank. This
second ridge is only recognizable when the test
is preserved.
Midway between two principal flanges
there is a bullae-fonn swelling ; this swelling
dies near the middle of the flank. The orna­
ment becomes. prominent almost suddenly;
the two ridges arise from a prominent conical
periumbilical tubercle and the interposed bullae
becomes at first the start of one rib, then of two.
All the ribs tend to be of equal strength.
B) Up to 33 mm D (C-310). The ornament
continues in the same pattern but the tubercles
corresponding to the original intercalary ribs
are the most prominent; inte:rspersed intercala­
ry ribs which die near the middle of the flank
may occur; the section becomes wide and has a
very high umbilical wall.
C) Up to 46 mm D (C-310). The living
chamber occupies about halt the last whorl; the
intercostal spaces as in the preceding stage are
about twice as wide as the ribs.
DIAGNOSIS. This species is characterized
by: A) its rounded subrectangular whorl section
in the adult, B) the increasing strength of its
ribs and the periumbilical mammillate tubercles.
Measurements D Euw H w DfEuw D{H
% a/o
C·310 17 6 7 11 35 41
C-3 10 33 11 14 22 33 42
C-310 46 ? on the middle of the flanks.
COMMENTS. Fagesia zanellai differs from
''Fagesi peroni var. colombiana" Fritzche (1923),
because the latter is smooth and flattened. Fa­
gesia thevestensis (Peron) (1896; p.23, pl.7,
figs. 2-3; Pervinquiere, 1907, p.325, pl.20, figs.
5a,b; 6a,b), is a similar species but it possesses
from 14 to 15 periumbilical tubercles whereas
in the. ,Colombian species the tubercles vary
fromJfto 10; further, the intercostal spaces are
wider'iand the conch is more involute at compa­
rable'dlameters in the Colombian species.
OCCURRENCE. La Frontera Fomiation.
Lower Turonian.
This species is named in honor of the geo­
logist Livio Zanella.
_____ _ __"""
(Pa na71U!m mit�
Pa \"
-
·
.
Genus
·
s Furon, 1935.
1972 Paromammites sp. Etayo-Serna, p. 47.
Holotype. L.Vs-7 (U.N.C.M.H.N.C.P.Ht.N.73).
Only the holotype is available; the inter­
nal whorls are not observable. The whorl section
is depressed, coronate; the conch is widely um­
bilicated. The ornament consists of flaring, com·
pressed triangular peg-like umbilical clavi that
begin at the umbilical seam; from these clavi
usually two but sometimes three ribs branch;
the ribs are compressed and acute and cross the
flank radially or with a feeble adoral concavity,
they bear a minute, almost imperceptible angu­
losity (or tubercle?) near the middle of the flank
and they have an acute bullate tubercle which
is slightly bent adapically on each side of a
narrow median ventral area; along the median
ventral line the ribs are very weak_, almost in­
terrupted but have a weak adoral arc. There are
16 periumbilical tubercles per whorl; the inter­
costal spaces are one and a half times as wide as
the ribs on the internal mold,they are subequal
on the test.
Toward the adapical half of the last whorl
there are deep oblique constrictions across the
flanks, they parallel the ribs and begin at the
umbilical seam. When the test is preserved, stri·
gations are observed on it.
DIAGNOSIS. This species is characterized
by: A) its wide umbilicus with vertical wall sur­
rounded by stake-like tubercles, B) its reniform
whorl section, and C) the very subdued tubercles
Measurements D Euw H W DfEuw D/H
. % %
L.Vz.· 7 25 12 9 13 48 36
COMMENTS. Paramammites colom bianus
differs form Paramammites gracilis (Pervinqui·
ere) [19 07 , p. 337, pl. 21, fig. 41, the species I
previously compared it with, by lhe more com·
pressed whorl section and strong tubercles near
the middle of the flank of the Tunisian species.
The ribs are more numerous in the Colombian
form: , 16 per h8lf whorl compared to 1 1 in the
Tunisi.lu\ species. The specimen described from
Vene;uela by Le�za (1968, p.200, pl. 1 ! fig. !- .
4) as "Paramamm1 tes polymorphus Pervmqm�­
re", has a more compressed whorl section and
has stt;�·ng, blunt ribs and tuberculation.
o ccuRRENCE. San Rafael Formation.
LowefTuronian.
· ·

ramammites Furon 1935, p. 58; .

.
Family COILOPOCERATIDAE ;·

: ��·�;
Vascoceras polymorphum Per­
)
TYPE SPECIES.
[8
vinquiere, subsequent desig· G,enus •· oplitoides von Koenen l
. ttation, R1!yitulnt 1954, foot· �
- note p. 255. · . Hoplitoides von
YP
Koenen, 1898, p. 53. :.:;:-. ., .
>:·
.
.f.aron.!.ammi�es_..£2!o_mbian!:f!. SPECIES. ffoplitoides latesellatus v'on·
.
T E
n. sp. .
Pl. 13, figs. 5, 13. .
Koenen, original designatjR,n�,
;
Hop1itoides b,e.!!!,!Zn moj�c� •
1968 Paroi7Uitnmites cf.P. gracilis (Pervinquiere), n. sp.
Etayo-Sema, p. 43, fig. 13, T. 1. Pl. 13, figs. 4 , 8.
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBiA
197 2 Hoplitoides ingens (v.Koenen), Reyment,
p. 9 15 , pl. 35, fig. 4a, 4b, 4c.
Holotype. C-310 (U.N.C.M.H.N.C.P.Ht.N.74).
paratype. C-3 10 (U.N.C.M.H.N.C.P.Pt.N.72)
..• \1 ,,, 1 �....
The innennost whorl observed on internal
mold is the thiid; the whorl section is subcircu­
lar at the ad apical portion, It has a slightly flat·
tened venter, the flanks are evenly conve�, the
umbilical wall is short an!i vertical; the tourth
whorl has a noticeable increase in height, it beco·
mes vaulted with weakly conve� sides; a sharply
beveled area connects the lateral carinae to the
flanks; this produces a narrow, deeply grooved
venter. The fifth whorl increases notably in
height and becomes lanceolate In section with
the ventral grQPve preserved.
In this particular specimen the siphon does
not run beneath the center of the ventral groove
but instead under one of the carinae (diagenetic
effect?). The only ornament discernible is low,
sinuous ribs that are visible only with oblique
illumination; on the adventral third of the flank
they are crescentic and they fade out at the ed­
ge of the periventral carinae. A larger fragment
(paratype, C·310) shows the change from a te·
nuously sulcate to a rounded venter.
The suture line has a wide, asymmetrically
bifid E/L saddle, it is very high and descendent
into the wide L lobe; L/U saddle is subrectangu­
lar.
DIAGNOSIS. This species is characterized
by: A) the rapid change in shape of the ventral
region from bicarinate to narrowly rounded, B)
its low sinuous ribs, C) the asymmetrical, descen·
dent E/L saddle and the wider, shallow and tri·
fid L.
Me811Urementl D Euw H W
C-310 34 3 20 10
CO MMENTS. Reyment (1972, pl. 35, fig.
4a, 4b, 4c) figured this species under the name
Hoplitoides ingens (v. Koenen) actual diameter
of Reyment's Sf.ecimen about Omm.] however, I venter widens quickly adorally (as observed on
using Reyment s description of von Koenen's
species one observes: A) the internal mold of
Hoplitoides ingens as illustrated by Reyment
(1955, pl. 20, figs. la, lb) from Nigeria shows a
lanceolate venter; at an equivalent diameter H.
hemanmojicae shows instead a wide sulcate
venter that just begins to round (see Reyment,
1972, pl. 35, fig. 4c), B) none of the suture
lines illustrated by Reyment of different
"subspecies" of Hoplitoides (1955, partim)
have a E/L saddle well bisected and subsymme­
trically bifid or an elongated steplike arrange­
ment ot the accessory elements causing a strong
descendent flexion of the �uture line and the
consequent asymmetry of L; for example,
compare the ontogeny of the suture line of
H. hemanmoiicae (="H. ingens ) in Reyment
(1972, pl. 35, fig. 4b) with that of Hoplitoides
rngens (von Koenen) in Reyment (1955, pl. 18,
fig, 4).
BY AMMONITES 91
OCCURRENCE. La Frontera Fonnation.
Lower Turonian.
This species named in honor of the geolo-
gist Heman Mojica R. . ,
�' .
- ��-- -
.
.
• ·
�
. -· .
Ho itoides lacabJJJmoo n. sp.
PI. 3, bg. 9. - -- -.
?1972 Hoplitoides ingens laeuis [Solger], Rey­
ment, p. 915 , pl. 35 , fig. 6.
Holotype. (U.N.C.M.H.N.C.P.Ht.N. 75).
The conch is discoidal, involute; the whorl
section is slightly inflated on its central and ad·
umbilical thiid, but compressed to rather exca­
vated on the adventral third. The venter is nar­
row, concave, boidered by two acute carinae.
The ornament consist of fine raised striae
whicti begin at the umbilical margin where they
describe a slight concavity, they then trend pror­
siradiate up to the adventral third of the flank
where they describe a crescentic concavity,
then the y extend onto venter, crossing it in a
slight adoral arc. Short, sickle-like swellings,
slightly oblique to the crescentic trend of the
growth striae , appear on the adventral third of
the flank ; the distance between the swellings
changes but it is always more than three times
the width of the swellings.
D IAGNOSIS. This species is characterized
by: A) its disk-like shape and grooved ventral
zone, B) the bundles of falcoid growth striae on
the adventral third of the flank are crossed
obliquely by claw-like swellings, C) the absence
of any periumbilical swellings.
Measurement D Euw H W D/Euw"/o D/H%
La Frontera 48 4 29 ? 8 60
D{Euw D/H COMMENTS. A species that resembles
% % the H. lacabagnae is figured by Peron (1896, p.
9 61 40, pl.l, fig.6,7) as "Discoceras cf. largilliertia­
nus d'Orb." from Algeria; his species is more
inflated on the central and adumbilical thirds of
the whorl than the Colombian species, and its
his figure), but this occurs slowly in the Colom­
bian species.
Reyment (1972, pl.35, fig.6) illustrated,
but did not describe, an apparently closely rela­
ted species from Colombia; he wrote (ib. p.915)
on the caption to figure 6 of plate 35, "individu
du type H. ingens laevis, c'est-a-dire des formes
sans acune ornamentation veritable"; the speci·
men illustrated by Reyment, however, has fine
but very well defined falcate riblets on the in­
ternal mold but in the Nigerian species descri·
bed by Reyment (1955, p1.18, fig.2) one obser­
ves puckers or folds; furthermore, on the Co·
lombian specimen figured by Reyment (op. cit.,
p1.35, fig.6), there are crescentic sweUings
which cross the trend of the growth striae on
the adventral third of the flank and especially
on the last third of the length of the last wliorl,
"
but this trait is not noted by Reyment in his
discussion.
92
OCCURRENCE. La Frontera Fonnation.
Lower Turonian.
f!
JL� itoides lgc,iraldae n. SE·
Pl:-3, 1fg."f.t�&· �- " - umbical bullae, contrary to the Colombian spe­
Holotype. C-264 (U.N.C.M.H.N.C.P. Ht.N.76)
The conch is discoidal, narro.wly umbili·
cated; the whorl section ·is lanceolate; the umbi·
lical · wall is vertical and rounds abruptly into
the flanks which are evenly convex on the
adumbilical and adventral thirds of the flank
but slightly concave to compressed on the adven·
tral third. The venter is narrow, pulley like, for·
ming an open V, paralleled on both margins by
an acute carina.
The ornament consists of seven bullae·li·
ke periumbilical swellings from which two bun·
dles of riblets diverge, each branch is slightly ar­
cuate; on the adumbilical third of the flank
each of these bundles form crescentic swellings
which fade out before reaching the ventral carl·
nae. One or two intercalary swellings occur be·
tween each couple of bifurcating ribs.
DIAGNOSIS. This species is characterized
by: A) its V like venter, B) the periumbilical
swellings from which bundles of riblets are
issued, C) its claw - like swellings on the in·
temal half of the adventral third of the whorl
side.
Measurements D Euw H W D{Euw "k DJH0/o
C-264 50 5 28 12 10
COMMENTS. This species differs from H.
lacabagnae _by the absence of periumbilical swe­
llings on the latter.
OCCURRENCE. La Frontera Formation.
Lower Turonian.
· : } ifop"litoides (?) sp. ind.
Pl.13, fig.15.
Reference specimen . C-310.'
Only half of a compressed specimen. The
whorl section on one side is compressed and
moderately shouldered; there is a keel along the
median ventral line; the ornament is characteri­
zed by ribs issued in pairs in a subdued manner
from the umbilical rim; they are rounded, pro­
minent and are radial or moderately sinuous on
the flank ending in a triangular prominence at
the ventral shoulder, these prominences are
slightly elongated in the spiral direction; between
two of the rib couples there are two (or three?)
intercalary ribs which begin approximately near
the middle of the flank and end in rounded bu·
llate prominences at the ventral shoulder.
The intercostal distances between the
primary ribs is twice the width of the ribs; the
&pace between the intercalary ribs is subequal
· _g..;,.nus,.
to the ribs.
COMMENTS. There is some resemblance
F. ETAYO · SERNA
between this specimen and Hoplitoides latese-
1/atus von K<>enen as figured by von Koenen
(1898, p1.6, fig.3a, b, only), but the ventral re­
gion of the latter is tabulate and there are peri­
cimen which is not bullate at the umbical rim
and has a median ventral keel.
OCCURRENCE. La Frontera. Lower Tu-
ronian.
{. Family COLLIGNONICEltATIDl\El /,
2!12!!! .0 -Wi���ocerasJ:llB,:S�J,* J�4�.
Niceforoceras Basse, 1948, p.694.
TYPE SPECIES. Niceforoceras columbianus
Basse, original designation.
.Jl.iceforocW!§_ boy..,g&ae�Jse.. n.
sp. 'Pf.14, fig.7.
Holotype. NRC-9 (U.N:C.M.H.N.C.Pllt.N.7 7)
The conch is involute, discoidal. On the
adapical segment of the last whorl the whorl
section is ridged, carinate, and has subparallel,
feebly convex sides, The ornament consists of
strong biconcave growth lines which begin at
the umbilical rim without any indication of tu­
berculation; they extend onto venter and join
the ventral median keel in an acute V that
points adorally; bundles of these lirae form
inconspicuous flat bands arranged in an imbri­
56 cating manner.
Periventrally there are tubercles, they are
oblique, claw shaped at the adapical part of the
last whorl but they tend to become parallel to
the median plane on the adoral segment of the
last whorl.
DIAGNOSIS. This species is characterized
· by: A) its smooth appearance, B)its low, ban�
ded ribbing, C) the change in trend of the
periventral tubercles from oblique to parallel
l
late in the ontogeny.
Measurements D Euw H W D /Euw % D/H 04
N RC-9- 50 5 29 15 10 58
COMMENTS. This species resembles N.
umbulaziforme Basse (1948, p.695, pl.25, figs.
4a, 4b, only), but on the latter species the ribs
are more prominently raised, almost acute, and
there are weak periumbilical tubercles; N. boya­
caense is also more inflated adapically than Bas­
se's species . .
OCCURRENCE. Cucaita Member. Lowe1
Coniacian.
e ro�(ce riJ .C!! , 9.,rossouv�,
.....!fi§§t
- - -···
�-
Peroniceras de Grossouvre, 1894, p.93.
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
TYPE SPECIES. Peroniceras Moure ti de Gros·
souvre, 1894, original desig­
nation.
1968 Peroniceras mouretii [de Grossouvre],
Etayo-Serna, Tab.l.
197 2 Peroniceras mouretii de Grossouvre, Eta­
yo-Serna, p. 48.
Holotype. VL6·Cu-5 (U.N.C.M.H.N.C.P. Ht .N.
78).
Only the holotype is available. The ante­
penultimate whorl has a ogival section (HW: 10;
W: 6mm� it is ventrally keeled and bears two
asymmetrical shallow grooves paralleling the
ventral median carina; the flanks have fine,
rounded sinuous riblets or ribs, separated from
each other by intercostal spaces twice as wide
as the ribs; the test is thick.
The penultimate whorl has a nearly qua­
drangular section: the umbilical wall is subver­
tical and rounds smoothly into the convex si·
des; the venter has three carinae separated by
shallow furrows; the space between the ventro­
lateral carinae and the periventral margins is
sharply truncated to excavated; on the flanks
there are prominent bar-like prorsiradiate ribs
which are feebly curved across the flanks; along
one third of the last whorl there are 14 riblets.
The ribs are issued at brink of the umbilical
wall where they develop a weak bullate swelling; .
along the periventral margin the ribs have
outwardly extended clavi.
From the periventral clavi the ribs extend
toward the ventrolateral keel and disappear
near it. The intercostal space is greater than
the width of the ribs, and increases adorally.
The last whorl has a subr�ctangular section. The
ribs are very prominent, rounded and separated
by channel like intercostal spaces twice as wide
as the ribs, the connection with the ventrolate­
ral keels has almost vanished and the periven­
tral clavi dominate over the periumbilical
bullae. The ventral carinae are conspicuous,
well raised and faintly undulated.
The suture line has a deep L lobe which
widens adapically but is constricted adorally by
E/L and L/U saddles.
DIAGNOSIS. This species is characterized
by the rapid change of its whorl section from
ogival to quadrate to subrectangular, B) the de­
velopment of rounded, well separated ribs on
which the periventral clavi become prominent.
COMMENTS. In ontogenetic development
of the whorl section this species is similar to
"Schlo enbachia Dravidica Kossmat" (1895, p .
190, pl.23, figs. 3a·d) in which the ribs on the
young are close to each other and the venter is
narrow, with only one distinct keel and the v.!;t·
93
trolateral keels only suggested, the change to
subquadrate whorl follows rapidly ; the speci­
men described by Kossmat has been re-studied
by Matsumoto who confirmed the on_togenetic
development (1965, II, p.213) and stresses the
"Gauthiericeras-like character'' of the early
whorls.
Kossmat's species, however, has an early
ontogenetic bifurcation of the ribs from the
umbilical tubercles, wherea the Colombian spe­
cies has only single, regularly spaced ribs; in this
characteristic the Colombian species resembles
"Peronicer(ZS subtrica rjn a tu m d'Orbigny var.
tridol'S4tum Schlii ter;'as figured by de Grosso­
uvre (1894, p.94, pl.lO, fig.3); although it is
difficult to compare it to the large French spe­
cimen, however, on the latter the ribs seem to
be persistently concave adorally whereas on the
Colombian specimen the ribs tend to be straigh­
tened and separated; on the specimen referred
to Schliiter's species by Grossouvre (1894, pl.ll,
figs. 1a, 1b) the ribs are very acute on the umbi·
lical rim and the periumbilical tubercles acute
and far from the umbilical rim whereas on the
Colombian species the ribs are bar-like with al·
most uniform width across the flanks.
Furthermore, if Grossouvre's assignment
of the specimen on his plate 11, fig.2 to Schlti·
ter's species is correct that species would have an
ontogenetic development with a very early oc­
currence of strong ribbing, the opposite of that
of the Colombian species.
OCCURRENCE. Cucaita Member. Lower
Coniacian.
This species is named in honor of the geo­
logist Roberto Garcia S.
Pero niceras diabloi �
n. s .
-· -'Pm, fig. 1; Tex1-figs. 1;·'9F;· 9a: ·
1968 Peronicerasaff. rousseauxi Gross., Etayo­
Sema, p.45, fig.13, Tab.l.
Holotype. NRC-3 (1965) (U.N.C.M.H.N.C.P.Ht.
N.79).
Only one specimen is available. The penul­
timate whorl has a very indistinct ornamenta­
tion of widely separated low ribs; on the folio·
wing whorl the ornamentation is rapidly streng­
thened; the ribs start at the periumbilical mar­
gin with a rounded tubercle, sporadically bulla­
te, on the flank they describe a well defined
adoral concavity, ending in compressed peri­
ventral clavi; the ribs do not show a distinct con­
nection to the ventral carinae. The ribs are flat,
low and depressed on middle of the flank.
The section of the last whorl is rounded
subrectangular slightly higher than wide; the
umbilical wall is subvertical, it rounds smoothly
into the flattened sides; the space between the
ventrolateral carinae and the periventral mar­
gins is slightly excavated to beveled, the median
ventral line is raised, and on each side of it the-
 F. ETA YO
94
·
re is an asymmetrical sulcus bordered by a con­
tinuous low carina. There are 15 ribs on two
thirds of the last whorl. The suture has a L
pouch·like, digitated, U3 is very oblique adum­
b ilically , narrow.
DIAGNOSIS. This species is characteri·
zed by : A) the rapid strengthening of its ribbing
and the dominance of the tubercles over the
ribs and the attenuation of the ribs on the mid­
dle of the flank, B) the conspicuous suture line
with extremely oblique U3.
Measurements D Euw H W D/Euw DIH
NR.C·3( 1965) 52 26 16 16 48 °k 30%
COMMENTS. This specimen resembles
those described and illustrated by Burckhardt
(1919, p.1 1 3 ; 1921, pl.29, fig.7-10) as "Peroni­
ceros cf. tricarinatum Schtilter sp. non auct.";
they are similar in the flattening of the ribs on
middle of the flanks and in the arcuate nature
of the ribs, but the Mexican species has a coar­
ser ornamentation earlier in the ontogeny than
in the Colombian species.
Peroniceras diabloi n. sp. also resembles
Peroniceros westfallicum as illustrated by de
Grossouvre ( 1 894, p.98, p1.12, fig. 1 ) but in the
French species · the periumbilical tubercles are
clearly bullate whereas in the Colombian
species they are dominantly mammillate.
OCCURRENCE. Cucaita Member. Lower
Coniacian.
�....1,.
�
,l'erv,nicer uerrai o, sp..
-·��· - "PJ.14; fig.;Text-fig. 91, 9J.
Holotype. NRC·Est.9 (U.N.C.M.H.N.C.P. Ht.N.
80).
Two fragments are available. The smallest .
one has a rectangular whorl section and although
the specimen has been slightly compressed, the
ven tral median carina is elevated, continuous
but widely undulated; it is paralleled by a lower
but continuous carina on each side; the carinae
are separated by a shallow, very asymmetrical
sulcus. The ribs are strong, prorsiradiate to fee­
hi y concave ad orally, rounded· they start at the
umbilical rim without devel�ping a tubercle
though a . slight bulging is insinuated , they end
periventrally at strong clavate to slightly obli­
que rounded tubercles; the intercostal spaces
are slightly wider than the ribs on the internal
mold.
Sporadically an intercalary rib, that ap­
pears near middle of the flank or branches
from the subsequent primary at about the same
point , may occur; the largest very flattened
(mechanically) specimen has a similar ornamen­
tation but with more definitely clavate periven·
tral tubercles.
DIAGNOSIS. This species is characterized
by �ts high whorl section, prominent ventral
med1an keel and strong,uniform bar-like prorsi-
SE RN A
radiate to concave ribs with weD developed pe·
riventral clavi and weak periumbilical bullae .
COMMENTS. This species resembles Pe­
roniceras moureti Grossouvre (1893, p. 100, pl.
1 1, figs, 3, 4 ; text-fig. 37-39), on the latter ho·
wever the ribs are slender and pointed adumbili·
cally and at the same time they develop conspi-
cuous elongated periumbilical bullae. ··
The suture line of Peroniceras moureti
Grossouvre (1894, p. 94, fig. 37, p. 101, fig.39)
is similar to that of the Colombian species in its
general aspect but in spite of the difference in
size the L lobe in the French species is narrower
and more deeply incised, whereas in the Colom­
bian species it is fan-like.
Venzo's ( 1936, p. 101, pl. 9, fig. 1a, 1b)
specimen of "Peroniceras Moureti De Gross�',
likewise has thin and acutely bullate adumbili­
cal end of the ribs.
The specimen figured by Schliiter (1876,
p . 38, pl. 41, fig. 3 , 5) as ·�mmonites cf. tridor­
satus " and referred by Grossouvre to his species,
has slender ribs with tubercles farther away
from the umbilical rim and a higher section th an
i s typical o f Grossouvre's species. The ontoge·
netic development of the whorl section shown
by Schliiter (pl. 41, fig. 5) adds height very
slowly but the tricarinate stage is rapidly �;�cquh
red.
·
The specimens referred by Young (1963,
p. 73, pl. 26, fig. 5 , pl. 27, fig. 1, t ext-fig. 13)
to Peroniceras moureti (Grossouvre , 1894), dif­
fer from the Colombian species by their crowded,
well rounded, strong and conspicuously ad orally
concave ribs; their whorl section is markedly
subquadrate also.
OCCURRENCE. Cucaita Member. Lower
.
Coniacian.
•.,1'._, .."�-
..
This species is named in honor of the geo·
logist Alvaro Guerra.
2���·- J-E-!-·o..-n-c;-<;�-c-l-oc_e_r_esJ
...�ati!,1���
.
Prionocyc/oceras Sapth, 1926, p. 80.
TYPE
-· ·. ,
SPECIES.
. . . . . ... _,�-
1950 Donjriarizceras longispinata Basse , p. 247,
Pt· 11, fig. 1·4.
1963 Donjuaniceras longispinata Basse , Young;
p. 67.
1964 Priononocycloceras longispinata (Basse ),
Et.ayo·Sema, p. 126.
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
1968 Prionocycloceras longispinata (Basse ),
Etayo-8erna, p. 45, fig. 13, Tab. 1.
1972 Prionoeycloceras longispinata
; ·
Etayo-Serna, p. 48. .,, . clavate and abventrally inclined in Basse's spe·
Hipotype. C-134 (U.N.C.M.H.N.C.P.Hy .N. 44).
Only an internal mold is available. Conch
involute, compressed, the umbilical wall is short,
rather sloping, merging into the side with a
rounded umbilical rim. The flanks are weakly
cpnvex, they connect to the planifastigate ven- . .
ter through weak shoulders.
The ornament consists of ribs beginning
at the umbilical rim with small bullate to coni­
cal tubercles; the ribs are radial on the adumbi·
lical third of the flank, they describe an arc on
the middle of the flank and an adoral concavity
on the adventral third; the ribs widen progressi­
vely toward the periventral margin where they
end at clavate, adventrally pointing tubercles
which adaperturally become parallel to the ven­
tral median line and comute . There are 6 ribs
on two thirds of the last whorl.
Wherever the test is preserved its surface
is covered with striae that parallel the ribs, they
cross the venter in an acu te V with its apex ado­
ral. Between the primaries there may be a riblet
with a similar trend but without tubercles. The
ventral median line is raised as a fine dentate
carina. The ribs tend to split longitudinally . The
intercostal distance varies from one to four ti­
mes the width of the ribs.
B) Up to 65 mm D (C-134). At this stage
the ribs are much wider, consequently , the in­
tercostal space varies from one to one and a
half times the width of the ribs; the ventral re­
gion becomes obtuse.
The suture of this specimen has a tall , ra­
mose, and bifid E/L saddle and a deep L lobe
with vertical notched sides; L/U saddle is trifid
with subquadrate accessory saddles.
DIAGNOSIS. This species is characterized
by its moderately convex to flattened subrec­
tangular whorl section and its broad flexuous
ribs.
Measurements D Euw · H W D/Euw%
c- 134 so 13 22 1 5? 26
c- 1 34 65? ? 30 22? ? ?
COMMENTS. Basse (1950, p. 245) pro­
posed Donjuaniceras as a subgenus of Peronice­
ras, with "Donjuaniceras longispinata Basse" as
type species; Basse considered Prionocyclus " only by prominent riblets on the periventral
�ayabanus Steinmann"as described by Ger­
hardt (1897, p. 197, pl. 5, fig. 22a, b) as to
"offre tous Ies caractl�res du genre Donjuanice­
ras et se place au voisinage des especes prece­
dentes". Basse's subgenus became a synonym of
Prionocycloceras Spath (1926).
. AHhough I am interpreting Basse's species
In a rather broad sense, in my opinion, it diffP-rs
95
from Prionocycloceras guayabanum (Steinmann)
[see Gerhardt, op. cit.] which has the periven­
tral tubercles inclined tOward the ventral median
(Basse ), line with a knob-like aspect; the tubercles are
cies, furthermore the latter is a more slender
species; the intercostal spaces are also larger.
• The specimen I previously referred to Basse' s
species was compared to plaster carts of Basse' s
species.
The specimen figured by Biirgl (1957, pl.
14, fig. 5, ex. spec. HB 31/10 SGN?vlP) as Prio­
nocycloceras guayabanum (Steinmann) from
the Nariiio Section of Girardot, has a more ra­
pidly widening venter and a more subquadrate
whorl section than the holotype of Basse's spe­
cies .
f
Priono$ loceras J2.0rtarum n. se. - .
-pr.·n, 'ij. l'2; Pn4; fij:1;"Text-fig: 9N.
Holotype. C-134 (U.N.C.M.H.N.C.P.Ht.N. 81).
Only the holotype is available. The conch
is moderately evolute, the whorl section is sub­
rectangular, it has an obtusely fastigate venter;
the umbilical wall is oblique and rounds smoothly
into the subparallel flanks that then round into
the venter without a shoulder.
The penultimate whorl has wide, flatte­
ned ribs of which every other or the third one is
stressed and develops a periven tral bullate to
conical tubercle; on the last whorl the interme­
diate ribs are considerably weaker and the orna­
ment appears as if only rounded prominent pri­
mary ribs were present; the ribs, usually but not
always, start at rounded prominent periumbili­
cal tubercles, they describe an adoral arc on the
adumbilical half of the flank and a sharp conca­
vity on the adventral half; the ribs end at pro­
minent cornute tubercles that are inclined toward
the median ventral line; the sinuosity of the ribs
may be more or less accentuated.
The growth lirae are biconcave; on venter
they form an acute adorally pointing V; on
the periventral margins the fine intercalary ri·
blets and growth striae ('?) may be so prominent
as to be easily observable. The intercostal space
increases adorally and is from two to three times
the width of the ribs: there are 6 ribs on the last
DfH % half whorl. The median ventral line is raised as a
44 fmely denticulated carinae.
DIAGNOSIS. This species is characterized
throughout its ontogeny by: A) the elimination
of the intercalary ribs that are later represented
margin , B) the rapid ontogenetic development
of comute periventral tubercles.
Measurements D Euw H W D/Euw% D/H%
C·134 49 18 20 16 37 41
COMMENTS. The nearest similar species
is "Prionocyclus medio tuberculatus Gerhardt
96 F. E T AYO - SERNA
(1898, p. 198, figs. 23a,b,c), which differs by
its more widely spaced ribs (13-14 per whorl);
the presence of intercalary riblets along the pe­
riventral margin in P. portarum, the subdued
mediolateral tubercles in Gerhardt's species.
The specimen referred to Gerhardt's spe­
cies by Biirgl (1957, pl.14, fig.4a,4b), from an
unknown locality, is not in my opinion conspe­
cific with Gerhardt's species because it has in·
tercalary ribs on the external half of the flank
which have strong centrolateral inflations and
the primary ribs are few.
Prionocycloceras portarum differs from
Prionocycloceras acutospinatum (Basse ) (1950,
p.247, pl. 11, fig. 5-6) by the strong omamenta·
tion early in the ontogeny and the strongly ar·
cuate ribs on the flank of the latter.
OCCURRENCE. Lower Coniacian.
�-'�··��-,.,....,
This species is.named in honor of the Geo­
logists Jaime and Nuria de Porta.
). cucaitaw..s._ ll·
Pl. 14, fig. 9, Text-M1g. 9M.
PrJangncloc;eras
1968 Prionocycloceras (?) aff. P. lenti (Ger­
hardt), Etayo-Sema, p.45, fig.13, Tab.l.
1972 Prionocycloceras aff. lenti (Gerhardt),
Etayo-Sema, p. 48.
Holotype. NRC-5 (1965) (U.N.C.M.H.N.C.P.
-Ht. N. 82).
Only the holotype is available. The conch
is discoidal, relatively involute, with the whorls
overlapping to the adumbilical fourth of the
flank; along the umbilical margin of the penul­
timate whorl there are small crescentic folds
but the flanks are smooth except for the
growth striae; on the last whorl the periumbili·
cal tubercles are mammillate with their apices
slightly inclined inwards; the ribs arise in pairs
from these tubercles; on the adapical half of the
last whorl there are fine ribs which describe an
elongated S on the flank and end periven trally
in tubercles with a chisel like point and a
q1,1adrate base and are obliquely inclined to the
median ventral line; at this point the intercostal
space is subequal to the ribs, but adorally ribs
become more distant and wider.
On the adoral half of th� last whorl the
periventral tubercles. are parallel to the median
ventral line and moderately enlarged laterally
and outwards. The intercostal whorl section at
the adapical end of the last whorl is fastigate; at
the adoral end it is subrectangular, it has an
accentuated overhanging umbilical wall which
merges through an angular umbilical rim into
the flanks which are inflated on the adumbilical
half and excavated to compressed on the adven·
tral half; the sides round smoothly into the al·
most tabulate venter which has smooth carina
along the ventral median line.
The specimen is com pletely septate
and missing its living chamber. The growth lines
follow the trend of the ribs, they cross vente!
forming an acute V with apex pointing adora·
Jly. On the adoral fourth of the last whorl there
are 8 tubercles.
DIAGNOSIS. This species is characterized
by its change in whorl section from Castigate
to truncate, by the numerous ribs which arise
in pairs from spinose umbilical tubercles and
end in comu te oblique tubercles.
MeB!IIlre ments 0 Euw H W 0/Euw % D/H0"
NRC-6(1965) 64 17? 26 18 28? 40
COMMENTS. Prionocycloceras (?) cucai­
taense n.sp., resembles "Gauthiericeras lenti" Ger·
h ardt (1897, p.77, pl. l, fig.4a, b; text-fig.3; non
Reyment 1958, p.12, non Matsumoto, 1965,
p.45); Gerhardt's species is more fmely costate
with 17-18 ribs per half whorl and has weak pe·
riumbilical tubercles; on the contrary, the new
species has a sparse ribbing.
_ sp.
''Ammonites barbacoensis" Karsten (1858,
p,111, pl.4, fig.5) has even greater similarities in
the shape of the whorl section, involution, type
of ribbing, and tuberculization, but they diffex
by the wavy carina at the adoral portion of the
conch ["dorso acutiusculo late tuberculato", ib.
p. 11]; P. (?) cucaitaense lacks the tubercles.
The association of Karsten's species with
"Ammonites Tucuyensis von Buch" is incorrect
the latter species comes from La Puya Membex
of Albian Age (Renz, 1972, p.704, 7 19) . Kars·
ten (1858, p. 111) noted that "A Barbacoensis
. .kommt selten neben dem Leonhardianus m.,
.
varicosus Sow., Tucuyensis Buch bei Barba·
• . .
coas . . . ," but later (1886, p.18) he reported a
more reliable association near Barbacoas (Vene·
zuela): "A. Leonhardianus Krst . . A. toroanlfS
.
Krst . . A. Mosque rae Krst . . .et A. Barbacoen­
.
sis Krst . . � {see also Karsten, op. cit., p. 62];
.
these ammonites probably come from the La
Luna Fomiation and perhaps represent two
different biostratigraphic horizons. "A. barba­
coensis" Karsten represents an unnamed genus
re:ated to · Prionocycloceras (as observed by
Gerhardt, 1898, p. 193), rather than being
congeneric with it.
OCCURRENCE. Cucaita Member. Lowe!
·
Coniacian.
�-
-
- -
Genus
f
t{;IOTiceEDs
ilia.: -- -�'-
:� :. /}'''d. /\ '
��r;·"lfi; {.�}.'-
..
TYPE SPECIES. Gloriace1Tl$ correai n. sp.
DIAGNOSIS. Conchs small to mocMrately
large. The earliest whorls observed are discoidal
with .fastigate, unicarinate median ventral re·
gion; the ornament consists of faint falcoid ox
biconcave ribs that cross the venter in an acute
adoral bend; the ribs mar-bifurcate on the urn·
bilical third of the flank, at this stage no tuber·
cles are developed.
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOM BIA BY AMMONITES
Shortly after, the ribs weaken on the me­
dian ventral line and develop a rounded some­
what spirally elongated external lateroventral
node; the admedian ventral end of the ribs
elongates longitudinally and which soon inter·
connect fonning almost continuous keels on
each side paralleling the median keel but the
ribs can still be observed crossing the venter.
The ribs sporadically develop weak periumbili·
cal swellings. With growth the ribs tend to stra·
ighten on flanks, become club-like and connect
the ventrolateral carinae.
The periventral tubercles are much stron·
ger than the periumbilical ones which someti·
mes are missin g. The suture line has E/L saddle
large, rectangular and bifid with the external
branch foliate and narrow at its stem; L lobe is
wide, deep, bifid (pseudotrifld); L/U2 saddle is
slender, taller than E/L saddle, bifid, phylloid ;
U2 lobe is digitated , short. U2/U3 and U3/U1
are subequal. U3 lobe is digitated, glove-like;
.U1 lobe is bifid , narrower but as deep as U2;
U1/l saddle is single, witn'a unique phylloid
head.
COMMENTS. This new genus resembles
Peroniceras Grossouvre (1894) from which it
differs by possessing an early initial stage with
flexuous bifurcating ribs that cross the carinate
venter forming chevrons, a character retained in
some adults but absent in Peroniceras [see for
instance Peroniceras dravidica Kossmat, (1895,
p.190, pl.23, fig.3)], and by having undulated
ventrolateral carinae. Major differences are pre­
sent on the suture line.
The suture line of Peroniceras as figured
by Grossouvre (1894, p.44, text-fig.37) has
a wide E/L with a narrow, highly incised digita­
ted L. U2{U3 saddle is accessory to L/U2; such
interdependency of L/U2·U2/U3 saddles is also
observable on "Peroniceras cf. dravidicum Koss­
mat sp." as illustrated by Spath (1921, p .295 ,
pl.23, fig. 1d) (which incidentally is the only
example of the entire suture of a Peroniceras
known to me);in the latter species U1/I saddle
is trifid .
"Peroniceras " leei Reeside (1927, p.40,
pl.43, fig.3) has a suture with U2 and U1 sepa­
rated by a tall bushy U2/Ul. The Gloriaceras al·
so resembles Pro texan ites Matsumoto (1955),
as exemplified by "Mortoniceras bourgeoisi d'
Orbigny" in de Grossouvre (1894, pl. 14 , figs.
2a, 2c), from which it differs by the early deve­
lopment of the keel , the more flexuous and nu­
merous initial ribs, the slenderer but more dis·
sected elements of the external suture and by
the better defined and raised umbilical saddles.
Of the non-Colombian species known to
me congeneric fomlS would be represented by
some of the specimens described by Burckhardt
(1919, p. 109) as "Peroniceras cfr. subtricarina­
tum Sturm, non D'Orb, et auct.", especially the
specimen showing the last· suture and the living
chamber (Burckhardt, 1921, pl. 27, fig. 5); the
:Mexican specimen seems more evolute than the
Colombian one and its E/L saddle is also m<�r�
97
ramose and rectangular.
Incidentally, it should be observed that
the suture line of a fragmentary specimen figu­
red and described by Burckhardt (op. cit., 1921,
p.llO, pl.26J fig.2) although "true" Peroniceras­
_
like in its u2/U3 saddle depending on L/U2
does not belong to the large specimen of that
illusqation nor to the specimen represented by
his plate 27, fig. 5 where not obliquity is shown
• by U2 and U2/U3 saddle is independentof i..JU 2.
On the Dlrckhardt.'s specimen here considered as
congeneric with Gloriaceras the lateral carina
are also wavy (see op. cit., pl. 27, figs. 9, 1 2 ; pl.
27, fig. 5 ) . For coments on Stunn's specimen
see below under G. paulinae.
This genus is named in honor oftJie late
biologist Miss Gloria Correa.
Gloriaceras corr_eai n. sl?_: --.. _ .,... . ,_ ..,..,,
""P1. t3, fig. 18; Pl. n. 1tg .'10; 'fext�1igs.
_
9 C , 9D, 9H.
1954 Peroniceras moureti De Grossouvre, Burgi,
p. 33 [ex.spec. HB-2 13. S.G.N.M.P.].
1954 Ba rroisiceras (Herleites) sp., Burgi, p. 33.
[ex. spec. juv. HB-211/8. S.G.N.M.P.].
1957 Peroniceras moureti De Grossouvre, BUrgi,
p . 1 3 8.
1957 Barroisiceras (Herleites) spec., Biirgl, p.138
1964 Peroniceras moure ti De Grossouvre, Etayo­
Sema, p. 126, ex Btirgl 1954 .
1964 Barroisiceras (Herleites) sp;. Etayo-Sema,
p. 128, ex BUrgi 1954.
1968 Peroniceras aff. P. Czomigi Grossouvre,
Etayo-Serna, p. 45, fig. 13, Tab. t.
Holotype. NRC.Ca-1-5 (U.N.C.M.H.N.C.P.Ht.N.
83).
A completely septate specimen is available.
The i!'nermost whorls are smooth, the earliest
visible ornament consists of low rounded ribs
which begin at the umbilical rim, on the adum­
bilical third of the flank they describe a weak
concavity and then a feeble arc up to the invo­
lution line, from there the ribs extend onto ven­
ter with an adoral concavity; the intercostal
space is nearly as wide as the ribs. Sporadically
a rib may bifurcate adumbilically.
On the adapical part" of the last whorl the
section is rectangular with moderately high
abrupt umbilical wall, slightly overhanging; it
rounds through the beveled periumbilical zone
into the flat, subparallel inner half of the flanks
and then merges into the acutely beveled adven­
tral fourth ; the venter is tricarinate; at this point
the ribs are subradial and weakly biconcave,
their adventral concavity accentuated by their
acute terminal connection to the lateral carina;
the ribs are widened on the adventral half of
98 F.
the flank and have a periventral ovate clavus
from which they extend onto the venter.
The median ventral carina is high, slightly
Wldulated and separated from the ventrolateral
carinae by shallow , asymmetrical sulci. Adaper­
turally the whorl section becomes lower; the ribs
may bifurcate {rom the umbilical rim or near
the middle of the flnak.
On the adoral part of the last whorl the
section becomes subquadrangular, the ribs are
club-like on the flanks and at the connection
with the ventrolateral keels, clavi-like undula·
tions of the carinae develop as if the latter were
tuberculated; the ventral margin clavi become
strenghtened and periumbilically the ribs tend
to develop clavate swellings.
DIAGNOSIS. This species is characterized
by the slow ontogenetic cl)ange of its ribbing
from fme and biconcave in the early whorls to
strong, straight, club-shaped ribs in the later
stages; likewise the change in whorl section is
slow.
Measurements D Euw H W D/Euw D/H No.R.
Ofo %
NRC· Ca· 1·6 66 27 23 20 41 36 14
COMMENTS. This species resembles that
described by Burckhardt (1919, p. 109; 1921,
pl. 27, especially on fig. 3) as "Peroniceras cfr.
subtricarinatum Sturm, non D'Orb. et auct.",
but the nbs are finer and sharper on the latter.
The Mexican species also has more closely spa·
(!ed ribs.
G. c orreai n. sp. differs from G.paulinae
n. sp. because on the latter the ribs change
through the ontogeny from biconcave to stron·
gly concave ad orally. Earlier (Etayo • Serna,
1968) I iden tified G. correai n. sp. as "Peronice­
ras cf. Czomigi", having compared it to figure
2a of plate 11 of Grossouvre's paper; in the
French species, however, the illustrations indi·
(!ate that the lateral keels are continuous, not
�onnecred to the marginal tubercles.
Furthermore, the ribs are concave ado·
rally on the adumbilical third of the flank up to
a size where they are completely strai�t on the
Colombian species which likewise has umbilical
bullae that are lacking in the Grossouvre's (op.
cit., p. 103) illustration.
OCCURRENCE. Cucaita Member. Lower
Coniacian.
Gloriaceras rgzulinae.!bJJ!...
.__,._,�Pl. U,'Iig.·t ·;Text-figS': 9K, 9L.
fiolotype. NRC-Cu5 (U.N.C.M.H.N.C.P.Pt.N.84)
Paratype. (U.N.C.M.H.N.C.P.Pt.N. 73).
There is one internal mold available. The
second (?) whorl is subeUiptical in section,sligh­
tly higher than wide, the maximum width is at
the umbilical rim, the flanks gently converge to­
ward the arched venter; on the third whorl the
section has doubled .its height: the periventral
ETA YO · SERNA
region Is broadly rounded and there is a keel
along the median ventral line paralleled by
weak grooves.
On the fourth whorl the section retains
the preceding proportions but the flanks are
slightly more flattened and two lateral carinae
paralleling the median keel are visible on the
adoral part of the third whorl; the flanks are
smooth but at the adapical part of the fourth
whorl, prorsiradiate, weakly biconcave ribs be­
gin to develop from the umbilical rim and ex­
tend onto the venter ending at the ventrolateral
carinae, at this stage the periventral clavi are on·
Iy weakly indicated; soon thereafter the ribs be­
gin to show clear periventral clavi and more or
less contemporaneously bullate swellings deve·
lop on the ribs along the umbilical margin; on
the flanks of the fourth whorl the ribs describe
a well defined adoral concavity . The intercostal
spaces are slightly wider than the ribs on the in·
ternal mold.
B) Up to 50 mm D fVL-6-Cu-1(1966)].
The conch is septate throughout, the ornament
of the penultimate whorl is the same as on the
small specimen although the concavity of the
ribs on the flanks is less obvious because the
test is still present; on the adapical third of the
last whorl the concavity of the ribs on the
flanks becomes conspicuous, the ventrolateral
clavi are depressed and outwardly inclined.
The living chamber occupies two thirds of
the last whorl and on it the ribs are strongly
concave and very prominent. On this specimen
all three ventral carinae are only slightly wavy .
DIAGNOSIS. This species is characterized
by the quick ontogenetic change of the ribs from
sickle shaped in the early wh orls to strongly
concave adorally on the later whorls; the whorl
section changes more rapidly in height than in
wid th.
Measurements. D Euw H w D/Euw D/H No.R.
% %
NRC· Cu • 6 38 16 13 1 0 39 3 4 21(1/'Z)
VLG • CU· 1 52' 20 1 9 1 5 38 3 6 33
(1966)
N OTE : Specimen VL6-CU-1-(1965) was measu·
red to the end of the phragmocone only.
COMMENTS. Gloriaceras paulinae n.sp.
differs from the specimen figured by Burckhard t
(1921, plate 27, fig.5) which is characterized as
noted by Burckhardt (1919, p.ll O) : "Un ejem·
plar (I am XXVII fie.5) se aparta de los otros oor
. ,
sus costillas mas rObustas y mas distanciada5 ya
en Ia juventud, y parece representar una especia
distinta, como lo indica tambien la sutura que . . .
se acerca a un tipo de linea sutural caracterlstico
para el genero Mortoniceras . . . ", because of
.
its smaller size, lesser involution and weaker ribs.
The other specimensaescn"bed by Burck·
hardt (1919) and illustrated later (1921, pl. 26,
fig. 1-12, but especially figs. 4,2, 'th e latter ex·
 ZONATION OF THE CRETACE OUS OF CENTRAL C OLOMBIA
elusive of the suture line) might be conspecific
with the Colombian species.
The specimen described and illustrated by
Stunn (1901, p. 60, pl. 3, fig. 5, 5a) as "Peroni­
ceras subtricarinatum d'Orb. ·5pec." to which
Burekhatdt compared his specimens (1919, p.
109) shows at the stage of growth illustrated
by Sturm (D:39mm approx.) a similar orna­
ment to that of the adult of Gloriaceras pauli­
nae n.sp. however, from Stunn's comments (op.
cit. p.60), "Den Beschreibungen durch frii here
Autoren ist nur hinzuzufligen, dass bei zuneh·
mender Gro55e der lndividuen die drei Riicken·
kiele immer mehr schwinden, so dass schliesslich
der- Durehschnitt des letzten Umganges spitz-:.
oval wird", that is to say that on the European
species the ventral carinae attenuate and the
whorl section becomes acute-oval with growth;
this signifies that the ontogenetic development
df Sunn's "species" is opposite to that of the
Colombian species.
OCCURRENCE. Cucaita Member. Lower
Coniacian. /"' ,7-:- , /l . .,· I jq v
- r-o .rv<'>H·&... o.,. tozr...,,, ,j{f , ··· · · !".-
. Genus J!u�ocerafJ.Basse 1947.
'>IW� oll!l@'.eg!!!i
_-u.,. !li ·�'-.""'r ·- "" .. ., · -
Zumpangoceras Basse , 194 7, p. 144.
TYPE SPECIES. Zumpangoceras burckhardti
Basse , original designation.
Lectotype of Z. burckhardti
the specimen described by
Burckhardt (1919, p.98 and
later illustrated as Barroisice­
ras sp. ind.num. 7 by the same
author ( 1921, pl . 23 fig. 1),
here designated.
COMMENTS. The "subgenus" Zumpan­
goceras was pro�osed by Basse (1948) in her
study of the European and Madagascan species
of Barroisiceras. By studying my material from
Colombia and through the perusal afBurckhardt's
paper, im seems clear that although all of Burck·
hatdt's "Barroisiceras , . � share some common
characteristics, not all of them share the same
ontogenetic development.
The ribs are dominantly biconcave (:<11e·
xuous in Burckhardt's tenninofogy), or if straigh­
tened the superposed growth striae are flexuous;
in all cases on which the ventral region is obser·
vable the ribs continue across the venter (see
Burckhardt, 1921, pl. 25, fig. 9) on the early
ontogenetic stages, in a subdued manner in the
later stages (see op. cit., pl. 23, fig. 1) or some·
times indicated by stressed growth striae; this
peculiarity accotding to Basse (1948, p. 144) is
not present on other "Barroisiceras ...
In Burckhardt's material there are at least
two well defined groups of specimens with diffe·
rent ontogenetic development of the ornament:
1) specimens without lateral tubercles at any
�tage of development (small and large specimens
tn Burckhardt's sample), 2) specimens with Ia·
terai tubercles at least at some stage of develo!J·
lllent. In the latter group a second division is
BY AMMONITES 99
observable and was previously noted by Burek­
hardt in the description of his different species:
those specimens on which the lateral tubercles
quickly disappear without having ever been pro·
minent, and those specimens on which the late·
ral tubercles notably increase their strength with
growth.
It is for the latter group that the genus .
Zumpangoceras is here restricted, and i� is in
accotdance with Basse's criteri on to elevate her
subgenus to generic rank . In her study , Basse
acknowledged the value of Reeside's ideas ·c:)n the
taxonomic importance of the lateral tubercles
in such group, �� �e stated ( 194.�� p.ll�):
"de tubercules med1o·lateraux ne sont pas ·stric-
tement homologues. . . "� ?
I feel that the absence of the lateral tu­
bercle, unless due to sexual dimorphism, is to
be considered taxonomically significant, conse·
quen tly, I propose to restrict Zumpangoceras to
the Mexican and Colombian species of ammoni-·
tes with "Forresteria-like" appearance, with fie-
�uous ri�bing or periodically emp�asi�ed growth
hrae, wh1ch cross venter With a linguiform pro·
jection; . a median ventral clavus whi�h alter-
nates w1th ventrolateral clavate to obhque tu·
bercles; the lateral tubercles become stronger
with growth , they tend to migrate toward the
center of the flank from a more internal posi·
tion and dominate over the periumbilical tuber­
cles ; the early whorls with tectiform ventral re·
gion which subsequently becomes truncated or
even weakly excavated with growth.
The specimen figured by Burckhardt on
his plate 22, fig. 16, is here interpreted as cons­
pecific with the lectotype of Z. burckhardti;
the specimens described as Barroisiceras sp. ind.
num. 7 (op.cit.,pl. 25, figs. 16, 17) are here in·
terpreted as probably congeneric but not cons·
pecific with the lectotype.
Zum a �inai n. sp ""' ·� · ='" ·---�·--..
1gT; ext-ffgs. 9o, 9P:.
l �
Prr, ��
1968 l''orresteria (Forresteria) cf. F. allaudi
(BLT). Etayo-Serna, p.45, fig. 13, Tab. 1.
1972 Forresteria (Forresteria) cf. allaudi (BLT),
Etayo-Serna, p. 48.
Holotype. NCR-1 (1965) (U.N.C.M.H.N.C.P.Ht.
N. 85).
Paratype. (U.N. C.M.H.N.C.P.Pt.N. 74).
A) Up to 30 mm D (VL-6-Cu5). At the
adapical fourth of the penultimate whorl the
section is fastigate, senate along the median line;
there are rounded periventral to clavate tuber·
cles, the sides are ornamented with flattened
flexuous ribs, they begin at the umbilical rim
and internally to the middle of the flank they
bifurcate; one intercalary rib appears near the
middle of the flank and between a pair of
bifurcating ribs. Midventral and periventral
tubercles alternate, the fanner being adoral to
the latter.
100 F. ETAYO - SERNA
On the adapical third of the last whorl
the ornament follows the pattern previously
described, the ribs are accentuated and have the
shape of an elongated S, they extend on venter
and interconnect at the median ventral clavi.
The whorl section becomes subrectangu­
lar; the umbilical wall is low and vertical, it
rounds abruptly intO the flanks which are con­
vex on their adumbilical third, then they weakly
converge toward venter forming a rounded ven­
tral shoulder; the venter is raised· along the me­
dian line but it becomes slightly excavated on
each side. At this stage some ribs have weak pe­
riumbilical bullate swellings.
On the adoral half of the last whorl the
section becomes more inflated on its adumbi­
lical third and the venter becomes wide, almost
truncate ; at the boundary between adumbilical
and central thirds of the flank some ribs de\lelop
a subtriangular prominent tubercle, at the same
time the periven tral tubercles enlarge outwards
but they are clearly compressed_ �d clavate.
B) Up to 50 mm D (NRC-1 (1965)). This
specimen includes the adult (?) liVing chamber.
It is involute, the intercostal whorl section is
suboval: the umbilical wall is short, subvertical,
it merges rather abruptly into the convex flanks
that in tum round bto the broadly arched
venter.
The ornament consists of ribs most of
which start at the umbilical brink in a claw-li­
ke manner,· they begin as sharp points and
thicken outwardly, they also bend backwards
slightly on the adumbilical third of the height
of the whorl; just internally to the middle of
the flank the ribs develop a prominent pyrami­
dal tubercle and from it the ribs run forward
ending .periventrall y in a thorny, clavate and
outwardly directed tubercle; the ribs may bi­
furcate from the lateral tubercle with one of
the branches joining either the following adoral
or the preceding adapical periventral tubercle
(which th(ln appears to join two ribs coming
tonn different nodes) in such a manner that the
ribs seem _ to form a W on the flank; this does
not alwayspccur on the opposite side, however.
Some ribs may have a slight periumbilical
node or they may bifurcate periumbilically
with one branch joining the subsequent subme­
dial tubercle. The ribs are broad, low, and tend
to split lQngitudinally on the adventral half of
the flank: The intercostal spaces are broad, one
and a half times as wide as the ribs; shallow.
Along the median ventral line there is a
row of prominent clavi, adoral to the p�riven­
tral tubercles at the vertex of the linguiform
projection of raised lirae (rib lets?) on venter.
The growth striae are impressed on the
ribs on the internal mold; on venter the ribs
themselves are absent but the raised growth
striae cross it in the wide adorally linguiform
arc; the preserved fragment represents the living
chamber, there are 10 periventral clavi and 8
subcentral lateral tubercles.
DIAGNOSIS. This species is characterized
by: A) its great involution, B) the ontogenetic
development of its ornament: the siphonal and
penventral tubercles appear first followed by a
weak periumbilical and finally, a sub-mediolate­
ral tubercle, the latter at first are nearer the wn­
bilical third of the flank and then slowly migra­
te upwards toward the middle of the flank but
never attaining an exactly medial position but
becoming stronger with growth, C) the change
in the shape of the venter from fastigate to al­
most truncate and wide.
MeastUem eata. D Euw H W D/Euw D/H
% %
VL6 • Cu5 30 5 16 12 17 63
NRC-1 (1966) 60 10? 22 20 20 44
COMMENTS. The small juvenile specimen
resembles Forresteria s tantoni Reeside (1932,
pl. 7, figs. 1·7) which has a very strong ribbing
earlier in the ontogeny than the Colombian spe­
cies. Furthermore, its ribs are club-sh aped and
more numerous than in Z. osplnai n.sp., the late­
ral tubercle en Reeside's species is centrolateral to
adventral, contrary to that of the new Colom­
bian species where it is at first adumbilical in
position, later migrating toward center of the
flank.
The large Colombian specimen resembles
Forresteria (F.) allaudi (Boule, Lemoine and
Thevenine)[1907, p. 32, pl. 8, fig. 6 ; malfor­
med?l, however, the whorl section in the most
similar specimens is very depressed as with the
specimen illustrated by Boule, Lemoine and The­
venine, or the one by Basse (194 7, pl.8, figs.3,3a)
from the "Con_glomerat ferrugineux Ambohi­
marina, Mont Carre " ; however, on the Madagas­
can specimens the vertical distance between the
periventral and the lateral tubercles is shorter
than the distance be tween the latter and the
umbilical tubercles, the opposite of the Colom·
bian species.
Likewise on the Madagascan species the
ventrolateral tubercles are heavy and conical
whereas in the Colombian species they are com·
pressed, clavus-like, upwards and outwardly
directed. ForresteriD {orresteri Reeside (1932,
pl.5, fig. 5 ) has a subquadrangular whorl section
rather than higher than wide as on the Colom·
bian specimens; the siphonal clavi ofF. forresteri
Ft.eeside are elongated, and the ribs club-shaped
whereas on the Colombian species the ribs are
curved and h ave a tendency to split longitudinal·
ly.
Matsumoto (1969, p. 308) has discussed
the Indo-Pacific species, he considered Reeside's
species a synonym of the Madagascan species.
Whatever may be the case, it is clear that the
Colombian species differs in the ontogenetic
development of its nodes and pattern of rib­
bing and belong to a different group than the
Indo-Pacific species.
 Z O N ATION OF TH E CRETACEOUS OF CENTRAL C O L O MBIA
The specimen referred to the Madagascan
species by Benavides-Citceres ( 1956, p.478, pt
61, fig. 1) is too poorly preserved to warrant a
comparison. On the specimen illustrated by
Burckhard t ( 1921, pl. 25) in his figure 17, the
ribs are raised, ridge-like between the lateral
and umbilical tubercles, consequently, the late­
ral tubercles are not well individualized, they
are fine, subrounded in the Mexican species; the
opposite of these characteristics occur in the
Colombian species.
Incidentally, this figure 17 of Burckh ardt's
plate 25 ( 1921) seems to show the migration of
the tubercles toward middle of the flank.
OCCURRENCE. Cucaita Member. Lower
Coniacian.
,.....,.
.. #/
... ·.J.- .....�.-
This species is named in honor of the geo­
logist Carlos Ospina.
•
Holotype. N.R.C - Sora (U.N.C.M.H.N.C.P.Ht.
N. 68).
The innermost whorl observed (third?) is
twice as high as wide; the maximum width on
the adumbilical third of the flank; externally
from the maximum width the flanks are flat to
excavated and converge toward the venter that
has inclined sides. The next whorl covers two
thirds of the height of the preceding whorl , the
adumbilical third of the flank is clearly convex;
the remaining part of the flank is flat and weakly
converges toward the venter into which it mer­
ges across the shoulder.
The ornament consists of ribs which be­
gin at the umbilical rim with a rounded to bulla­
te tubercle, they trend radially to near the mid·
die of the whorl where they develop a weak
round node from which the ribs may bifur..:ate,
the adventral branch is slightly adorally arcuate
across the venter; at the ventrolateral margins
the ribs develop clavi that are subparallel to the
median ventral line; riblets interconnect the me­
dian ventral and the lateroventral tubercles.
Early in the ontogeny , the ribs are narro·
wer than the intercostal spaces, later they beco­
me broad , club-shaped, wider than the intercos·
tal spaces especially on the adventral half of the
flank. The increase in umbilical width with
growth is quite •significant.
D IAGNOSIS. This species is characterized
by: A) the decrease in the distance between the
ribs with growth, B) the ephemeral development
of the small, rounded lateral tubercles, C) the
stressing with growth of the periumbilical tuber·
cles and the increase of the umbilical width.
Measurements D Euw H W D{Euw D/H
Sora NRC 40 7 22 14 17% 1:1 5 %
NOTE : The measurements refer to the entire
part of the conch only.
BY AMMONITES 101
COMMENTS. The only species�·known to
me that is comparable is ''Barroislc.� aff. als·
tadenense Solger, non auct�', de�cribed by
Burckhardt (1919, p.105) and subsequently
illustrated (1921, pl.25, tig.10) but on the
Mexican species [=Zumpangoceras ] the ribs are
finer, more crowded and more prominent and
the periventral tubercles are very attenuated,
likewise its umbilicus does not seem to widen
• with growth.
OCCURRENCE. Cucaita Member. Lower
Conician.
Genus
TYPE SPECIES. Buenoceras loboi n. sp.
DIAGNOSIS. Conch discoidal, moderate·
ly involute, maximum inflation near middle of
the flank from which the flattened sides round
toward the umbilicus and the periventral mar­
gins; the venter is tectiform; the early whorls
are ornamented with fine ribs that " appear
smoothly at the umbilical margin; they are
weak on the flank but on the adventral third
they are stronger; there are ventrolateral
tubercles that are inclined toward the ventral
median line, the ribs then cross the venter in an
acute V with an acute clavus at the vertex; on
later whorls weak ribbon like biconcave riblets
arise in bundles from tenuous periumbilical
swelling; they may bifurcate near the middle of
the flank where they are a little wider but do
not develop a tubercle . In the adults all ribs end
in oblique claw-like tubercles which are connec·
ted through the prolongation of the ribs with
the ventral median clavi.
COMMENTS. This genus differs from
Zumpangoceras Basse ( 194 7), by the absence of
lateral tubercles which are stressed with age.
From "Schloenbachia " flexuosa Gerhardt
( 1898, p. 157, pl. 4, fig. 1a,1b,1c) from Upper
Coniacian beds of Colombia which has a whorl
section with the inner half of flanks weakly ex­
cavated to flat, whereas the moderately excava­
ted external halt" of the whorl converges toward
the venter; the inner whorls of "S. " {lexuosa
have venter fastigate, later in ontogeny the ven­
tral zone widens, the ventral median keel is de­
pressed and the venter becomes almost trunca•
ted ;on the internal whorls the lateral ornament
consists of low flexuous ribs, which develop
oblique ventrolateral claw-like tubercles and
continue to midventer; on subsequent stages,
the ornament is composed of bundles of fal­
coid to falcate raised lirae that define very low
ribs, they extend onto venter forming an acute
siphonal V, the ribs develop a row of ventrola­
teral tubercles subparallel to the ventral median
line.
These ventrolateral tubercles are .elevated
adapically and thin and project slightly oblique­
ly and adorally; on later stages the rib; and peri­
ventral tubercles are suppressed and only raised
lirae with an accentuated concavity on the ad·
102 F. ETA Y O · SERNA
ventral third of the whorl side are present (the
two later stages are illustrated on Gerhardt's
figure 1; plate 4, 1898); the sum of the prece­
dign characteristics differentiate clearly both
taxa.
This genus is named in honor of Professor
Jesus A. Bueno of the Department of Geocien·
cias U. Nal. of Colombia.
·-·-,� .• .liJ��rGergs I,Qboi n. m.
--· .
. PI.l ;fig. 2; Text-fig. 9R, 9U.
Holotype. NRC-4 ( 1965) (U. N. C. M. H. N. C.
P. Ht. N.87).
Paratype. (U. N. C. M.·
H. N. C. P. Pt. N. 75).
The adapical segment of the second whorl
h as a subcylindrical section; at an equivalent
position on the third whorl the section is twice
as high as .wide and ventral median clavi occur
on the adoral portion of this whorl; the flanks
have an even curvature from the ventral median
clavi to the umbilical rim.
The umbilical wall is curved and bends
toward the venter of the preceding whorl. On
the fourth whorl (D: 15 mm) the venter is tecti­
form and has diverging flat ventral sides which
round smoothly into flanks; the latter have
their maximum width adumbilically near the
middle.
Every whorl overlaps to a point slightly
adumbilical of the middle of the flank of the
preceding whorl.
·
The ornament on the last PJ't!&erved whorl
consists of ribs that arise from rounded swe­
llings at the umbilical rim, describe a weak
concavity on the adumbilical third of the flank,
then trend prorsiradiate to near the middle of
the flank, from which point they describe a wi·
de adoral concavity. The ribs are flat, ribbon­
like; at the periventral margins the ribs develop
an oblique claw-like subelliptical thickening,
from these nodes the riblets or Iirae connect
with the ventral median clavi in a manner such
that venfA!r is crossed by an array of acute Vs.
The suture line has L asymmetrical; the
sanddles are subrectangular, symmetrically to
subsymmetrically bifid; U2 is oblique.
The laceration of the different elements
of the suture line is very variable even on both
sides of the same suture.
- . ' �- . ;..·.. .
'
DIAGNOSis: This species is characterized
by: A) its biconcave to falcoid ribs that coales­
ce at low rounded tubercles on the umbilical
rim, some of them bifurcate near the middle
of the flank without forming a tubercle , B) the
claw-like periventral tubercles and elevated me­
dian ventral row of clavi.
Measurements D Euw H w
NRC-Sora ? ? 16 10
NRC·4 ? ? 30 18
COMMENTS. The only foreign species
·
known to me that compares with this species is
Reesidites minimus (Hayasaka and Fukada) ty­
pe species of Reesidites Wright and Matsumoto
(1954), however, the latter has prominent
strong ribbing very early in the ontogenetic de­
velopment (see Hayasaka and Fukada, 1951,
pl.l), the opposi� is true of the Colombian spe­
cies, on the latter the ventrolateral nodes are
oblique to the median ventral line but in Reesi­
dites minimus they are parallel to the siphonal
line (See Matsumoto, 1965, p 61, pl.64, fig.
34a; holotype, pl.14), finally (and not critical?)
the suture lines are differen t : in the Japanese
species E/L saddle is asymmetrically bifid with
the adventral branch conspicuously shorter
than the external branch (see Matsumoto,
p .64, fig. 34b, holotype).
More similar, though , and here interpre­
ted as congeneric, is · the Mexican species repre­
sented by "Barroisicera sp. ind.num.3", of
Burckhardt (1919, p.102) and subsequently
illustrated by him ( 1921, pl.24, fig. 10); from
the illustration, the Mexican species h as closer
periventral tubercles and a more accentuated
bend of the low ribs at middle of the flank than
on the Colombian species.
Biirgl, 1954a, recorded a "Barroisiceras
subtuberculatum (Gerhardt)" (ex.spec. HB-216
SGNMP), which seems to be a member of Bue­
noceras loboi n. sp., however the former seems
thicker and has stronger ribs than the latter.
OCCURRENCE. Cucaita Member. Lower
Conician.
"\""' ' \
' . c , ..
This species is named in honor of the
Engineer Ramiro Lobo of the Ministerio de Mi-
nas y Petroleos. ·
.._,
Genus.!Pt!lt . IlL
1.
ns � �ygnjtesh�atsumoto,
������
��.� '�-
Qlr �).
-��
-�·.
Protexanites Matsumoto, 1955, p.3.
TYPE SPECIES. ' 'Mortoniceras" bourgeoisi
(d'Orbigny) Grossouvre
(1894, p.73, J)l. 13, fig.2 ; pl.,
14, fig.2-5) [sic], ori�al desig-
nation.
·
fro{g�!'n.ti£st_,ucqiV,Je��-�
Pl.l , fi'gs. ,4; 'rexr.Ji �
1968 Protexariites caiiaense (Gerhardt), .Etayo­
Sema, p.45, fig.13, Tab. I.
1972 froiexanites canaense (Gerhardt), Etayo­
Serna, p. 48.
Holotype. VL6-Cu-3 (U. N. C. M. H. N. C. P.
N.88).
��
The first whorl of the conch has a vaulted
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY
section; it is slightly wider at the umbilical mar­
gin and has weakly convex or subparallel sides
' that evenly round into the broad fastigate ven­
ter; the umbilical wall is oblique and rounds
abruptly into the sides; the second whorl is sub­
q uadrate, the flanks are flat between the umbi­
lical and the ventral margins, -the venter is wea­
Idy keeled along the median line, arched from
the keel to the periven tral margin; the flanks
overlap only the external half of the preceding
whorl.
On the third whorl the venter tends to be­
come flat but keeled along the median line; up
to the second whorl the sides are smooth, but
on the third whorl broad, well separated ribs
begin to appear.
On the adoral half of the fourth whorl
there are eight low ribs most of them with bu­
llate periumbilical tubercles; the ribs extend
�;adially to the periventral margin where they
develop a long triangular spinose tubercle, upon
which abub; against the following whorl.
On the fifth whorl the section is quadra­
te, with a broadly rounded umbilical rim and
the innermost part of the umbilical wall weakly
converges toward the venter of the preceding
whorl; the ventral margins are broadly rounded
along the intercostal spaces, they raise into
rounded bulges from which they connect with
the ventral median keel.
There are sixteen ribs on the fifth whorl,
they are separated by intercostal spaces twice
as wide as the width of the ribs, and the peri­
umbilical bullae are conical; the ribs are wider
and weaker near the middle of the flank. On the
sixth and last whorl the section is as in the pre­
ceding whorl; the growth of the whorls is relati­
vely slight.
The ribs begin at brink of the umbilical
wall and are inclined adapically, they bend slight­
ly just outside the umbilical rim and then trend
radially to weakly arcuately across the flank,
they develop periventral tubercles that project
outwards and backwards like a thorn; from this
marginal tubercle the rib continues onto the
venter as a broad triangular ridge at the end of
which one or two clavate, bead like tubercles
are pre,se nt; between these tubercles and the
raised median ventral kell there is a wide shallow
sulcus.
The growth striae begin a� the umbilical
se am, they slant down about 45 adapically on
the umbilical wall, they then tum forming a
shallow concavity on the adumbilical third of
the flank, describe a weak arc on the central
third and a concavity on the adventral third ;
they trend toward the ventral median line for­
ming a linguiform projection.
At some places on test, especially where
this is limonitized the growth striae can be seen
· to anastomose with each other and form a net­
work; at places where the test is preserved but
not limonitized the internal structure of the test
AMMONITES 103
appears to consist of rhombic compartments but
this is due In twinning of the calcite shell material; ' ·
DIAGNOSIS. This species is characterized
by: A) its quadrate whorl section, B) ib; radial
ribs with periventral spinose nodes and C) the
shallow and moderatelly narrow sulci . that para-
llel the ventral median keel. · :; > · ·
.
Me&S\lremeots D Euw H w D/Euw D/H No.R.
% %
VL-6-Cu-3 38 15 15 13 39 39 20
70 30 26 24 43 �-7 19
90 36 30 29 40 33 23
NOTE : The height of the whorl was measured ·
over the carina.
COMMENTS. Protexanites cucaitaense �p.
differs from "Mortoniceras bourgeoisi (D'Orbig­
ny)" in de Grossouvre [1894, p. 73, pl. 14, figs.
3a1 3b ) ,which has the ventrolateral clavi taller
th an the ventral median keel on the excavated
venter, contrary to the condition of the Colom­
bian species; further, on the French species the
umbilical tubercles are radially elongated but
in the Colombian species they are pustuliform;
in addition, the ribs tend to attenuate and widen
near the middle of the flank on the Colombian
species, a trait that is not observed on the French
species.
Pro texanites cucaitaense differs from Pro­
texanites cagnaense (Gerhardt) [usually mis!pe­
lled canaense] which has a high, rectangular
whorl section whereas the former has a quadra­
te whorl section and shallower ventral sulci
paralleling the ventral median carina; further­
more, on P. cucaitaense n.sp., there is a persistent
correspondence of two successive ventral clavi
to each marginal tubercle, a trait that is obser­
ved only late in the ontogeny of Gerhardt's
species (1898, pl. 1, fig. 2a, 2b).
Young (1963, p. 66-67) discussed the sta­
tus of Protexanites Matsumoto and Prionocy­
cloceras Spath and seemed to dismiss the valut.!
of the ventrolateral clavate tubercles of Prote­
xanites.
In my opinion, based on the Colombian
specimens available to me, it appears that the
ventrolateral tubercles greatly enlarge with
growth in Prionocycloceras, but on Protexani­
tes, the tubercles do not enlarge as much and
if they enlarge, they do not obliterate the ven­
tral clavi, or both weaken concomitantly (see
Grossouvre, 1894, pl. 13, fig. 2; D : 150mm).
As observed by Young, and confirmed
by the Colombian material, there are seconda­
ry ribs or riblets intercalated along the periven­
tral margins on Prionocycloceras, but in Prote­
xanites, the intercostal spaces are smooth, the
ribs are more distant and rectiradiate; in addi­
tion, the ventrolateral clavi seem to appear
earlier in the ontogeny of Protexanites than in
Prionocycloceras. Incidentally, the type loca­
lity of the holotype of the type species of Prio ­
nocycloceras is not near Chejende, Venezuela,
104 F. E T AYO · SERNA
but "Fundort: Von Stiibel im Bett des Rio Gua­
yabo an der Siidwestseite des Cerro Pelado im
siidlichen Columbien gefunden". (Gerhardt,l898
p. 197; see also Steinmann, 1882, p.169).
OCCURRENCE. Cucaita Member. Lower
Coniacian.
�.���TH:fH�A
Family
....
CYMATOCERATIDAE
Genus Heminautilus Spath, 1927
Heminau ti���ath, 1927, p. 22.
4 357 3 1 ) in whorl profile, although on the latter
TYPE SPECIES. Nautilus saxbii Morris, ori·
ginal designation.
Heminautilus etheringtoni
Durham.
Pl.4, fig. 8; Pl. 5, fig. 9.
1946 Heminautilus e theringtoni Durham, p.432
pl. 62, fig.4; pl. �. figs. 3,5-7, pl.64, fig.
1-3; Text-fig. 3c-f,5 .
1947 Heminautilus e theringtoni Durham, Dur­
h am, p. 576, figs. 1-3.
1956 Heminautilus etheringtoni Durham, Kum·
mel, p. 335, 436, 437, fig. 201.
1964 Heminautilus etheringtoni Durham, Kum­
mel, K407, fig. 29K.
1964 Heminautilus e theringtoni Durham, Eta­
yo-Serna, p. 129.
Hypotype. F- 15 (U.N.C.M.H.N.C.P.Hy.N.45). ·
, Heminautilus saxbii (Morris) the h olotype
Only a completely septate internal mold
is available. The conch is very involute, the whorl
section is high, inverted cordate; the umbilical
wall is inclined , it ascends smoothly to the high.
periumbilical area and then merges into the
slightly excavated convergent flanks that bend
abruptly at the periventral margin to form a
rounded rectangular shoulder; the venter is
truncated.
The suture line is adorally arcuate on the
umbilical wall, on the periumbilical area it bends
sharply and trends rursiradiate for a shorter dis·
tance than the internal branch , and then it chan­
ges progressively to form a wide, very regular
* *
adoral concavity on the flanks, it crosses the
venter straight, normal to the median ventral
line. In general the shape of the suture line is
that of a sickle with a short handle.
Measurements D Euw H W D/Euw D/H
% %
F·15 115 5 75 62 4 65
COMMENTS. The specimens studied by
Durham, excepting that coming from his loca·
lity C (Paratype, U.C.M.P.N. 35735) are slightly
more inflated than the present which, however,
agrees well with Durham's holotype (U.C.M.P.N.
the flanks are relatively straight whereas in my
specimen they are weakly contracted towards
the adven tral third of the flank.
On this specimen the suture lines cross
the venter normal to the median ventral line
without any indication of a siphonal sinus or
at most only a suggestion of it; in the collection
of the University of California there is a topoty­
pe, internal mold, of Heminautilus etheringloni
· Durham (U.C.M.P.N. loc. C-180) which does
not show the ventral sinus, a trait previously re·
corded by Durham ( 1946, p. 432) for one para­
type (U.C.M.P.N. 337 33); on this specimen the
suture line is weakly arched adorally and at one
point it is slightly oblique to the ventral median
line.
On the holotype (U.C.M.P.N. 35731) the
venter of the internal mold is flat, almost trun·
cated but where the test is preserved it shows a
feeble siphonal sulcus: a ventral sutural lobe is
present.
of the type species of the genus as illustrated by
Kummel (1956, pl. 10) shows an adventral con­
traction of the flanks and its section is less infla·
ted and the ventral sinus is well marked compa­
red to some equivalent specimens (immature?)
of H. e theringtoni.
The specimen illustrated by Douville {19 16,
pl. 17, fig. 2b; see fig. 6,4) as "Nau tilus lallie ri
d'Orbigny" has suture line trending straight or
with an adoral bow on the venter; the venter on
the latter species is flat to weakly excavated
with bicarinate margins and with maturity a cen·
tral carina begins to develop, but this trait is
not observed on Durham's species.
*
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
·
--� ·� ..
.. J_:··
._. ...,,
SUMM ARY
In two regions of the Eastern Cordillera
of Colombia, namely Villa de Leiva and Apulo,
the lithologic sequences were ascertained and
carefully mapped. The ammonites collected bed
by bed from the lithologic sequences were taxo.­
nomically investigated and this study constitu·.
tes the body of the present report.
102 species of ammonite.s are described
and illustrated, of these 8f11o of them are recog­
nized as new taxa; 52 g_eneral of ammonites are
described of which 21 u;o of the total is reported
new as follows: Colomboceratoides n. sg. [type
species Toxoceratoides (Colomboceratoides)
renzoni n. sp.J; Pseudoptychoceras n.g. (type
species Pseudoptychoceras gilberti n. sp. ]; Car­
loscaceresiceras n.g. [type species Car/oscacere­
siceras caceresi n. sp. ] ; Corteziceras n. g. (type
species Corteziceras cortezi n. sp. ]; Jaumaportai­
ceras n. sg. [ type species Trochle!Ceras (Jauma­
portaiceras) ho{{stetteri n. sp.]; Zambranoites n.
sg. [type species Vectisites (Zambranoites)
zamoranoi n. sp.]; Juandurhamiceras n.g. (type
species Juandurhamiceras juandurhami n. sp.];
Stoyanowiceras n.g. (type species A mmonites
tre{fryanw; Karsten, 1858]; Riedelites n.g.
f type species Riedelites esthersemae n. sp. j;
'Rinconiceras n.g. f type species Rinconiceras
rinconi n. sp. ]; Ralphimlayites n.g. [type species
Acanthoceras prorsocurvatum Gerhardt, 1898];
Codazziceras n. g. [type species Lyelliceras
scheibei Riedel, 1938]; Franciscoites n. g. [type
species A mmonites toroanus Karsten, 1858];
Gloriaceras ng. [type species Gloriaceras correai
n. sp.l; Buenoceras n. g. [type species Buenoce­
ros loooi n. sp. ].
The new species described are the follo­
wing: Holcophylloceras hemandezi n. sp., Hy­
pophylloceras wiedmanni n. sp ., Eogaudryceras
helmsi, n. sp., Eogaudryceras (Eote tragonites)
cabaricoi n. sp., Toxoceratoides (Colombocera­
toides) renzoni n. sp., Pseudoptychoceras
gilberti n. sp., Aconeceras gutierrezpalmae n. sp.,
Melchiorites palmeri n. sp., Valdedorsella (?)
rolombiam n.sp., Pseudohaploceras Ieiuaeme n.sp.
Carloscaceresiceras caceresi n. sp., Carloscacere-
siceras monteroi n. sp., Corteziceras cortezi n.
sp., Pse"(ldosayne/la ralphimlayi n. sp., Lewesi­
ceras (?) ulloai n. sp., Trochleiceras (Trochleice-.
ras) juliuerti n. sp., Trochleiceras (Jaumaportm­
ceras) ho{fstetteri n. sp., Cheloniceras (Epiche·
loniceras) dognaeluirae n. sp., Cheloniceras
(Epich eloniceras) camachoi n. sp., Cheloniceras
(Epicheloniceras) carlosacostai n. sp., Cheloni­
ceras (Epicheloniceras) jimenoi n. sp., Cheloni·
ceras (Epicheloniceras) ban:eroi n. sp., Cheloni­
ceras (Epich eloniceras) pardoi n. sp., Cheloni:
ceras (Epicheloniceras) leonhardtriedeli n. sp.,
Cheloniceras (Epicheloniceras) huertasi n. sp.,
Vectlsites (Zambranoites) zambranoi n. sp.,
Vectisites (Zambranoites) cruzi n. sp., Vectisi­
tes (Zambranoites) mateusi n. sp., Vectisites
(Zambranoites) cadenai n. sp., Vectisites
(Zambranoites) duquesi n. sp., Hypacanthopli­
�es (?) dognaliciae n.sp., Jua�durhaf1!icer!V!
Juandurhami n. sp., Juandurham1ceras gzrald01
105
· : ".i;..'� ,
sp., A canthohoplites bigoureti(orin.e n. sp.,
n.
Acanthohoplites seunesiforme n. sp., Acantho­
hoplites quitasuegno n. sp., Acanthohoplites
odiosus n. sp., Acanthohoplites pluricostatum
n. sp., Acanthohoplites elegan tean te n. sp.,
Acanthohoplites luisperezi n. sp., Acanthoho­
plites (?) lep tocerati{orme n.sp. , Acarrtf!.fiOplites
(?) serpenti{orme n. sp., Colombiceras {?) fore­
roi n. sp., Colombiceras (?) sarmientoi n. sp.,
• Gargasiceras (?) juanwyatti n. sp., Douvilleiceras
tarapacaense n. sp., Douvilleiceras abozagloi n.
sp., Eodouvilleiceras pedrocarvajali n. sp., Eodo­
uvilleiceras (?) tequendamai n. sp., Du{renoyia
boteroi n. sp., Du{renoyia hansbuergli n. sp.,
Neodeshayesites albertoalvarezi n. sp., Neodes­
hayesites ciPgulatum n. sp., Stoyanowiceras
cin tatum n.. sp. , Riedelites esthersernae n. sp.,
Riedelites quebradanegra n. sp., Riedelites
{ilosum n. sp., . Parahoplites hubachi n. sp.,
Parahoplites triston n. sp., Engonoceras olgalu­
ciae n. sp., Engonoceras (?) duartei n. sp.,
Platiknemiceras colombiana n. sp., Knemiceras
pegnai n. sp., Rinconiceras rinconi n. sp.,
Lyelliceras carvaja/orum n. sp., Lyelliceras
pseudolyelliforme n.sp., Lyelliceras isaacleaz· n.
sp., Ralphimlayites apuloense n. sp., Tegoceras
benauidescaceresi n. sp., Codazziceras {ina n.
sp., Mammites nodosoidesappelatus n. sp.,
Mammites {ugax n. sp., Mamites scutulatus n.
sp., Franciscoites suarezi n. sp., Im/ay{ceras (?)
ralphimlayi n. sp. Fagesia zanellai n.>sp., Para­
mammites colombianus n . sp., Hoplitoides
hernanmojicae n. sp., Hoplitoides lacabagnae
n.sp., Hoplitoides lagiraldae n. sp., Niceforoce­
ras boyacaense n. sp., Peroniceras robertogarciae
Peroniceras diabloi n. sp., Peroniceras guerrai n.
sp., Prionocycloceras portarum n.sp., Prionocy­
cloceras (?) cucaitaense n. sp., Gloriaceras co­
rreai n. sp., Gloriaceras paulinae n. sp., Zum­
pangoceras ospinae n. sp., Zumpangoceras (?)
sorae n. sp., Buenoceras loboi n. sp., Protexani·
tes cucaitaense n. sp.
These different species of ammonites repre­
sent a biostratigraphic sequence ranging in age
from Upper Aptian to Lower Santonian, based on
them, tb> following zonation ("Assemblage-zone"),
has been proposed : 1) Zone of Du{renoyia sanc­
torum and Stoyanowiceras tre{{ryanus, below;
2) Zone of Parahoplites (?) hubachi and Acan­
thohoplites (?) leptoceratiforme, above for the
Upper Aptian; 3) Zone of Neodeshayesites co­
lumbianus and Douuilleiceras solitae for the
Lower Albian; 4) Zone of Platiknemiceras co­
lombiana, Rinconiceras rinconi and Lyelliceras
pseudolyelli{orme for the Middle Albian; 5)
Zone of Mammites nodosoidesappelatus and
Franciscoites suarezi for the Lower Turonian;
6) Zones of Gioriaceras correai, Pro texanites cu·
caitaense and Codazziceras scheibei for the
Lower Coniacian; 7) Zone of Lenticeras baltai
and Paralenticeras sieuersi for the Lower Santo·
nian.
Assemblages representing the Upper Al­
bian, Cenomanian, Upper Turonian, and Upper
Coniacian were not recognized. Detailed descrip·
tions of some lithostratigraphic units are given,
ar;.d their faunal content indicated.
1 06 F. ETA YO · SERNA
RESUMEN
En dos regiones de Ia Cordillera Oriental
de Colombia, en Villa de Leiva y Apulo, se de­
terminaron las secuencias litol6gicas que fueron
cludadosamente cartografiadas. Las amonitas
colectadas estrato por estrato de tales secuen­
cias litologicas fueron taxon6micamente inves­
tigadas y este estudio constituye el objetivo
principal de este escrito.
Se describen e ilu!l;ran 102 especies de
amonitas, de las cuales ssu;a:: onfonnan nuevas ta­
xa; se d e �ben 52 generos de amonitas de las
cuales 2lu/odel total se dictaminan como nuevas
asi: Colomboceratoides n. sg. [especie tipo To­
xoceratoides (Colomboceratoides) renzoni n.
sp. ]; PseudoptychocerO$ n. g. [especie tipo Pseu­
i
f,
doptychocer:O$ gilberti n. sp. ]; Ca rloscaceres ce­
rO$ n . g. especie tipo Carloscaceresiceras cacere­
si n. sp. ; Corteziceras n. g. [especie tipo Corte­
ziceras cortezi n. sp. ]; Jaumaportaicer(l$ n. sg.,
respecie tipo Trochleiceras (JaumaportaicerO$)
hof{ste tteri n. �.J;Zambranoites n. sg. [especie
i
tipo Ve ctis tes (Zambranoites) za.mbranoi n. sp . ];
Juandurhamiceras n. g. [especie tipo Juandurh a­
miceras juandurhami n. sp . ] ; StoyanowicerO$ n.
g. respecie tipo A mmonites treffryanw Karsten,
18158 J; Riedelites n. g. [especie tipo Riedelites
esthersemae n. sp. J; Rin con iceras n. g. r especie
i
tipo R in conicerO$ rinconi n. sp. ]; Ralph m layi­
tes n. g. respecie tipoAcanthocerasprorsocurua·
1898];
tum· G!:!rfiardt, Codazziceras n . g. [ espe<:ie
ti L lli sheibeiRiedel, 1938J;Franciscoi­
po ye ceras
tes n. g. r especie tipo A mmonites toroanw
1S58
Kan;ten, ]; G loriaceras n. g. [especie tipo
Gloriaceras correai n. sp.l; Buenoceras n. g.
[ especie tipo Bueno cerO$ loboi n. sp. ]. .
Las nuevas especies descritas son las si­
guientes: Holcophylloceras hernandezi n. sp. _
Flypophylloceras wiedmanni n. sp., Eogandry­
r:eras helmsi n. sp., Eogaudryceras (Eo te tragoni­
tes) cabaricoi n. sp., Toxoceratoides (Colombo­
r:eratoides) renzoni n. sp. , Pseudoptychoceras
�ilberti n. sp., Aconeceras gu tierrezpalmae n. sp.
lf.elchiorites palmeri n . sp., Valdedorsella (?) co­
�mblana n.sp., Pseudohaploceras leivaense n.sp .
�arloscaceres1ceras caceresi n. sp., Carloscace­
·esiceras monteroi n. sp., Corteziceras cortezi
1. sp., Pseudosaynella ralphimlayi n. sp., Lewe­
:iceras (?) ulloai n. sp., Trochleiceras (Trochlei­
:eras} juliverti n .sp., Trochleiceras (Jaumaportai­
:eras) hoffste tteri n. sp., Cheloniceras (Epiche­
oniceras) camachoi n. sp., Cheloniceras (Epiche·
oniceras) carlosaf;ostai n. sp., Cheloniceras
Epicheloniceras) jimenoi il. sp., Cheloniceras
'Epicheloniceras} barreroi n. sp., Cheloniceras
'Epicheloniceras ) pardoi n. sp . , Cheloniceras
'Epicheloniceras) leonhardt riedeli n J>p.Chelo i­ n
:eras (Epicheloniceras} huertasi n. sp., Fe ctisi­
·es (Zambranoites) zambranoi n. sp . , Ve,·tisites
'Zambranoites) cruzi n. sp. , \Tectisites (Za m b ra­
lOites} mateusi n. sp . , Vectisites (Zambranoi­
·es} cadenai n. sp . , Vectisites (Zaml•ranoites}
luquesi n. sp., Hypacanthoplites ('t) dogna­
icine n . sp. , Juandu rhamiceras juandurhami n.
:p ., Juancfurhamiceras giraldo i n. sp., Acantho­
lqplites . bigoure ti{orme n . sp., A canthoho­
,z,tes od1osus n . sp., Acan thohoplites p/uricos·
tatum n. sp., Acanthohoplites elegan tean te n .
sp., A canthohoplites luisperezi n . sp., A cantho­
hoplites (?) leptoceratiforme n. sp., Acantho­
hoplites (?) serpen ti{orme n. sp., Colombiceras
(?) foreroi n . sp., Co/ombiceras (?) sarmientoi
n . sp., Gargasiceras (? ) juanwyatti n. sp., Dou vi­
lleiceras tarapacaense n. sp., Douuilleiceras abo·
zagloi n. sp., Eodouvilleiceras pedrocarvajali n .
sp., Eodouvilleiceras (?) tequendamai n . sp . ,
Dufrenoyia bo teroi n. sp., Dufrenoyia hans·
buergli n. sp . , Neodeshayesites a/bertoaluarezi
n. sp ., Neodeshayesites cingulatum n . sp .,
Stoyanowiceras cin tatum n. sp., Riedelites
que bradanegra n. sp., Riedelites filosum n. sp.,
Parahoplites hubachi n. sp., Parahoplites tristan
n. sp., Engonoceras olgaluciae n. sp., Engonoce­
ras (?) duartei n. sp., Platiknemiceras c o lom­
biana n. sp., Knemiceras pegnai n. sp. , Rinconi­
ceras rinconi n. sp., Lyelliceras caruajalorum n.
sp., Lyelliceras pseudolye/liforme n. sp . , Lyelli­
ceras isaacleai n . sp., Ralphimlayites apuloense
n. sp., Tego ceras benauidescaceresi n. sp . ,
Codazziceras {ina n. sp . , Mammites nodosoide­
sappelatus n. sp ., Mammites fugax n. sp., Mu m­
mites scu tulatus n.sp., Franciscoites suarezi n.
sp. , Imlayiceras (?) �alph imlayi n. sp., Fagesia
zanellar' n. sp., Para mammites colombianus n .
sp., Hoplitoides hernanmojicae n . sp . , Hoplitoi­
des lacabagnae n . sp . , Hoplitoides lagiraldae n.
sp., Niceforoceras boyacaense n . sp . , Pe roniceras
robertogarciae n. sp., Pe roniceras · diabloi n. sp .,'
Peroniceras _guerrai n. sp., Prionocycloceras
portarum n .sp., Prionocycloceras (?) cucaitaense
n. sp., Gloriaceras paulinae n. sp., Zumpangoce­
ras ospinae n.sp., Zumpangoceras (?) sorae n .
sp., Buenoceras lobo'i n .sp., Protexanites cucai­
taense n. sp.
Estas diferentes e species de amoni tas re­
presentan una secuencia bioestratigratica que se
extiende en edad desde al Aptiano Superior,
hasta el Santoniano Inferior, dentro de la cual
el autor propone Ia siguiente zonacion (''Assem­
blage-zones") : 1) Zona de Dufrenoyia sanc torum
y Stoyanowiceras treffiJ'anus, subyacente ; 2)
Zona de Parahop/itcs (?) hu bachi y A canthoho­
plites (?) lep tocera tiforme, superyacente : Ap­
ti an o Superior; 3) Zona de Neodeshayesites co­
lombianus y Do uvilleice ras solitae p ara el Albia­
no Inferior; 4} Zona de Platiknemiceras colom­
biana, Rinconir:P.ras rinconi y Lyelliceras pseu­
dolyelliforme para el Albiano Medio; 5) Zona
de Mammites nodosoidesappelatus y Francis­
coites suarezi para el Turoniano Inferior; 6) Zo­
na de Gloriaceras correai, Pro texanites cucai­
tae/lse y Codazziceras scheibei p ara el Coniacia-
. no Inferior; 7) Zona de Len ticeras baltai y Para­
lenticeras sieversi p ara el Santoniano Inferior.
No - fueron reconocidas las faunas repre·
sentativas del Al bi an o Superi o r, Cenom aniano,
Turoniano Superior y Coniaciano Superior.
Se dan descripciones detall adas de algunas
unid ades l ito estra tigrlificas y se indica Ia fauna
contenida en elias.
 ZONATION OF THE CRETACEOUS OF CEN TRAL COLOMBIA BY AMMONITES
RESUME
Les sequences lithologiques des regions de
Villa de Leiva et de Apulo, dans la Cordillere
Orientale de Colombie, cartographb!es en detail
ont ete detenninees. Les ammopites, recueillies
strate par strate ont ete clllll.Wes taxonom�
ment; cette etude constitue l'objectif du present
travail.
•
102 especes d'ammonites sont decrites et
illustrees, parmi lesquelles 85 o/o confonnent un
nouveau taxon; 52 genres d'ammonites sont
decrits panni lesquels les 21% suivants sont
considerees comme nouveaux: Colomboceratoi­
des n. sg. respece type Toxoceratoides (Colom­
boceratoicfes) renzoni n.sp.];Pseudoptychoceros
n. g. fespeee type Pseudoptychoceras gilberti n.
sp.j; Carloscaceresiceras n. g. [espece type Car­
loscaceresiceras caceresi n.sp. ]; Corteziceras n.g.
[qspeee type Corteziceras cortezi n.sp.J ; Jauma­
portaiceros n. sg. r espeee type Trochleiceras
(Jaumaportaiceros) hof{stetteri n.sp.l; Zambra­
noites n. sp. [espeee type Vectisites (Zambranoi­
tes) zambranoi n. sp.j; Juandurhamiceras n. g.
[espece type Juandurhamiceras juandurhami
n. sp.]; Stoyanowiceras n.g. respece type Ammo­
nites tref{ryanus Karsten 1S58]; Riedelites n.a.
respece type Riedelites esthersemae n. sp.1 ;
1linconiceros n.g. respece type Rinconiceras rin­
coni n.sp. ]; Ralp'himlayites n.g. [ espece type
Acanthoceras prorsocurvatum Gerhardt, 1898};
Codazziceras n.g. [espeee type Lyelliceras schez­
bei Riedel, 1938]; Franciscoites n. g. [espece ty­
pe Ammonites toroanus Karsten, 1858]; Gloria­
ceras n .g. [ espe ce type Gloriaceras correai n.sp. ];
Buenoceras n.g. [ espece' type Buenoceras loboi
n.sp . ].
Les nouvelles especes decrites sont les sui­
vantes: Holcophylloceras hemandezi n. sp., Hy­
pophylloceras wiedmanni n. sp., Eogaudryceras
helmsi n.sp., Eogaudryceras (Eotetragonites) ca­
baricoi n.sp., Toxoceratoides (Colomboceratoi­
des) renzoni n.sp., Pseudoptychoceras gilberti
n.sp., Aconeceras gutierrezpalmae n.sp., Melchi­
orites palmeri n.sp., Valdedorsella (?) colombia­
na n.sp., Pseudohaploceras leivaense n.sp., Car­
loscaceresiceros caceresi n.sp., Carloscaceresice­
ras monteroi n.sp., Corteziceras cortezi n.sp.,
Pseudosaynel/a ralphimlayi. n.sp., Lewesiceras (?)
ulloai n.sp., Trochleiceras (Trochleiceras) juli­
uerti n . sp., Trochleiceras (Jaumaportaiceras)
hof{stetteri n.sp., Cheloniceras (Epicheloriiceras)
camachoi n.sp., Cheloniceras (Epicheloniceras)
carlosacostai n.sp., Cheloniceras (Epichelonice­
ras) jimenoi n. sp. , Cheloniceras (Epichelonice­
ras) barreroi n. sp., Cheloniceras (Epichelonice­
ras) pardoi n. sp., Cheloniceras (Epicheloniceras)
le onhardtriedeli n.sp., Cheloniceras (Epicheloni­
ceras) huertasi n.sp., Vectisites (Zambranoites)
zam branoi n. sp., Vectisites (Zambranoites) cru­
zi n.sp., Vectisites (Zambranoites) mateusi n.sp.,
Vectisites (Zambranoites) cadenai n. sp., Vecti­
sites (Zambranoites) duquesi n. sp., Hypacan­
thoplites (?) dognaliciae n.sp., Juandurhamice­
ras juandurhami n.sp., Juandurhamiceras giral­
doi n.sp., Acanthohoplites bigoureti{orme n ..:p.,
107
Acanthohoplites teunuiforme n. sp . , Acantho­
hoplltu quitasuegno n. sp., ACtJnthohoplitu
odiosru n. up., Acanthohoplites pluricostatum
n. sp.,Acanthohoplites eleganteante n. sp.,ACtJn·
thohoplites luisperezi n. sp., Acanthoh¢plitu
(?) leptocerotlforme n. sp. , Acanthohoolites (?)
serpenti{orme n. ap., Colombiceras (?) foreroi
n. sp., Colombiceras (?) sarmientoi n. sp., Gar­
gasiceras (?) juanw.vatti n. sp ., Douvilleiceras
tDrapacaense n. ap., Douvilleiceras abozagloi n.
aD., Eodouvilleicel'dll pedroCtJrvajali n. sn., Eodo­
uuilleiceras (?) tequendamai n. sp., DutrenoyilJ
boteroi n. sp., DutrenoyilJ hansbuergli n. ap.,
Neodeshayesites albertoalvarezi n. sp., Neodu­
hayesites cingulatum n. ap., Stoyanowiceras
cintatu m n. so., Riedelites esthersemae n. sp.,
Riedelites quebrodaneea n. sp., [liedelites fila ­
sum n. sp., Parohoplites hubachi n. ap., Paraho­
plites tris tan n. s� . Engonoceros o[Haluciae n.
t
so., Engonoceras (? J duartei n. sp., Platiknemi­
ceros colombiana n. sp., Knemiceras pegru,i n.
sp., Rinconiceras rinconi n. sp., LyP-llicera8 car­
vaialorom n. sp., Lyelliceras pseudoljelli{orme
n. sp., Lyelliceras isCUJcleai n. sp., Ralphimlayites
apuloense n. ap., Tegoceras ·benavldescaceresi.
n. sp., Codazziceras {ina n. sp., Mammites nodo­
soidesappelatus n. sp., Mammites {ugax n. sp.,
Mammites scutulatus n.sp.,Franciscoites suarezi
n. sp. Imlayiceras (?) rolphimlayi n. sp., FagesilJ
zanet'fui n. sp.,Paramammites colombilJnus n. sp.,
Hoplitoides hemanmojicae n. sp., Hoplitoi­
des lacabagnae n. sp., Hoplitoides lagiraldae n.
ap., Nice{oroceras boyacaense n. sp.,Peroniceras
robertogarcwe n. sp., Peroniceras diabloi n. sp.,
Peroniceras guerrai n.sp.,friono.cycloceras por­
taro m n. sp., Prionocycloceras (?) cucaitaense
n. sp., Gloriaceras correai n. sp., Gloriaceras pau·
linae n. sp., Zumpangoceras ospinae n. sp., Zum­
pangoceras (?) sorae n. sp., Buenoceras loboi n.
sp., Protexanites cucaitaense n. sp.,
Ces especes differenms d'ammonites repre­
sentant une sequence bio stratigraphique dont
l'ige s'etend de 1'Aptien Superieur jusqti' au San­
tonien Inferleur. On pror.ose pour 12t intervafie
Ia zo'lation suivante : ('�mblage-zone;"), 1)
Zone de Du{renoyia sanctorom et Stoyanowice­
ras tref{ryanus sous Ia; 2) Zone de Parahoplites
(?) hubachi et Acanthohoplites (?) leptocerati­
forme au dessus de I' Aptien Superieur; 3) Zone
de Neodeshayesites columbianus et Douvilleice­
ros solitae pour 1' Albien Inferieur; 4) Zone de
Platiknemiceras colombiana, Rinconzceras rin­
coni et Lyelliceras pseudolyelli{orme pour 1' Al­
bien Moyen; 5) Zone de Mammites nodosoide­
sappelo.tus et Franciscoites suarezi pour le Tou·
ronien Inferieur; 6) Zone de Gloriaceras correai,
Protexanites cuca1taense et Codazziceras schei­
bei pour le Coniacien inferieur; 7) Zone de Len­
ticeras baltai et Paralenticeras sieversi pour le
Santonien lnferieur. Les Ammonites representa­
tives de l'Albien Superieur. du cenomanien , du
Turonien Superieur et du Coniacien Superieur
n'ontpas ete identifiees.Ce travail conUent flnale­
ment Ia description detaillee de quelques unites
litho logiques ainsi que Ia faune qu'elles con­
tiennent.
 108 F. ETAYO · SERNA
RIASSUNTO
In due regioni della Cordigliera Orientale
di Colombia, Villa de Leiva e Apulo si sono
stabilite alcune sequenze litologiche rilevate poi
con cura. Le ammoniti, raccolte strato a strato
secondo l'ordine della successione litologica,
sono state oggetto della ricerca tassonomica che
viene presentata in questo scritto.
Si descrivono e illustrano 102 specie di
ammoniti, fra le cuali 85 o/osi asse gnano a novi
taxai si descrivono 52 generi di ammoniti fra le
quali l'autore e del parere che 2lo/osiano nuovi:
J
Colomboceratoides n. sg. specie tipo Toxoce­
ratoides (Colombocerato es) renzoni n. sp. ];
·
Pseudoptychoceras n. g. [species tipo Pseudo­
ptychoceras gilberti n. sp.1; Carloscaceresiceras
n. g. f specie tipo Carloscaceresiceras caceresi n.
sp.j; torteziceras n. g. [specie tipo Corteziceras
cortezi n. sg. J; Jaumaportaiceras n. sg. rspecie
tipo Trochleiceras (Jaumaportaiceras) hoffste­
tteri n. sp.J; Zambranoites n. sg. [specie tipo Vec­
tisites (Zambranoites) zambranoi n. sp.l; Juan­
durhamiceras n. g. [species tipo lllllndurhamice­
ms.juandurhami n.sp.J ; Stoyanowiceras n.g. [spe·
cie tipo Ammonites tre{fryanus Karsten 1858];
Riedelites n. g. [specie tipo Riedelites estherser­
nae n. sp. ]; Rir,coniceras n. g. [specie tipo Rin­
coniceras rinconi n. sp. ]; Ralphimlayites n. g.
fspecie tipo Acanthoceras prorsocurvatum Ger­
hardt, 1898 ]; Codazziceras n. g. f specie tipo
Lyelliceras scheibei Riedel, 1938 j; F'ranciscoites
n. g. specie tipo A.mmonites toroanus Karsten,
J
1858 ; Glorillcera n. g. [specie tipo Gloriaceras
correai n. sp.]�Buenoceras n. g. [specie tipo Bue­
noceras loboi : sp. ].
.
. .
.
·
Le nu�ve specie. descritte sono Je seguenti:
Holcophylloceras hemandezi n. sp.Hypophyllo•
ceras wiedmanni n. sp., Eogaudryceras helmsi
n. sp., Eogaudryceras (Eo tetragonites) cabaricoi
n. sp., Toxoceratoides (Colomboceratoides) ren­
zoni n. sp., · Ps�udoptychoceras gilberti n. sp.
Aconeceras gut1errezpalmae n. sp. Melchiorites
palmeri n.sp., Valdedorsel/a (?) colombiana n.sp.
Pseudohaploceras leivaense n. sp., Carloscacere­
siceras caceresi n. sp., Carloscaceresiceras mon­
teroi n. sp., Corteziceras cortezi n.sp., Pseudo­
soynella ralphimlayi n.sp., Lewenceras (?) ulloai
n.sp., Trochleiceras (Tt'ochleiceras) juliverti n.sp.,
Trochleiceras (Jaumavortaiceras l hnff.�tP.tteri n .
sp., Cheloniceras (Epicheloniceras) dognaetVIrae
n. sp., Cheloniceras (Epicheloniceras) camachoi
n. sp. Cheloniceras (Epicheloniceras) carlosacos­
tai n .' sp., Cheloniceras (Epicheloniceras) jime­
noi n. sp., Cheloniceras (Epicheloniceros) ba"e­
roi n. sp., Cheloniceras (Epicheloniceras) pardoi
n. sp., Cheloniceras (Epicheloniceras) ieonhar­
dtriedeli n. sp., Cheloidceras (Epichelonb�ras)
huertasi n. sp., Vectj$ites (Zambranoites) zam­
branoi n. sp., Vec ti&jJ{Is (�ambranQ.ites) ,cruzi
iJiP-[
n. sp., Vectisites (Za. 4ttoite$) mateust n . sp.,
. .
Vectisites (Zambraii0MI1); cadenai n; sp., Vec­
tisites (Zambranoites) duquesi n. sp., Hypacan·
thoplites (?) dognaliciae n. sp., Juandurhcmice­
ras juandurhami n. sp., Juandurhamiceras giral­
doi n.sp., Acantholwplites bigourctiforme n. sp.;
Acanthohoplites seunesiforme n. sp., Acantho­
·
hoplites quitasuegno n. sp., Acanthohoplites
odiosus n. sp., Acan thohoplites pluricosta tum
n. sp.,Acan thohoplites eleganteante n. sp., A can­
thohoplites luisperezi n. sp., Acanthohoplites
(?) leptoceratiforme n. sp., Acan thohoplites (?)
serpenti{orme n. sp., Colombiceras (?) foreroi
n. sp., Colombiceras (?) sarmientoi n.sp., Garga­
siceras (?) juanwyatti n. sp., Douuilleiceras tara­
pacaerise n. sp., Douuilleiceras abozagloi n. sp.,
Eodouvilleiceras pedrocarvajali n. sp., Eodou­
villeiceras (?) tequendamai n. sp., Du{renoyia
boteroi n. sp., Du{renoyia hansbuergli n. sp.,
Neodeshayesites albertoalvarezi n. sp., Neodes·
hayesites cingulatum n. sp., Stoyanowiceras
cin tatum n. sp., Riedelites esthersemae n. sp.,
Riedelites quebradanegra n. sp., Riedelites {flo­
sum n. sp. Parahoplites hubachi n. sp., Paraho­
plites t,.iston n. sp., Engonoceras olgaluciae n. sp.,
Engonoceras (?} duartei n. sp., Platiknemiceras
colombiana n. sp., Knemiceras pegnai n. sp.,
Rinconiceras rinconi n. sp., Lyelliccras carvaja­
lorum n. sp.,Lyelliceras pseudolyelli{orme n.sp.,
Lyellieceras isaacleai n. sp., Ralphimlayites
apuloense n. sp., Tegoceras benavidescaceresf
n. sp., Codazziceras {ina n. sp., Mammites nodo­
soidesappelatus n . sp., Mammites {ugax n. sp. ,
Mammites scutulatus n . sp. Fhnciscoites suarezi
,
n . sp., Imlayiceras (?) ralphimlayi n. sp., Fagesia
zanellai n. sp., Paramammites colombianus n.sp.,
Hoplitoides hemanmojicae n. sp., Hoplitoides
lacabagnae n . sp., Hoplitoides lagiraldae n. sp.,
Nice{oroceras boyacaense n. sp., Peroniceras ro­
bertogarciae n. sp., Peroniceras diabloi n. sp.,
Peroniceras guerrai n. sp., Prionocycloceras por­
tarum n. sp., PrionocyclocerU$ (?) cucaitaense n.
sp., Glorillceras correai n. sp., Gloriaceras pauli­
nae n. sp., Zumpangoceras ospinae n. sp., Zum� , ·
pangoceras (?} sorae n. sp., Buenoceras loboi n.
sp., Protexanites cucaitaense n. sp.,
�ueste diverse specie di ammoniti rappre­
sentano una sequcnza biostratigrafica che si
stende in eta dall Aptiano Superiore fino al San·
toni?..no Inferiore dentro Ia quale l'autore pro·
pone la seguente zonificazione , \'Assemblage·
zone") : 1) Zona a DufrenOJiiz sanctorum y Sto­
yanowiceras treffryanus sottostante; 2) Zona a
Parahoplites- (?) hubachi y Acanthohoplites (?)
leptoceratiforme, soprastante (?) L'Aptiano Su­
periore ; 3) Zona a Neodeshayesites columbianus
y Douvilleiceras solitae per I' Albiano Inferiore;
4) Zona a Platiknemiceras colombitzna, Rinconi­
ceras rinconi y Lyelliceras pseudolyelli(orme per
l'Albiano Medio; 5) Zona a Mammites nodosoi­
desappelatus y Franciscoites suarezi per il Turo­
niano Inferiore; 6) Zona a Glorillceras correai,
Protexanites -cucaitanse y Codazziceras scheibei
per il Coniaciano Inferiore; 7) Zona a Lentice­
ras baltai y of?aralenticeras sieversi per il Santo.- '
niano· Inferio,:Q.. -Non si sono riconosciute le fau·.
·
ne reopresentiltive dell' Albiano Superiore, del
Cenoina:1iano, Turoniano Superiore e Coniacia·
no Superiom.
Si descrivono minuziosamente alcune unita
litostratigrafiche e si indica Ia fauna I( conteta.
 ZONA "nON OF THE CRETACEOUS OF
AQ
ZUS
· · ; .( : .
ENFASSUNG
In den Umgebungen von Villa de Leiva und
Apulo , zwei Gebieten auf der Ostlichen Kordille­
re Kolumbiens, wurden die lithologischen Abfol­
gen der Kre,�de aufgJ)nommen und sorgf3Itig Kar­
tographiert. Die aus jeder Schich_t aufgesammenl­
ten Anmoniten wurden taxonomisch untersucht.
Die Ergebnisse dieser Untersuchung stellen das
Hauptziel der vorliegenden Arbeit dar.
Es werden 102 Ammoniten-Arten beschrie_­
ben und abgebildet, aarunter 85 o/oDeueTaxa. Es
werden auch 52 Ammoniten-Gattungen besch­
rieben, unter denen sich 21 %neue Gattungen
finden : (Colomboceratoides) n. sg. [Typus Art
Tgxocerato itJes (Colomboceratoides) renzoni.
n� sp. ] ; Pseudoptychoceras n . sg. (Typus Art
Pseudoptychoceras gilberti n. sp· ]; Carloscace­
resiceras n. g. (Typus Art Carloscaceresiceras
caceresi n. sp.J; Cortezicer{ls n. g. (Typus Art
C.,orteziceras cortezi n. sp. ]; JauTTUJportaiceras
n. sg. fTypus Art Trochleiceras (Jaumaportai­
ceras) hof{stetteri n. sp . ] ; Zambranoites n.sg.
(Typus Art Vectisites (Zambranoites) zambra­
noi n. sp. ] ; Juandurhamiceras n. g. [Typus Art
Juandurhamiceras juandurhami n. sp.] ; Stpya­
nowiceras n. g. fTypus Art A mmonites treffrya­
nus Karsten 185 81; Riedelites n. g. (Typus Art
Riedelites esthersemae n. sp. ] ; Rmconiceras
n . g. [ Typus Art Rinconiceras ri11coni n. sp.];
Ralphimlayites n. g. {Typus Art Acanthoceras
prorsocurvatum Gerhardt 1898] ; Codazziceras
n. g. (Typus Art Lyelliceras scheibei Riedel,
1938 ] ; Franciscoites n. g. fTypus Art Ammo­
nites toroanus Karsten 185S]; Gloriaceras n. g.
[Typus Art Gloriaceras correai n. sp. ] ; Bueno­
ceras n. g. (Typus Art Buenoceras Ioboi n. sp. �
Bei den neun aufgestellten arten handelt es
sich urn: Holcophylloceras hemandezi n. sp.,
Hypophylloceras wiedmanni n. sp., Eogaudry­
ceras helmsi n. sp., Eogaudryceras (Eo tetrago­
nites) cabaricoi n. sp., Toxoceratoides (Colom­
bocera toides) renzoni n. sp., Pseudoptychoceras
gilberti n . sp., A coneceras gu tierrezpalmae n. sp.,
Melchioritespalmeri n . sp., Valdedorsella (?) co­
lombiana n. sp., Pseudohaploceras leiuaense n.
sp., Carloscaceresiceras caceresi n. sp., Carlosca­
ceresiceras monteroi n. sp., Corteziceras cortezi
n. sp., Pseudosaynella ralphimlayi n. sp., Lewe­
siceras (?) ulloai n. sp., Trochleiceras (Trochlei­
ceras) .juliuerti n. ·s p. , Trochleiceras (Jaumapor­
taiceras) ho((stetteri n. sp., Cheloniceras (Epi­
cheloniceras) camachoi n. sp., Cheloniceras
(Epicheloniceras) carlosacostai n . sp., Cheloni­
ceras (Epicheloniceras) jimenoi n. sp., Cheloni­
ceras (Epicheloniceras) barreroi n. sp., Cheloni­
ceras (Epicheloniceras) Pflrdoi n. sp., Chelonice·
ras (Epicheloniceras) leonhatdtriedeli n.sp, Che­
l�niceras (Epiche/oniceras) huertasi n. sp., Vectt­
liltes (Zambranoites) zambranoi n. sp.,Vectisi·
tes (Zambranoites) cruzi n. sp., Vectisites (Zam­
branoites) mateusi n. sp., Vectisites (Zambra­
.
noltes) cadenai n. sp., Vectisites (Zambranites)
duquesi n . sp., Hypacanthoplites (?) dognaliciae
n. sp., Juandurhamiceras juandurhami n. sp.,
Juandurhamiceras giraldoi n. sp.,Acanthohopli­
tes bigoure ti{orme n. sp., Acanthohoplites
seunesiforme n. sp., Acanthohoplites qui-
CENTRAL COLOMBIA BY AMMONITES 1 09
{[ ,;�£�> ·
..._ : ·
tasuegno n. sp., Acanthohoplites odiosus n. sp.,
Acanthohoplites pluricostatum n. sp ., A cantho­
hoplites eleganteante n. sp., Acanthohoplites
luisperezi n. sp., Acan thohoplites (?) leptocera­
ti{orme n. sp., Acanthohoplites (?)serpen ti{o nne
n. sp., Colombiceras (?) (oreroi n. sp., Colombi­
ceras (?) sarmientoi n. sp:;Bargasiceras (?) juan­
wyatti n. sp., Douuilleiceras tarapacaense n. sp.,
Douvilleiceras abozagloi n. sp., Eodouuilleiceras
pedrocarvajali n. sp . , Eodou uille iceras (?) tequen­
damai n. sp., Dufrenoyia boteroi n. sp., Du(re­
noyia hansbuergli n. sp., Neodeshayesites alber­
toalvarezi n. sp., Neodeshayesites cingulatum n.
sp., Stoyanowiceras cintatum n. sp., Riedelites
quebradanegra n. sp ., Riedelites filos �., n. sp.,
Parahoplites hubachi n. sp., Parahoplites tristan
n. sp., Engonoceras olgaluciae n. sp., Engonoce­
ras (?) duartei n. sp., Platiknemiceras c olom­
biana n. sp., Knemiceras pegnai n. sp., Rinconi­
ceras rinconi n. sp., Lyelliceras caru<Ualorum n.
sp., Lyelliceras pseudolyelliforme n. sp . , Lyelli­
ceras isaacleai n. sp., Ralphimlayites apuloense
n. sp., Tegoceras benauidescaceresi n. sp., Co­
dazziceras {ina n. sp. , Mammites nodosoidesap­
pelatus n . sp., Mammites fugax n. sp.,Mammites
scutulatus n. sp., Franciscoites suarezi n. sp.,
Imlayiceras (?) ralphimlayi n. sp., Fagesia za­
nel/Qi n. sp., Paramammites colombianus n. sp.,
Hoplitoides hern.anmojicae n. sp., Hoplitoides
lacabagnae n. sp., Hoplitoides lagiraldae n. s p .,
Nice(oroceras /Joyacaense n. sp., Peroniceras
robertogarciae n. sp., Peroniceras diabloi n. sp.,
Peroniceras guerrai n. sp., Prionocycloceras
portarum n. sp., Prionocycloceras (?) cucaita­
ense n. sp., Gloriaceras paulinae n. sp., Zumpan­
goceras ospinae n. sp., Zumpangoceras (?) sorae
n. sp., Buenoceras loboi n. sp., Protexanites
cucaitaense n. sp.,
Die obengenannten Ammoniten·Arten stel­
len eine biostrat igraphische Abfolge dar, die den
Zei traum vom Oberen Aptium bis zum Unte­
ren Santon1um �:epriisentiert. Folgende Ammo­
ni ten-Zonen (Faunen Zonen) werden vorgeschla­
gen : 1) Zone des Du(renoyia sanctorum und
Stoyanowiceras treffryanus im liegenden; 2)
Zone d'!s Parahoplites (?) hubachi und Acan­
thohoplites (?) leptocerati{orme, im unmittel­
baren Hangenden des Oberen Aptiums; 3) Zone
des Neodeshayesites columbianus und Douui­
lleiceras solitae fiir das Untere AJbium; 4) Zone
des Platiknemiceras colombiana, Rinconiceras
rinconi und Lyelliceras pseudolyelliforme fiir
das Mittlere Albium; 5 ) Zone des Mammites
nodosoidesappelatus und Franciscoites suarezi
fiir das Untere Turonium; 6) Zone des Gloriace·
ras correai, Protexanites . cucaitaense und
Codazziceras scheibei ftir das Untere Coniacium ;
7) Zone des Len ticeras baltai und Paralen ticeras
sieuerse flir das Untere Santonium.
Flir das Obere AJbium, das gesamte Ceno­
manium, das Obere Turonium und das Obere Co­
niacium wurden keine Ammoniten-Zonen er­
kannt. Schliejllich, werden einige lithostratigra­
phische Einheiten im de tail beschrieben und
Anweisungen tiber ih ren Fossilinhalt gegeben.
110 F. ETAYO
REFERENCES
A NDERSON, F. M. 1 938.- Lower Cre taceous
deposits in California and Oregon, Geol.
Soc. A mer. Spec. Paper, No. 1 6, XX+339
pp., p ls. 1 -8 4.
A NTHULA, D.J. 1899.- Uber die Kreidefossilien
des Kau hasus mit einem allgemeinen
Ue berblick u ber die Entwicklung der Sedi­
mentiirbildu ngen des Kau kasus. In Fr.
Frech and G. von A rthaber, Neue ForB·
chungen in den Kaukasische� Liindern,
A b t. 1, Beitr. Paliiont. Geol. Osterr. - Un­
garns, v. 1 2, Heft III, pp. 55-1 59; pls. 2- 1 4.
BASSE, E., 1 928. · Quelques invertebres cretaces
de Ia Cordillere A ndine. Bull. Soc. g eol.
France, ser. 4, v. 28, nos. 3, 4, 6, pp. 1 1 3-
1 4 7, pis. 7-8, Te:tt. figs. 1-20.
-- 1 946. - Sur deux A mmonites nouvelles du
Coniacien du Sud-Ouest de Madagascar:
Subbarroisiceras n.g. · mahafalense n. sp.,
Bull. Soc. geol. France, ser. 5, vol. 1 6, nos.
�-3,pp. 11- 76, pl.2.
-- 1 94 7. - Les peuplements malgaches de Ba­
rroisiceras, A nn. Paleont., v. 33, pp. 9 7-
1 77, pls. 1 ·9, Text-fig. 1 · 7.
1 948. - Quelques ammonites nouvelles du
--
Cretace de Colom bie (A m. Sud.). Bu ll.
Soc. geol. France. ser. 5, vol. 1 8, nos. 8 · 9,
pp. 691 -698, pl. 25.
, - 1950.- Quelques mollusques du Cretace de
- pp. 71 7-788, p is. 25·29, Text-figs. 1 - 1 7.
Colo m bie. Bull. Soc. geol. France, ser. 5,
vol. 20, nos. 4, 5, 6, pp. 245·255, pl. 1 1,
Tex t-fig. 1 .
BATALLJidR, J. R. , 1 954.- Los engonoceratidos
en Espana. Dr. D. Francisco Padillo Va­
quer.Homenaje postumo. Univ. de Barce­
lona. Fac. de Ciencias, pp. 1 73- 1 78, 3 figs.
BENA VIDES-CA CERES, V. E., 1 956. - Creta�
ceous System in Northern Peru. Bull.
A m er. Mus. Nat. Hist. , v. 1 08, art. 4, pp.
359-493, pls. 31 -66, Tex t-fig: 1-58.
B OHM, J., 1 898.· Ue ber Ammonites Pedemalis
v: Buch. Zeitsch . Deu tsch. Geol. Ges,.el.,
v. 5 0, pp. 1 8 3·201, pls. V- Vlll.
.fJOSE, E. 1 923.· A lgunos faunas cretacicas de
Zacatecas, Durango y · Guerrero. Bol. lnst.
Geol Mex ico, no, 42, pp. 1·21 9, pls. 1-1 9.
BOTERO, A . G. , 1 936. - Bosquejo de Paleonto­
logl'a Colom biana. . An. de Ia Esc. NaL de
Minas, Medellin, n. 35, pp. 86, 70 figs.,
1 0 pis.
BOULE, M. , LEMO I1:J!J;
P. and THE VENIN, A.
1 906-1 9 0 7. - Paleontologie de Madagascar
3. Cephalopodes cretaces des environs de
Diego-Suares. A nn. Paleont., v. I, (asc. 4
( 1 906), pp. 1 73:.!.92, pis. 14·20. Ibid. v. 2,
fasc. 1 ( 1 90 7);· pjiiil -56; pis. 1 - 8, Paris.
· SERNA
BREISTR OFFER, M. , 1 936.· Sur quelques
cephalopodes du Cretace de Colombie.
C. R. Soc. geol. France, fasc. 9, pp. 1 55-
1 5 7.
-- 1 94 7. Sur les zones d 'amm onites darn;
l'A lbien de France et d 'A ngleterre. Trav.
Lab. geol. Grenoble, v. 26, pp. 1 -88.
-- 1 952.- Sur quelques ammonites de !'A l­
bien in{erieur de Madagascar. C. R. Soc.
geol. France, fasc. 1 5, pp. 266·268.
-- 1 952.- Sur Ia decouverte de Knem icerati­
nae (A mm onites albiennes) en Equateur,
en Colom bie et au Venezuela C. R. A cad.
Sci. Paris, v.234, no. 27, pp. 2633-2635,
-- 1 952.- Sur la position systematique du
genre Ptychoceras d 'Orb. Trav. Lab. geol.
Greno ble, v. 29 ( 1 951), pp. 4 7-54.
-- 1 965.· Vues sur les zones d 'ammonites de
l'A lbien. Mem. B.R.G.M., no. 34, pp. 3 1 1 -
3 1 2, 1 ta ble, In, Colloque sur le Cretace
inferieur (Lyon, 1 963).
BREISTROFFER, M. and VILLO UTREYS, 0.
1 953. Les A mmonites Albiennes de Peille
(Alpes-Maritimes) . Trav. Lab. geol. Gre­
no ble, v. 30, pp. 69· 74.
BR UGGEN, H. 1 9 1 0. - Die Fauna des unteren
Senons von Nord-Peru. In, G. Steinmann.
Geol. und Pal. v. Siidamerika 1 6, Neues
Jahrb. Min. Geol. und Pal. B.-Bd. vol. 30,
B UCH, L. von 1 850.· Uber Ceratiten. A bhandl.
k. preuss. Akad. Wiss. zu Berlin ( 1 848),
pp. 1-30, pls. 1 · 7.
BURCKHARDT, C. 1 9 1 9 · 1 92 1 . Faunas Jura­
sicas de Symon (Zaca tecas) y fau nas Cre·
tacicas de Zu mpango del R io (Guerrero).
II. El Cre taceo Superior del Zu mpango
del R io (Estado de Guerrero). lnst. Geol.
Mexico. Bol. No. 33, pp. 7 7· 1 35 ( 1 9 1 9),
pis. 22-32, ( 1 921).
-- 1 9 25.· Faunas del Ap tiano de Nazas (Du·
rango) . lnst. Geol. Mexico. Bol., n. 46, pp.
1 · 7 1 , pis. 1 · 1 0.
B UR GL, H. , 1954a. - El Cre tticeo Inferior en los
alrededores de Villa de Leiva, Boyaca, lnst.
Geol. Nal. Colom bia,Bol. Geol., v. 2, n. 1,
pp. 5-22, pls. 1 ·4.
-- 1 954b.· El Cretaceo Superior en la region
de Gir,ardot, Inst. Geol. Nal. Colom bia,
Bo'l. G�ol., v. 2, n. 1, pp. 23-48, pis. 1-8,
Tex t-fig's. 1 - 1 2.
-- 1 955.- El Anticlinal de Apulo. lnst. Geol.
Nal. Co�om bia , Bol. Geol. v.3, n. 2, pp. 2·
22, pls. �1;4,
 ZONATION OF THE CRETACEOUS OJI' CENTRAL COLOMBIA BY AMMONITES
-- 1 956b.- La variabilidad de Ia amanita
Duf:renoya texana Burckhardt. lnst. Geol.
NaL Colo m b ia , Bol. Geol., 11. 4, nos. 2-3,
pp. 1·22, pls. 1-3.
-- 1 95 7. · Bioestratigrafia de Ia Sabana de
Bogota y alrededores. fiist. Geol. Nal. Co­
lombia, Bol. Geol., v. 5, n. 2, pp. 1 1 3-185,
pls. 1-20.
-- 1 96 0.- Geologia de Ia peninsula de la Gua­
jira.. Inst. Geol. Nal. Colom bia, Bol. Geol.,
v. 6, nos. 1 -3, pp. 1 29-1 68, pis. 1-8.
-- 1 961.- HiBtoria Geologica de Colom bia.
Rev. A cad. Col. Cienc. Ex. FiB. Nat. vol.
XI, n. 43, ·pp. 1 3 7-1 91, Text-fig. 1-41,
4 tab.
-- 1 96 1 b.- Geologia de los alrededores de
Ortega. Boletin de Geologia, Uniuersidad
Industrial de Santander, Bucaramanga.
no. 8, pp. 2 1 ·38, Tex-fig. 1, 1 map.
-- 1 96 1 c.- Sedimentacion ciclica en el geo­
sinclinal cretticeo de Ia Cordillera Orien­
ta/ de Colombia. Inst. Geol. Nal. Colombia,
Bol. Geol., u. 7, nos. 1-3, pp. 85-1 18, Tex t­
fig. 9.
-- 1 963. - Die rhythmischen Bewegungen der
Kreidegeosynklinale der Ostkordillere
Kolumbiens. Geol. Rundschau, 11. 53, n. 2,
pp. 706-731, Text-figs. 1-9.
CACERES GIR ON, C. and ETA YO SERNA, F.
1 969.- Bosq uejo geologico de Ia region
del Tequendama. 1er. Cong. Col. Geol.
Opusculo gu ia, Exc. pre-congr., pp. 1-22,
Tex t-figs. 1-6.
CAMPBELL, C. J. and B UR GL, H. 1 96'5.· Sec­
tion through the Eastern Cordillera of
Colombia, South A merica, Geol. Soc.
A m. Bull. , u. 76, n. 5, pp. 56 7-590, pl. 1,
figs. 1 -6. List of fossils in Document no.
8393, L ibrary of Congress, Washington.
CANTU CHAPA, A. 1 963.- Etude bioestratigra­
phique des ammonites du centre et de
l'est du Mexique (Jurassique superieur et
Cretace). Mem.. Soc. geol. France, ser. 5,
v. 42, fasc. 4, no. 99, pp. 1-1 02, pls. 1-8,
Tex t-figs. 1-3, 6 charts.
CASEY, R. 1 949.· The ammonite genus Uhli·
gella in the English A lbian. Geol. Mag. ,
1101. 86, n. 6, pp. 333-345, Tex t·fig. 1.
- 1 954. - New genera and subgenera of
Lower Cretaceous ammonites. J. Was­
hingto n A cad. Sci., u. 44, n. 4, pp. 106-
1 1 5, Tex t-figs. 1-1 0.
111
CASEY, R. 1 96 0.- A monograph of the ammo­
noidea of the Lower Greensand. Pt. 1, pp.
i-nx u i + 1 - 44, pi& 1 -1 0, Te:ct-figs.
1 - 1 3, Pelaeontogr. Soc. London (1 959).
1 96 1 . · A monograph of the ammonoidea
--
of the Lower Greensand. Pt. 2, pp. 45- 1 1a
pis. 1 1 ·25, Tex t-figs. 1 4-38, Palaeon togr.
Soc., London (1960).
-- 1961 b.- The Stratigraphical Paleontology
of the Lower Greensand. Paleontology.
v. 3, part. 4, pp. 487-621, pl. 7 7-84, Text-
.
figs. 1-14.
1 961 c.- The Cretaceous (A lbian) Ammonite
genus Platlknemiceras Bataller. Ann. Mag.
--
Nat. Hist., Ber. 1 3, u. 1 4, pp. 353-357, Te:c t·
figs. 1-2.
-- 1961d. - A monograph of the ammono idea
of the Lower Greensand. Pt. 3,pp. 1 1 9-21 6,
pls. 26-35, Tex t-figs. 39-68, Palaeontogr.
Soc., London (1 961).
-- 1 962.- A monograph of the ammonoidea
of the Lower Greensand. Pt. 4, pp. 21 7-288,
pis. 36-42, Text-figs. 69-1 03. Palaeon togr.
Soc. London (1 962).
-- 1 964.- A monograph of the ammonoidea
of the Lower Greensand. Pt. 5, pp. 289-398,
pls. 43-66, Tex t-figs. 1 04-1 44. Palaeon togr.
Soc. London ( 1 9 63).
-- 1 965.- A monograph of the ammonoidea
of the Lower Greensand. Pt. 6, pp. 399-546,
pis. 6 7-90, Tex t-figs. 1 45-206, Palaeon togr.
Soc. London ( 1 964).
-- 1 966. · A monograph of the ammonoidea
of the Lower Greensand. Pt. 7, pp. 54 7-582,
pis. 91 -9 7, Text-figs. 207-219. Palaeontogr.
Soc. London ( 1 966).
'·.
COLLIGNON, M. 1 9 3 7.- Paleontologie de Mada­
gascar. 22. Les ammonites pyriteuses de
L ' Aptien d' A n tanatanamirafy. A nn. Pa­
Mont., u. 26, pp. 1 0 7-1 32, pls. 1-3, Text­
figs. 1-13.
-- 1 949.- Recherches sur les faunes albiennes
de Madagascar. I. L 'A lbien d' A m bari­
maninga. Ann. glol. Sero. des Mines
Madagascar, fasc. 16, p. 1 - 1 28, pls. 1-22,
Text-(igc. 1 -29.
-- 1 950.- Recherches sur les faunes albiennes
de Madagascar. 3. L 'A lbien de Komihevi­
tra. A nn. g eol. Sero. Mines, fasc. 1 7, pp.
2 1 -54, pls. 3-9, Text-figs.. l-4.
-- 1 962.- A tlas des fossiles caracteristiques
de Madagascar (Ammonites) . Fasc. 9 (Ap­
tien). Seruice Geolog. Tananarive, pp. 1 -
64, pls. 2 1 5-240.
112 F. ETAYO - SERNA
1 963.- A tlas des fossiles caracteristiques
--
de Madagascar (Ammonites). Fasc. 1 0 (Al­
bien) .. Seroice Geolog. Tananarive, pp. 4-
1 84, pls. 241-31 7.
COQUAND, H. �
851. · Position des Ostrea ca­
lumba et b1aurlculata dans le groupe de la
craie interieure. Bull. Soc. geol. Fr., t. 1 4,
ser. 2, pp. 745- 770.
DIENER, C. 1 925.- Fossilium catalogus. I: A ni­
malia. Pt. 29, A mmonoidea neocretacea,
pp. 1 -244.
,
DOUVILLE, H. 1 904.- Mission scientifique en
Perse par J. de Morgan. 3. Etudes G eolo·
giques, pt. 4, Paleontologie, Mollusques
fossiles. 380 pp. , 25 pls. Paris.
1 9 1 6. - Les terrains secondaire& dans 1e
--
Massif du Moghara. Paleontologie. Mem.
Acad. Sci. Inst. de France, v. 54, ser. 2,
pp. 1 - 1 84, pls. 1 -21, Tex t-figs. 1-50.
DOUV
·
E
ILL , R. 1 906.- Sur des ammonites du
Cretace Sud-Americain. Ann. Soc. roy.
Zool. et Maloc. de Belgique, v. 41, pp. 142·
1 55, pls. 1 -4, Text-figs. 1 -5.
-- 1 9 1 1.- Palaeontologia Uniuersalis. Ser. Ill,
fasc. 3.
DR USHITS, V. V. and KHIAMI, N. 1 9 70. - Struc­
ture of the t.�ep ta, protoconch walls and
initial whorls in early Cretaceous ammo­
nites. Paleont. Joum. n. 1, pp. 26-38, p ls.
1 -2, Tex t-figs. 1-5.
DUBOURDIEU, G. 1 953. - A m monites nouvelles
des Monts du Mellegue. Bull. Seru. Carte
geol. A lgerie, Pa.leont. ser. 1, n. 1 6, 76 pp.,
pls. 1 -4, Tex t-figs. 1-20.
D URHAM, J. W. 1 946.- Upper Aptian nau ti­
loids from Colombia. J. Paleontology, v.
2 0, n. 5, pp. 428-434, pls. 6 2-64, Tex t­
figs. 1 -5.
-- 1 947.- Nuclear whorls of Heminautilus
etherl ngtoni Durham. J. Paleontology,
vol. 21, n. 6, pp. 5 76- 5 7 7, Tex t-figs. 1-3.
DIMITRO VA, N. 1 9 6 7. - Les fossiles de Bulga­
rie. rv: Cretace in{erieur. Cephalopoda
(Nau tiloidea et A mmonoidea). Academie
Bulgare des Sciences, pp. 1 -424, 93 figs.,
93 pls., Sofia.
ETA YO SERNA, F. 1 964.- PosiciOn de los faunas
en los depositos cretcicicos colombianos y
su valor en la su bdivision cronol6gica de
los mismoi. BoL Geol., UniL•. Ind. Scmtan­
der, n. 1 6-1 7, pp. 5-1 4 1, 8 fig8,1 Bucara­
manga.
-- 1 968 a.· Sinopsi8 entratigrafic a de la region
de Villa de Leiva y zonas proximas. Bol.
Geol. Univ. Ind. Santander, n. 21, pp. 1 9-
32, Tex t-figs. 1 -2, 1 map.
' -- 1 968 b.- El Sistema Cre taceo en la region
de Villa de Leiva y zonas proximas. _Geol.
Colom bianD, n. 5, pp. 5-74, Text-frgs. I-
1 8, 1 map.
-- 1 969. - Lenticeras baltai Li8son en Colom­
bia y su probable posicion zonal santonia­
na. Geol. Colom biana, n. 6, pp. 1 7-29, pls.
1 -3, Tex t-figs. 1 ·4.
-- 1 9 72.· Cretaceous section : Villa de Leiva
to Loma Piedra Gorda. 1 3 A nn. Field Con/,
Col. Soc. Pe tr. Geol. Geoph., pp. 35-59,
1 m ap.
D.BARBERO.
ETAYO SERNA, F., G. RENZONI,
1 9 76.- Contornos sucesivos del mar Creta­
ceo en Colombia. Mem. Primer Congr. Co­
lom bwno de Geologia, p.. 2 1 7-252, figs.
1 - 1 8.
FALLOT, E. 1 885.- Etude geologique sur les
etages moyen& et superieur& du· terrain
Cre tace dans le Sud·Est de la France. A nn.
sci. geol., 1Jol. 1 8, pp. 264, pls. 1-8.
FALLOT, P. 1 9 1 0.- Sur quelques fossiles pyri­
teux du Gault des Baleares. Trav. Lab.
geol. Fac. Sci. Grenoble, v. 9, fasc. 2, pp.
62·90, pis. 1 -3, Tex t·figs. 1-8.
-- 1 920a.- O bservations sur diverses especes
du Gargasien bathyal alpin et en particu­
lier sur Ia faune de Bliex. In Kilian and
o thers: Con trib. etude ceph. Paleocret. S.
E. de Ia France. Mem. Explic. carte geol.
det. France, pp. 229·266, pls. 1-3, Text­
figs. 1 -21.
-- 1 920b. La faune des marnes ap tiennes et
albiennes de Ia region d 'A ndraitx (Major­
que). Trab. Mus. Nac. de Cien. Nat. ser.
Geol. , n.26, pp. 1-68, pls. 1 -3, Tex t-figs.
1 -25.
FALLOT, P. and TERMIER. H. 1 923. A m moni- .
tes nouvelles des lies Baleares. Trab. Mus.
Nac. de Cienc. Nat., ser. Geol., n. 3 2, pp.
1 -83, pls. 1 -6, Text-figs. 1-32, Madrid.
FORBES, E. 1 845.- Report on the fossils (rom
Santa Fe de Bogota, presen ted to the
Geological Society by Evan Hopkins, Esq.
F. G.S. Quart. Jour. Geol.. Soc. London,
v. 1, pp. 1 74·1 79, Tex t-figs. B.
FRITZCHE, C.H. 1 924.- Neue Kreidefaunen
aus Sudamerika (Chile, BoliiJia, Peru , Co­
lumbia). 5. Amm oniten aus Turon und
Senon IJOn Colum bien. In G. Steinmann,
Geol. und Pal. v. Sudam erika. 2 7, Neues
Jahrb. Min. Geol. und Pal. B-Bd., vol. 5 0,
pp. 324·333, pl. 4, Tex t-figs. 7-9.
GABB, W. M. , 1 862. Synopsis of the Mollusca
of the Cre taceous FormDtion, Including
the geographical and stratigraphical range
and synonymy. Proc. A mer. Phil. Soc.,
vol. 8, pp. 5 7-25 7.
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
-- 1 8 64.- Descrip tion o f Cretaceous fossils.
ln Paleontology o f California, u. 1, sect.
4. GeoL Suru. California, pp. 5 7-243, p is.
9-32.
-- 1 8 69.- Descrip tion of new, and revision
ofpreuiously descri11ed, Cretaceous fossils.
ln Paleon tology of California, u. l , sect. 2.
Geol. Suro. California, pp. 1 2 7-205, plJI.
1 9-36.
-- 1 8 8 1. - Descrip tion of a collection of fos­
sils, made by Dr. A n tonio Raimondi in
Peru. Jour. A cad. Nat. Sci. Ph il. , 2nd. ser.,
uol.8, part.3, pp. 2 63-336, pls. 35-43.
GERHARDT, K., 1 8 9 7a.· Beitrag zur-Kenn tniss
der Kreide-formation in Venezuela und
Peru. lri G. Steinmann, Geol. und Pal. Su­
dameriha. 5. Neues Jahrb. Min. Geol. und
pal. B-Bd. 11, pp. 65-11 7, pls. 1 -2. Tex t-figs.
1-6.
-- I 8 9 7b.- Be itrag zur Kenntniss der Kreide­
formation in Colum bien. Ibid., pp. 1 1 8·
208, pls. 3-5, Tex t-figs. 7·20.
GlGNO UX, M. 1 9 2 0. - Les Phylloceratides du
·
PaleocrrHace. In Kilian and o thers. Con trib.
•.
.
Etude Ceph. Paleocret. S. -E. FrancP..
Mem. Explic. carte giwl. det. France, pp.
85- 1 01.
GLASUNO VA, A. E. 1 953. Ap tian and A lbian
ammonites from Kopet Dag, Lesser and
Greater Balhhans and Mangyshlah [in
Russian l Trud. Vses. Nauchno·Issled.
Geol. Inst. Moscow. 9 7 pp,, 2 7 pls.
GROSSO UVRE, A. De 1 8 94.- Recherches sur
Ia Craie superieure. Part 2. Paleon to logie.
Les amm onites de la Craie superieure.
Mem. carte geoL det. France (1893).
pp. 1 -264, pls. 1 ·39, Tex t-figs. 1 ·89.
HA YASA KA , /, and FUKADA, A . 1 95 1. On
the on togeny of Barroisiceras minimum
Ya be from the Upper A m monite bed in
Hohkaido. Jour. Fac. Sci. Hokhaid6 Uniu.,
ser. 4 (Geol. and Min. ), uol. 7, n.4, pp.
324--330, pls. 1·2, Tex t·fig. 1 -2.
HITZEL, E. 1 905,- Sur les fossiles de l'e tage
A lb ien recueillis par M.A. Gue bhard dans
la region d 'Escragnolles (A lpes-Maritimes) .
Bull. Soc. geol. France, ser. 4, uol. 2, pp.
8 74-880.
HOUSA, V. 1 9 6 7. Lewesiceras Spath (Pachydis­
cidae, A m monoidea) from the Turonian
of Bohemia, Sbornik Geol. Ve 'd. Paleont.
u, 9, pp. 7·50, pls. 1 -8, Tex t-figs. 1·8.
HUBACH, E. I 931. Exploracio n en Ia region de
Apulo·San A n tonio-Viola. Bol. Min. Petr.,
t·4, n. 25-27, pp. 4 1 -60, 1 map.
113
HUMPHRE Y, W. E. 1 949.· Geology of the Sie­
n-a de Los Muertos A rea, Mexico ( With
descrip tions of Aptian Cephalopods from
the La Peiia Formation). Geol. Soc. A m.
B u ll., u. 6 0, n. l, pp. 89·1 76, 2 figs., 1 8 p ls.
HYATT, A. 1 900.· In Zittel-Eastman Tex tbook
of Paleontology, 1st. English edition, pp.
502-604,Text-figs. 1049-1259, MacMillan,
London.
-- 1 903. Pseudoceratites of the Cre taceous.
Edited by T. W. S tanton. Monogr. U. S.
Geol. Suro. , uol. 44, pp. l -351, pls. I-4 7.
IMLA Y, R. W, 1 944.- Cretaceous formations of
Cen tral A merica and Mexico. Bu ll. A m.
A ss. Petr. Geol., uo/. 28, n. 8,pp.1 0 7 7·11 95,
Tex t-figs. 1 · 1 6.
-- 1 945. · Ammonites from the Dierks limes­
tone of Sou thern A rkansas. Journal of
Paleon tology, uol. 1 9, n. 3, pp. 2 7 7-28 1,
pls. 42-43, Tex t·(igs. l-4.
-- 1 96 0. · Early Cretaceous (A lbian)ammoni.
tes (rom the Chitina Valley and Talkeetna
Moun tains, A laska. U. S. Geol. Suro. Prof,
Pap er 354-D, pp. 8 7-1 14, p ls. I l -1 9, Tex t·':
figs. 2 1 -24 .
JA COB, Ch. 1 905. · Etude sur les ammonites e t
sur [ 'Horizon Stratigraph ique d u G iseme n t
d e Clansayes. Bu ll. de I a Soc. geol. de
France, ser. 4, u. S, pp. 399-432, pls. 1 2· 1 3,
Tex t-{igs. 9.
·.
-- 1 90 7. · Etudes paleon to logiq ues e t strati·
graphiques sur Ia partie moyenne des te­
rrains cretaces dans les A lpes francaises e t
les regions uo isines. These, pp. 1 ·3 1 4, pls.
1 · 6, Tex t-figs. 1 · 1 4, Greno ble.
-- 1 9 08. - E tude sur quelques ammo­
nites du Cre tace moyen. Mem. Soc. geo l.
France, Paleont., u. 1 5, n. 38 ( 1 9 0 7) , pp.
1 ·64, pls. 1-9, Tex t-figs. 1 ·44.
JA COB, Ch. et TOBLER, A. 1 906. E tude
stratigraphique et paleon tologique du
Gault de Ia Vallee de Ia Engelberger AA,
Mem. Soc. Pateont. Suisse, u. 33, pp. 1 ·26,
pls. l -2, Tex t·figs. 1·3.
JUL/VER T, M. 1 968. Colombie (prem iere par­
tie), Precambrien, Paleozoique, Mesozoi�
q u e et in trusions d 'dge mesozoique·Ter­
tiare. In Lexique Strat. /nternat., uo/. 5,
fasc. 4a, A merique latine (sous Ia direction
de R. Hoffstetter) , . pp. 1 ·651, Tex t·fig, 1 -
2 7, Paris.
KARRENBERG, H. 1 93 5. · A m monitenfaunen
aus der Nordspanischen O berkreide. Pa­
leon tograph ica. Vol. 82, Part A, pp. 1 25-
1 6 1 , pls. 30-33, Text-{ig. 5.
 1 14 F. ETAY O · SERNA
KARSTEN, H. 1 8 58.- Vber d ie geognostischen
Verhaltnisse des westlichen Colum bien,
der heu tigen Republihen Neu-Granada
und Equador. A m tl. Ber. iiber die 32. Ver­
Jammlg. d. Deu tsch. Natur/orsch. und
Arzte zu Wien 1 856, pp. 80·1 1 7, pis. 1 - 6,
Sections, 1 m a.P
_ :.
.
B
: -- . ... . ·· J 86. · C:eologie de l'ancienne Colom bie
.
Bolivarienne, Venezuela, Nouvelle-Grena­
de et Ecuador, pp. 1 ·62, pls. 1 -6, sections,
1 m ap., Berlin.
-- 1 94 7.- La situacion geogn6stica de Ia
Nueva Granada. Conferencia dictada en
Viena- 1 8 56. (Trod.). Rev. A cad. Col. Cien.
Ex. Fis. Nat. v. 7, n. 2 7, pp. 3 6 1 - 3 8 1 , p ls.
1 -6, sectio ns, I map.
KILIAN, W. 1 9 07-1 9 1 3. - Unterkreide (Palaeo·
cre tacicum). In F. Frech: Le thaea Geog·
nostica. 2 Teil, Mesozoikum, vol. 3,
Kreide. Lief. I, 1 9 0 7, pp. 1 - 1 68; L ief, 2,
1 91 0, pp. 1 6 9-28 7, pls. 1 -8; L ief, 3, 1 9 1 3,
pp. 289-3 98, pis. 9· 1 4.
KILIAN, W. and REB O UL, P. 1 9 1 5. Contribu­
tion a /'etude des faunes Pateocretacees
du Sud-est de Ia France. 1. La faune de
l 'Ap tien in{erieur des environs de Monteli­
mar (Drome). (Carriere de l'Home d 'A r·
mes), Mem. Expl. carte geol. det. France,
pp. 1-221, pis. 1 -9.
KNECHTEL, M.M. 1 94 7. In M. M. Knech tel, E.
"'· R ichards, and Mary V. Rathbun. Meso·
zoic fossils of the Peruvian A ndes, Pt.3,
Cephalopoda. Johns Hopk ins Univ. Stu­
dies in Geol., no. 1 5, pp. 8 1 - 1 50, p ls. 1 3-
4 7, Text-figs. 1 -24.
KOENEN, A. von 1 8 9 7. - Ue ber Fossilien der
Unteren Kreide am Ufer des Mungo in
Kame run, A bh. honig. Ges. Wiss. zu G o t­
tingen, math. -phys. Kl. , N. F. , Bd. 1 , N. 1 ,
pp. 1-48, pls. 1-4.
-
· ·-.- 1898.- Nach trag zu Ue ber Fossilien der
Unteren Kreide am Ufer des Mungo in
Kamerun; ib., pp. 49-65, pis. 5- 7.
KOSSMAT, F. 1 8 95- 1 898. · Untersuchungen
ii ber die siidindische Kreideformation.
Be itr. Paliiont. - Geol. Oster.· Ungarns u.
des Orie n ts, Band 9 ( 1 8 95), pp. 9 7-203,
pl. 1 5-25; Band 1 1 ( 1 89 7) , pp. 1 -4 6, pls. 1 -
8 , Tex t-fig. 1 - 3 , Band 1 2 ( 1 8 98), pp. B9-
1 52, pis. 1 4� 1 9.
KUMMEL, B. 1 956.· Post-Triassic nau tiloid ge­
nera. Museum Comp. Zool. Harvard, Bull••
v. 1 1 4, n. 7, .p. 324-484, pl. 1 -28, Tex t-figs.
1 -35.
· -,��':-. . .. .
-- 1 964. · In R. C. Moore (ed. ) : Trea tise on
invertebrate paleontology, Pt. K, Mollus­
ca 3, Nau tilQ.idea, Nau tilida, pp. K383-
K4 66, figs. ?tJ
0-33 7.
K UMMEL, B. and DECKER, J. M. 1 954.- Lo­
wer Turonian ammonites fro m Texas and
Mexico. Jour. Paleon tology, v. 28, n.3, pp.
3 1 0-3 1 9, pis. 30-33, Text-fig. 1 0.
LANE, F. W. , 1 960. · Kingdom of the Oc topus,
pp. i·xxvii +l -3 00, pis. 1 ·48, Tex t·figs, I -
1 3, Sheridan House, N. Y.
LA UBE, G. C. and BR UDER, G. 1 8 8 7. - A m mo·
niten der bohm ischen Kreide. Palaeonto­
graphica, v. 33, pp. 2 1 7-239, pls. 23·29.
LEANZA, A.F. , 1 968.· A lgunos ammonites
nuevos o poco conocidos del Turoniano
de Colom bia y Venezuela. Act. Geol. L i­
lloana, Tucuman. vol. 9, ,pp. 1 89-2 1 3, pls.
1 · 7.
-- 1 9 68 b.- Peroniceras buergli n. sp. (A mm.
Collignoniceratidae) del Coniaciano de
Colom bia. A ct. Geol. L illoana, Tucumcin,
vol. 9, pp. 231 -235, p l. 1 , Tex t·fig. 1 .
LISSON, C. I. 1 925.· A lgunos fosiles d e l Peru.
Bol. Soc. Geol. Peru, t. l , pp. 23-30, pls. l -
3.
-- 1 9 3 7. - U n ammon ites holo tipo d e G a b b
cuyo genera se ignoraba. Revista d e Cien­
cias. Ano 38, n. 4 1 9, pp. 23-26, pls. 1-3.
LORIOL, P. de 1 882.- Etudes sur Ia faune des
couches du Gault de Cosne (Nievre).
Mem. Soc. Pal. Suisse, vol. 9, pp. 1 - 1 1 8,
pls. 1 - 1 3.
MAHMO UD, IBRAHIM GAMAL EL-DIN. 1952.
Sur le genre albien d 'A m monites Knemi·
ceras J. Bohm, Trav. Lab. geol. Grenoble,
vol. 29 ( 1 951), pp. 7 7-95, pis. 1 -3.
MARCOU, J. 1 8 75. - Explication d 'une seconde
edition de la Carte Geologique de la Terre,
pp. 1 -222, 1 map. [ Chapitre X VIII (Geo­
logie de L:4. merique du Sud), pp. 1 62- 1 8 1] ,
Zurich.
MA TS UMOTO, T. 1 9 55.- Evolu tion of Peroni­
ceratidae. Trans. Proc. Paleont. Soc. Ja­
pan, N.S., n. l 8, pp. 3 7-44, Tex t-figs. 1-3.
-- 1 965.- A monograph of the Collignonice­
ratidae from Hokkaido. Part I. Studies of
the Cre taceous A m monites from Hokkai­
do and Saghalien - 1 4. Mem. Fac. Sci.
Kyushu Univ., Ser. D., Geology, vo/. 1 6,
n. l , pp. 1 -80, pls. 1 - 1 8, Tex t-figs. 4 0.
1 965b.· A monograph of the Colligno-·
�
nic�J!l-tidae fro m Hpkkaido. Part · f·'-l
.
Studies of the Cretaceous Am monites ··
from Hohhaido cmd Saghalien-1 5. Mem.
Fac. Sci. Kyushu Univ. , Ser.D, Geo logy,
vol. 1 6, n. 3, pp. 209-243, pls. 36·43, Tex t­
figs,,,_l-20.
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA BY AMMONITES
-- 1 966. Notes on Ammonites bourgeoisi
d 'Orbigny {rom the Upper Cre taceous of
France. Trans. Proc. Paleon t. Soc. Japan,
N.S., n. 61, pp. 2 0 I · 206, pis. 26.
-- 1 969.- A monograph of the Co /lignonice­
ratidae from Hokkaido. Part.3. StudieS ·
of the Cretaceous Ain monites from
Hohkaido and Saghalie n-20. Mem. Fac;
-·
;Sci.. Kyushu Univ. Ser. D, Geology, u. 1 9,. .
n. 3, pp. 2 9 7 .. 330, pis. 39-45, Tex t-fi'gs:
1 - 1 4.
-- 1 9 7 0. • A monograph of the Collignonice·
ratidae from Hokkaido. Part IV. Studies
of the Cretaceous A m monites from Ho­
k kaido and Saghalie n - 2 1 . Mem. Fac. Sci.
Kyushu Univ., Ser D, Geology, u. 20, n. 2,
pp. 225-304,. p is. 30- 4 7, Tex t-figs. 1-2 7.
-- 1 9 71.- A monograph of the Collignonice­
ratidae from Hokkaido. Part 5. Studies of
the Cre taceous A m monites from Hokkai­
do and Saghalie n-23. Mem. Fac. Sci. Ky u·
shu Uniu. , Ser. D, Geo logy, uol. 2 1 , n. l,
pp. 1 29-1 62, p is. 2 1 -24, Tex t-figs. l - 1 1.
MATSUMOTO, T. , OBA TA, I.. MAEDA, S.,
and SA TO, T. 1 964. - Yabeiceras (Creta­
(!eous ammonites) from Futaba,Northeast
Japan. Trans. Proc. Paleont. Soc. Japan,
N.S., n. 56, pp. 322 - 3 3 1 , Tex t-figs. 1 ·3,
pl. 48.
MA TS UMOTO, T. and NODA, M. 1 966.- Notes
o n · Ammonites bravaisianus d 'Orbigny
from the Cretaceous of France. Trans.
Proc. Paleont. Soc. Japan, N.S., n. 64, pp.
359-365, pl. 4 0.
MA YER-EYMAR, C. 1 8 93. Ueber Neocomian­
Versteinerungen aus dem Somali/and.
Viert. Naturf. Gesell. Zurich, u. 38, pp.
249-265, p is. 1 -2.
MORRIS, J. 1 848. - Descrip tion of a new spe ­
cies o f Nautilus from the Lower Green­
sand of the Isle of Wigh t. A nn. Mag. N.
Hist., ser. 2, v. I, pp, I 06- 1 0 7, 1 woodcut.
ORBIGN Y, A. D ', 1 84 0- 1 8 4 2. Paleontologie
Francaise. Terrains Cretaces. u. 1 , Cepha·
. lopodes, pp. 1 - 1 20 ( 1 8 4 0), pp. 1 2 1 ·430
( 1 8 4 1), pp. 431 - 662 ( 1 842), 1 48 pis.
Paris.
-- 1 842.- Coqu illes e t echinodermes fossiles
de Colom bie (Nouuelle Grenade), recuei·
llis de 1 8 2 1 a 1 833, par M. Boussingault,
pp. 1 -64, pls. 1·6, Paris.
-- 1 8 4 3.· C o n s id e r a tio n s ge o lo-
giques e t geologico-geographiques sur l'en­
sem ble des mollusques Gasteropodes des
terra ins crtHaces. Bull. Soc. geol. Fran­
ce, vol. I 4, pp. 460-485.
- 1853.- No te sur quelques coquilles fossiles,
recueillies dans les montagnesde Ia Nouve­
lle-Grenade, par M. le general Joaquin
1 15
Acosta. Journal de Conchyliotogie; u. 4,
pp. 208- 2 1 4, plate n.4.
PARONA, C. F. and BONARELLI, G. 1 8 9 7. ·
Fossili albiani d 'Escragnolles, del Nizzar­
do e della L iguria occiden tale. Paleontogr.
!tal., u 2, pp. 5.'3· 1 1 2, pis. 1 0- 1 4.:.,• -
PAVL OW, MAP., l 9 01 . · De quelq ues m cwens qui
pourraie n t contribuer a /'elaboration de la
classification genetique /es fqssil.ffff;(;(o ngres
Geologiqu e Interna tional, C.R., VIlle Ses­
sion, en France. Ier. Fasc., pp. 349-352.
PERON, M. 1 896. · Les am monites du Cretace
superieur de /'Algerie. Mem. Soc. · g�ol.
France, Paleont., n. 1 7, pp. 1 -88; pis. 1 - 1 8.
PER VINQ UIERE, L. 1 90 7. - Carte geologique
de Ia Tunisie. Etudes de paleon to/ogie tu­
nisienne. I. Cephalopodes des terrains se­
condaires, pp. 1 -438, pis. I -27. Paris.
-- 1 9 1 0. - Sur quelques ammonites du Creta­
ce A lgerien. Mem. Soc. geol. France, Pate­
ant., ser 1 7, fasc. 2-3, n. 42, pp. l :86, pls.
1 · 7, Text-figs. 38.
REDTENBA CHER, A. 1 8 73. · Cephalopoden­
fauna der Gosauschichten in den nordos­
tlichen A lpen. A bh . K.K. geol. Reich­
sanstalt, Wien, u. 5, n. 5, pp. 9 I -I 4 0, pls.
22·30.
REESIDE, J. B. jr. 1 9 2 7. - Cephalopods from
the lower part of the Cody Shale of Ore­
gon Basin, Wyo m ing. U. S. Geol. Survey,
Prof. Paper 1 50, pp. 1 - 1 9, pls. 1 -8.
1 9 2 7 b. - The cephalopods of the Eagle
Sandstone and related forma tions in the
Western In terior of the United States. U. S.
Geol. Suroey, Prof. Paper 1 5 1, pp. 1 -8 7,
pis. 1 -45, Tex t-fig. 1.
-- 1 932. - The Upper Cre taceous am monite
genus Barroisiceras in the United States,
U. S. Geol. Suroey, Prof, Paper 1 70 B, pp.
9-29, pls. 3- 1 0.
RENZ, 0. 1 9 70. - Ober d ie Gattungen Parengo­
noceras Spath, Knemiceras Bohm und
Neophlycticeras Spath (A m monoidea) aus
den A nden Venezuelas. Eclogue geol. Helv,
u. 63, n. 3, pp. 1 02 1 · 1 05 7, p is. 1 · 1 2, Text­
figs. 1 2.
-- 1 9 7 1 . - D ie Gattungen Hysteroceras Spath
undMortoniceras Meek (A mmonoidea) aus
den A nden Venezuelas. Eclogue geoi. Helv.
u. 64, n.3, pp. 569-609, pls. I·X l, Tex t-figs.
7, I Table.
-- 1 9 72.- D ie Gattungen Puzosia Bayle, Bhi­
mai tes Matsu moto und Desmoceras Zittel
(A m m onoidea) im O beren Albien Vene·
zuelas. Eclogue geol. Helv. , v. 65, n.3, pp.
701 - 724, pis. 1 - 1 0, Tex t-fig. 9.
REYMENT, R. A. 1 954 a.- New Turonian (Cre­
taceous) ammonite genera from Nigeria.
116 F. ETAYO · SERNA
Colon. Geol. min. Resourc., uol. 4, n.2, pp.
1 49-1 64, pls. 1-4, Tex t-figs. 1-5.
-- 1 954 b.- Some new Upper Cretaceous
ammonites from Nigeria. Colon. Geol. m in.
resour., v. 4, n. 3, pp. 248-2 70, pis. 1 -5,
Text-figs. 1 1.
-- 1 955.· The Cretaceous amm onoidea of
Southern Nigeria and the Southern Came­
TOO'JlS Bull. geol Suro. Nigeria, n. 25, pp.
1 · 1 1 2, pls. 1 ·25, Tex t-figs. 1 -46.
-- 1 958.· Some factors in the distribu tion of
fossil cephalopqds. Stockh. Con tr. Geo.,
vol. I, p t. 6, pjJ. 9 7- 1 8 4, pls. 1 - 7, . Text-figs
1-24.
-- 1 958.- Uber einige A m moniten aus dem
Coniac Kolumbiens und Venezuelas, Suda­
merika, Stockh. Contr. Geol. v. 2, p t. 1,
pp. 1 ·25, p ls. 1·4, Text-figs. 1 · 1 0.
-- 1 9 72a. - No te sur le genre Hoplitoides dans
l'A tlantique du Sud. Bull. Soc. geol. Fran­
ce, ser. 7, vol. 1 2, n. 5, pp, 913-9 1 5, pl.35.
1 9 72b. · Vermuteter Dimorphismus bei
--
der A mmonitengattung . Benueites, Bull.
geol. lnstn. Univ. Upsala N. S., v. 3, n. 1,
(1 9 71), pp. 1 · 1 8, pls. 1 · 1 0, Tex t-figs. 5.
_
-- 1 9 72c. · Some Lower Turonian ammonites
from Trinidad and Colom bia Geol. For.
S tockh olm Forh., v.94, part.2, pp.357-368,
Tex t-figs. 1 ·8.
RIEDEL, L. 1 938.- A mmonitas del cretacico in­
ferior de Ia Cordillera Oriental. Estudios
geol(>gico s y paleontol6gicos so bre Ia Cor­
diltera Oriental de Colom bia, pt. 2, De­
pbrt. Minas y Pe trol, Min. de Ind. y Trab.
Rep u blica de Colom bia, pp. 7-8 0, pls. 3-14.
R OCH, E. 1 926. · Sur quelques ammonites du
Revest, pres de Toulon. Bull. Soc. geol.
France, ser 4, v. 26, pp. 28 7-294, pl. 1 8,
Tex t-figs. 1·3.
ROD, E. and MA YNC, W. 1 954. - Reuision of
Lo wer Cretaceous stratigraphy of Vene­
zuela. Bull. A mer. Ass. Petr. Geol. v. 38,
n. 2, pp. 1 93�283, Tex t-figs. 1 -3 0.
SARASIN, Ch. 1 893. - Etude sur les Oppelia du
groupe du Nisus et les Sonneratia du
groupe du bicurvatus et du raresulcatus.
Bull. Soc. geol. de France, ser. 3, vol. 2 1 ,
pp. 149-1 64, pls. 4-6, Tex t-figs. 1 - 1 2. .
SA RASIN, Ch. and SCHONDELMA YER, Ch.
1 902.- Etude monographique des ammo·
nites du cretaciq ue inferieur de Chatel­
Saint-Denis. Mem. Soc. pal. Suisse, vol.
29, pp. 95- ��jP. f
l� 1 2-2 , Tex.t· figs. 3 � 6. _
SA YN, G. 1 8 90.· Description des amm onitides
du Barremien du Dje bel-Ouach pres Cons­
tantine, A nn. Soc. Agric. Hist. Nat. Lyon.
Ser. 6, v. 3, pp. 1 35-208, pis. 1 ·3.
-- 1 920.- Les Phy lloceras gargasiens du Sud­
Est de Ia France (Especes nouvelles ou
peu connues). In Kilian and o thers: Con­
trih. Etude Cevh. Paleocret. S. -E. Franr.P..
Mem. Explic. carte geol. det. France, pp.
1 9 1 ·2 03, pls. 1, tex t-figs. 1·3.
SCHLUTER, C. 1 8 7 1 · 1 8 76.- Die Cephalopoden
der o beren deutschen Kreide. Palaeonto·
graphica, Band 21, ( 1 8 71), pp. 1 -24, pl.
1 -8, Band 22, (18 72), pp. 25-1 20, pl. 9-35,
Band 24 (18 76), pp. X+121 ·264, pls. 36-55
SCOT!', G. 1 93 7.- A mmonites of the genera
Sonneratia and Douvllleiceras from the
Cretaceo�s of Colo m bia. J. Paleont., v. 2,
n. 1, pp. 34·3 7, pl. 6, Tex t-figs. 1·2.
-- 1 940.· Cephalopods from the Cre taceous
Trinity Group of the Sou th-Centra l Uni­
ted States. The University of Texas Publi·
cation. No. 3945, pp. 969- 1 1 06, p is. 55·
68, Tex t-figs. 1 39-1 79.
SEUNES' J. 1 8 8 7.- Note sur. q uelques A mmoni­
tes du Gault. Bull. Soc. geol. de France,
ser. 3, vol. 1 5, pp. 567-571, pls. 1 1 · 14.
SINZO W, I.
1 9 06.- Die Beschreibung einiger
Douvilleiceras·Arten aus dem Oberen
Neocom. R usslands. Verh. russ. k. Min.
Gesell. St. Petersburg, ser. 2, v. 44, pp.
1 5 7- 1 9 7, p ls. 1 -5, Tex t-fig, 1 ·5.
-- 1 90 7. - Untersuchung einiger A m m oniti­
den aus dem Un teren Gault Mangyschlaks
und des Kaukasus. Verh. russ. k. Min. Ges·
sell. St.. Petersburg, ser. 2, v. 46, pp. 455-
5 1 9, pis. 1 -8, Tex t-figs. 1 - 7.
SOMMERMEIER, L. 1 9 1 0. · D ie Fauna des Ap­
tien und A lbien in nordlichen Peru. In, G.
Steinmann, Geol. und Pal. u. Sudamerika,
1 5, Neues Jahrb. Min. Geol. und Pal.,B. -Bd,
v. 30, pp. 3 1 3 • 382, pis. 7-1 5, Text-figs.
1 - 3 7.
SORNA Y, J. 1 955.· A m monites nouvelles du
Cretace de Ia region des Monts du Melle­
gue (Constan tine).Bu ll. Sero. Carte geol.
A lgerie, ser. 1, Paleon t. , n. 1 8, pp. 1 ·4 0,
pls. 1 -2, Tex t-figs. 1 · 1 9.
-- 1 95 7. · A m monites du Coniacien du Massif
de Ia Haute Medjerda (Constantine, A lge·
rie). Bull. Soc. geol. France, ser. 6, u. 8,
pp. 1 8 7- 1 96, pl. 1 6, Tex t-figs. 1 ·2.
SORNA Y, J. ed., 1 95 7b.· Lexique Stratigraphi·
que International, vol. I, Europe, Fascicule
4a, France, Belgique, Pays-Bas, Luxem·
bourg, Fascicule 4a- VI, 5 m aps. Int. Geol.
Congr. Comm. S trat. Paris.
.
·� :�'
-- 195Tc. Cretace (France). In Lexique stra·
tigraphique International. v. 1, Europe,
fasc. 4a, 403 pp. , Geol. Congr. Comm.
Strat. Paris.
 ZONATION OF THE CRETACEOUS OF CENTRAL COLOMBIA
-- 1 968.- Lexique Stratigraphique Interna­
tional, volume VIII, Termes Stratigraphi­
ques Majeurs, article Ap tien, pp. 1-1 09,
Geol. Congr. Comm. Strat. Paris.
SO WERBY, J. 1 8 1 2. - The Mineral Conchlogy
of great Britain, vol. 1,� pp. 1 · 234, pls.
1 - 1 02, London.
·�
SPATH, L. F. 1 921. O n Cretaceous Cephalopoda
from Zululand. A nn. S. African Mus., v.1 2,
•
part 7. n. 1 6, pp; 2 1 7-321, p ls. 1 9-26.
1 922. - On Cretaceous ammonoidea from
A ngola, collected by Prof. J. W. Gregory,
--
D. Sc., F. R.S. Trans. R oy. Soc. Edin., vol.
53, part. 1, n. 6, pp. 9 1 - 1 6 0, pls. 1-4, Tex t­
figs. 1-4.
-- 1 923.- On the A m monite Horizons of the
Gault and Contiguous Deposits. Mems.
Geol. Surv. Summary of Progress, Great
Brit. ( 1 922), pp. 1 3 9-1 49.
-- 1 9 23 b. · A monograph of the ammonoidea
of th e Gault. Part I, pp. 1- 72, pls. 1 -4,
tex t-figs. 1 4, Palaeontogr. Soc. London.
-- I 925a. - A monograph of the ammonoidea
of the Gault, Part II, pp. 73-1 1 0, pls. 5-8,
Tex t-figs. 1 5-25.
_
! - 1 925b.- On Upper A lbum ammonoidea
{rom Portuguese East Africa. A nn. Trans·
vaal Mus. , vol. 1 1, part. 3, pp. 1 79-200,
pls. 28-37.
-- 1 93 0a. - On some A m monoidea from the
Lower Greensand. A nn. Mag. Na t. Hist.,
ser 1 0, v. 5, n. 29, pp. 4 1 7-464, pls. l 4-1 7.
-- 1 930b. - A monograph of the ammonoidea
of the Gault. Pat. VII, pp. 26 7-3 1 1 , pls.
25-30, Text-figs. 8 7-1 02.
-- 1 93 0c.- The Lo wer Cretaceous ammo·
no idea; with no tes on A lbian cephalopoda
from Hazara. The {ossl1 fauna of the Sa­
mana Range and some neighbouring areas,
Part. V., Palaeont. Indica, n.s., v. 1 5, pp.
5 0-66, pls. 8-9.
-- 1 93 1 . · A monograph of the ammonoidea
of the Gault. Part. VIII, pp, 313-378, pls.
3 1-36, Tex t-figs. 1 03-1 24.
- 1 934. - A monograph of the ammonoidea
of the Gault. Part. XI, pp. 443-4 96; pls.
49-56, Text-figs, 1 53-1 73,
- 1 942.- A monograph of the ammonoidea
of the Gault. Part. XV, pp. 669- 720, Text-
figs. 245-248, 4 t�bles.
·
- 1 943,- A monograph of the ammonoidea
of the Gault. Part. XVI, pp. 721-78 7,
STENZEL, H. 1 964.- ln R. C. Moore (Ed.) Trea­
tise on inuertabrate paleontology, Pt. K,
Living Nautilus, pp. K59-K93, figs. 43-68.
BY AMMONITES 1 17
STEINMANN, G. 1 88 1 . · Ue ber Tithon und
Kreide in den peruanischen A nden. Neues
Jahrb. Min. Geol. und Pal. , u. 2, pp. 1 3 0-
1 53, pls. 6 - 8, Tex t-r�g. l .
-- 1 882.- Ue ber Jura und Kreide in den A n­
den. Neues Jahrb. Min. Geol. und Pal., v. l
pp. 1 66· 1 70.
-- 1 929. - Geologie von Peru, x ii+448 ppjJ .
pls., Heidelberg.
STIELER, C. 1 920.- Uber sogenannte Mortoni­
ceraten des Gau lt. Centralblatt. Min. Geol.
und Paleont.,n. 1 ·2·, pp. 345-352, 392-400,
Tex t-figs, 1 -9.
STO YANO W, A. 1 949.- Lo wer Cretaceous Stra­
tigraphy in south-eastern A rizona, Geol.
Soc. Amer. Mem., n.38, pp. vii+ 1 69, pis.
1-27.
-- 1 958.- Su ture of Acanthohoplites aschil­
taensis (Anthula), Geol, Soc. A m erica
Bull., u. 69, n. 5, pp. 607-61 0, pis. 1 -2.
STURM, F. 1 90L- Der Sandstein uon Kiesling­
swalde in der Grafschaft Glatz Fauna.
Jahrbuch h. P. geol. Land. Bergakademie
zu Berlin ( 1 9 00), pp. 39-98, pls. 2-1 1 ,
Tex t-figs. 2.
S UESS, E. 1 866.- tiber A m moniten. Sitz. -Ber.
k. Akad. Wiss., math. -nat. Kl., uol. 52,
A bt. I (1 8 65), pp. 71-89, Wien.
S UTTON, F. A., 1 946.- Geology of Maracaibo
Basin; Venezuela. Bull. A mer. Ass. Petr.
Geol., v.30, n. I O, pp. 162 1 - 1 739, pis. 1-9,
Tex t-figs. 1-1 1.
THO MEL, G. 1 964.- Les zones d 'A mmonites de
l'Aptien des Basses·A lpes. C.R. A cad. Sc.
Parts, t. 258, n. 1 7, Gr. 9, pp. 4308-431 0,
TIETZE, E. 18 72.- Geologische und palaontolo­
gische Mittheilungen aus dem siidlichen
Theil des Banater Gebirgsstockes.
Jahrb. k.k. geol. Reichsanstalt, v.22, p .
35-1 42, pis. 2-9.
TO VBINA, S. Z. 1 9 71 (1 9 70). A new Aptian
genus of the ammonoid fam ily Paraho­
plitidae. Paleontological Jour. v.4, n.3,
pp, 33 7-345, 11 figs.
UHLIG, V. 1883,- Die Cephalopodenfauna der
Wernsdorfer Schiechten. Denkschr. k.
Akad, Weiss. Wien, math. -nat. kl., v.46,
A bt. 2, pp. 1 2 7-29 0, pls. 1-32.
WEDEKIND, R. 1 9 1 6.- Uber Lo bus, Suturallo­
bus und Inzision. Centralbl. Min. Geol.
Pal., pp. 1 85-1 95, Tex t- figs. 1 -6.
VENZO, S. 1 936. Cefalopodi del Cretaceo me­
dio-superiore dello Zululand. Pal. Italica.
u. 36, n.s., pp. 59-1 44, pls. 5-12.
118 F. ETA YO
WIEDMANN, J. 1 962.· Ammoniten aus der vas­
c ogotischen Kreide (Nordspanien). I. Phy·
lloceratina, Lytoceratina. Paleontograph i­
ca, A, v. 1 1 8, pp. 1 1 9-237, Text-figs. 1 058,
pis. 8-1 4.
-- 1 962b.- Unterkreide - A mmoniten Von
Mallorca. . 1. Lieferung: Lytoceratina, Ap­
tychi. Akad. Wiss. Lit. Abh. math. -nat.
K1., n. 1, pp. 1 - 1 48, pis. 1 · 1 0, Text-figs.
1 ·36, I Table.
-- 1 962c.· Habitus, Skulptur und Sutur bei
Kreide-A mmonoideen. N. Jb. Geol. Pa­
laont. A bh., v. 1 14, n. 3, pp. 3 1 7-332,
pl. 1 8, Text-figs. 1-4.
-- 1 962d.- Die systemati8che Stellung von
Hypophylloceras Salfeld N. J.B. Geol.
Palaont. A bh., v. 1 1 5, n.2, pp. 243-262.
pl. 1 6, Text-figs. 1-5.
-- 1 964.- Unterkreide·A mmoniten von Ma­
llorca 2. Lieferung: Phylloceratina. A bh.
Akad. Wiss. u. Lit., math. -nat. Kl.,
n. 4, pp. 1 5 7·264, pls. 1 1·21, Text-figs.
3 7-64.
-- 1 966.- Stammesgeschichte und System
der posttriadischen A m monoideen. Ein
Uberblick. 2. Teil. N. Jb. Geol. Palaont.
A bh., v. 1 2 7, H. 1, pp. 13-81, figs. 1 4-4 7,
pls. 3-6.
WIEDMANN, J. and DIENI, I. 1 968. Die Krei·
de Sardiniens und ihre Cephalopoden,
Pal. Italica, vol. 64, pp. 1-1 71, pis. 1-1 8,
Text-figs. 1 - 1 01.
WILLEY, A. 1 896, Zoological observations in
the South Pacific : Quart. Jour. Micro.
Sci., n.s., v. 39, n. 154, pp. 2 1 9·231, pl. 1 3.
* *
• SERNA
-- 1 9 02.· Contibution to the natural histo­
ry of the pearly Nautilus: in Zoological
results based on material collected in New
Britain, New Guinea, Loyalty Islands and
Elsewhere, Part 6, pp. 691 -830, pis. 75-83,
1 map., Text-figs. 1 · 1 5. Cambridge U.
Press.
WRIGHT, C. W. and MA TSUMOTO, T. 1 954,
Some doubtful Cretaceous ammonite ge­
nera from Japan and Saghalien. Mem.
Foe. Sci., Kyushu Univ., Ser. D, vol. 4,
n.2, pp. 1 07- 1 34, pls. 7-8, Text-figs. 1 0 7-
1 34.
WRIGHT, C. W. 1957.- In W. J. Arkell, Bernard
Kummel, and C. W. Wright: Mesozoic A m·
moidea. In R. C. Moore (ed.) : Treatise on
invertebrate paleontology, Pt. L, Mollusca
4, Cephalopoda, A mmonoidea, pp. LBO·
L490, Text-figs. 124-555. Univ. Kansas
Press.
YOUNG, K. 1 957.· Upper Albian (Cretaceous)
ammonoidea from Texas. J. Paleont.
v. 31, n. 1, pp. 1-33, pls. 1-1 0, Text-figs.
1·4.
-- 1 963.· Upper Cretaceous A m monites
from the Gulf Coast of the United States.
Univ. Texas Pu bl. no. 6304, ix +1 42 pp.,
Plates 1 ·82, Text-figs. 1 ·34.
-- 1 966.· Texas Mojsisouicziinae (A mmonoi·
dea) and the zonation of the Fredericks­
burg. Geol. Soc. A m. Mem., n. 1 00, pp. uii
+225, pls. 1 -38, Text-fig. 1 2 1 .
-- 1 9 72.- Cretaceous paleogeography: impli­
cations of endemic ammonite faunas.
Bureau Econ. Geol. Uniu. of Texas at A us­
tin. GC 72-2, pp. 1-12, Text-figs. 1 -4.
*
 · .

F I G URE S 3 - 17
120 F. ETA YO · SERNA
FIGURE 3
A) Terminology utilized in this paper when
describing whorl sections; M vl : median
ventral line AV: adventral : M: middle;
AU: adumbilical.
B) Terminlogy utilized in describing the ribs;
Ao: adoral ; Aa: adapical; Cc: concavity;
Bp: branching point; Cv: convexity ; Pr:
proyection.
C) Whorl section of Phylloceras buchianum
forbes); C-217. a) hypotype at a whorl
height (=H.) of 3 mm (x 5 2/3); b) hypo­
type at H:6 mm (x 5 2/3).
D) Growth striae and constriction ridge pro­
file of Holcophylloceras hemandezi n.sp.,
h olotype C- 1 3 at H : 8 mm (x 1 5/8).
E) Rib profile of Hypophylloceras wied­
manni n.sp. holotype C-225, at H : 14
\
mm (x 1 1/3J.
F) Terminology utilized in this work descri·
bing the elements of the suture line: E :
external lobe; L: lateral lobe; U: umbili­
cal lobe, 1: internal lobe. E/L: seddle be­
tween ; E and L lobes.
G) Types of ribbing described in this work.
H) Whorl section of Holcophylloceras hernan­
dezi n. sp., holotype C-13, at H : 8 mm
(x 2 6/8).
I) External suture line of Holcophylloceras
hernandezi n. sp., holotype C-13, at H: _
8 nim (x 2 6/8).
L) Whorl section of Eogaudryceras helmsi
n. sp., holotype C-223, a) at H : 4 mm
(x 1 1/2); b) at H : 14 mm (x 1 1/2).
M) Whorl section of Eogaudryceras (Eo tetra­
gonites) cabaricoi n. sp., holotype C-513,
at H: 14mm (x 1 1/2).
N) Part of external suture line showing L, of
Eogaudryceras (Eote tragonites} cabaricoi
- n. sp., holotype at H : 8mm (x 1 1/2).
0) Whorl section of Toxoceratoides (Colom­
bocera toides)renzoni n. sp., holotype LB·
1 0. a) at H: 9.5 mm (x 2 8/9); b) at H: 3.5
mm (x 2 8/9).
P) Complete suture line of Toxocera toides
(Colomboceratoides) renzoni n.sp.,morph
I, C-220, at H : 5 mm.
Q) Whorl section of Melchioriles media Rie­
del, hype type C-127, at H: 6mm (x 3 1/6).
R) External suture line of Melchioriles media
Riedel, hypotype C-127, at H : 6 mm
(x 2 1/6).
S) Whorl section of Pseudoptychoceras gil­
berti n. sp., holotype C-223, at H:5 mm
(x 3).
T) Complete last suture line of Pseudoply­
choceras gilberti n. sp., paratype C· 223,
at H: 6 mm (x 3).
U) Whorl section of Pseudohaploceras leiua­
ense n. sp., holotype VL-10-17, at H :
19.5 mm ( x 1 1/2).
V) Profile of the adoral ridge of a constric­
tion in Valdedorsella (?) colom biana n. sp,
paratype Sta Ana, at H: 12 mm (x 1 1/4).
W) Whorl section of Valdedorse/la (?) colom­
biana n. sp., paratype Sta. Ana, at H: 13
mm (x 1 1 /5).
X) Whorl section of A coneceras gu tierrezpal­
mae n. sp., paratype C-223. a) at H : 3mm;
b) at H : 5 .5mm (x 6).
Y) _Part Qf external suture.line of A coneceras
gulierrezpalmae n. sp., holotype C-223 at
H: 8 mm.
Z) Profile of a growth striae of Aconeceras
gutierrezpalmae n. sp., holotype C-223,
at H: 8mm (x 2 7/8).
$) Whorl section of Melchiorites palmeri
n. sp., holotype C-223, at H : 6mm (x 2 1/3)
Whorl section of Corteziceras (?)n.sp.ind.,
C-5 13, at H: 14 mm (x 1 1/3).
fi) External suture line of Pseudohaploceras
leiuaense n. sp. holotype at H : 13 mm.
 FIGURE 3

'..:. (
FIGU RA 3

J
f .
3ds
M
z

I
�
D
.l
·I
E

r/lo-+
AU

A
1 c

Pr ,,�\

~
e
I

a
8
t
���.
I I
\.� - I
.• J G H

o
Tn•olribWIII

o
P'IP(P':!pP)p,.· . . • · · · · \
r,HI'� Pl • • • • · · • n
)I'P
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 122 . F.
A) Whorl section of Carloscacercsiceras cace­
resi n. sp., paratype C-13-27, at H:20mm
(x 1 1/2).
B) External suture line of Carloscaceresiceros
caceresi n.sp.,paratype C-13-37, atH:lBmm.
C) Whorl section of Corteziceras cortezi n.
sp., . Sta. Ana, holotype at H :20 mm
{x 1 1/2).
D) a) Whorl section of Lewesiceras (?) ulloai
n. sp., c-310, holotype, at H : 8 mm (x
1 1/2); b) profJle of the adapical ridge of
a constrictions, ib. at H : 9.5 (x 1 1/2).
E) External suture line of Carloscaceresiceras
monteroi n. sp., holotype C-13-33, at H. :
16 mm (x 1 1/2).
F) . Whorl section of Corteziceras latecostata
(Riedel), hypotype C-225, at H : 21 mm
(x 1 1/2).
G) Whorl section of Trochleiceras (Jauma­
portaiceras) hoffstetteri n. sp., holotype
C-215-5, at H: 8mm (x 1 1/2).
H) Part of external suture line of Trochleice­
ras (J.) ho{{stetteri n. sp., C-215, paraty­
pe, at Ji: 5 mm.
I) Whorl section of Trochleiceras (Trochlei­
ceras) juliverti n. sp., holotype C-215-2,
at H : . 4 mm (x 3). · W)
J) . Whorl section of Carloscaceresiceras · (?)
chimuense (B.C.) n. sp.,h)lpotype VL-Est.
33a, at H: 34mm (x l l/7).
K) Part of external suture line of Carloscace­
resiceras ('!) chimuense (B. C.) n.sp., holotype
VL-Est. 3 aa, at H:18 mm (x 1 1/2).
L) Whorl; section of Trochleiceras (Jauma­
portiiiceras) hotfstetten· n. sp., C.215 para­
. type at H: 3 mm (x 3).
. '.: ·­
�) . Part of external suture line of Trochleice·
ras (Trochleiceras) juliverti n. sp., C-215-1
· paratype,(x 3), ·· ·
N) Whorl section of Pseudosaynella ralphim-
ETAY O · SERNA
FIGURE 4
layi n. sp., paratYpe C-223x, at H : 10 mm
(x 1 1/2).
0) External suture line of Pseudosaynella ral­
phimlayi n. sp., holotype SV-28 at H : 18
mm (x 1 1/2).
P) Whorl section of Cheloniceras (Epichelo·
niceras) barreroi n. sp., holotype C-514,
at H:17.5 (x 1 1/6).
Q) External suture line of Cheloniceras (E)
barreroi n. sp., holotype C-514, at H: 9
mm (x 1 1/2).
R) Whorl section of Cheloniceras (Epichelo­
niceras) carlosacostai n.sp., paratype
C-223-13, at H : 19 mm (x 9/10).
S) External suture line of Cheloniceras (Epi­
cheloniceras) carlosacostai n. sp., holoty­
pe C-223-1 6, at H : 16.5 (x 1 1/2).
T) Whorl section of Cheloniceras (Epichelo­
niceras) camachoi n.sp., holotype VL-10·
17, at H :21.5 mm (x 1 1/2).
U) Whorl section of Cheloniceras (Epichelo­
niceras) leonhardtriedeli n. sp., C�23, at
H : 13.5 (x 1 1/3).
V) Whorl section of Cheloniceras (Epichelo­
niceras) amazonarum (Burckhardt), bolo·
type C-5 14, at H: 20mm (x 8/10).
Whorl section of Cheloniceras (Epichelo­
niceras) dognaelvirae n. sp., holotype VL·
12-2, at H : 15 mm (x 1 1/5).
·
X) Whorl section of Cheloniceras (Epichelo­
niceras) jimenoi n. sp., holotype C-504,
at H:21 mm (x 4/5).
Y) Whorl section of Cheloniceras (Epichelo­
niceras) huertasi n. sp., paratype V17-Sa-
11, at H : 11.5 mm (x 1 1/2).
$) Complete suture line of Ch. (E.) huertasi
n.sp., paratype VL7 SA-11, at H:6 mm
(x 1 1/2).
Z) Whorl section of Cheloniceras (Epichelo­
niceras) pardoi n. sp., holotype C-504, at
H: 22 mm (x 9/10).
FIGU RA 4 FIGURE 4

Q (. 0

() G
V) n I
H

F
0 L
--u""\.
M ,

~
K
N
� 0

R
124 F. ETA Y O • SERNA
FIGURE 5
A) Whorl section of Vectisites (Zambranoi·
tes) cruzi n. sp., holotype C-504, at H:10
mm (x 3).
B) External suture line of V. (Z. ) cruzi n.sp.,
holotype C-504, at H: 5 mm (x 3).
C) Whorl section of Juandurhamiceras juan·
durhami n. sp., paratype Santa Ana, at
H: 10 mm (x 2 8/10).
D) External suture line of Parahoplites (?) in·
constans Riedel?, C-501-13, at H: 8 mm
(x 1 1/2).
E) Whorl section of Gargasiceras (?) juanwya·
tti n. sp., holotype C-151, at H:11.5 mm
(x 1 1/2).
F) Part of the external suture line of Paraho·
plites hubachi n. sp., morph I, C-127, at
H: 13 mm (x 1 1/2).
G) External suture line of ParahopUtes tris·
ton n. sp., h.olotype C-127, at H: 6 mm
(x 3).
H) Whorl · section of Vectisites (Zambranoi­
tes) zambranoi n. sp., holotype C-504, at
H: 17.5 mm (x 1 1/2).
I) Whorl section of Vectisites (Zambranoi­
tes) duquesi n. sp., paratype C-514, at H:
1 1 mm ( x 1 11-2).
J) External suture line of Juandurhamiceras
juandurhami n. sp. Santa Ana-25, at H : 13
mm (x 3).
K) External su ture line of Vectisites (Zam­
branoites) zambranoi n. sp., C-504, at
H: 12 mm (x 1 1/2).
L) Whorl section of Vectisites (Zambranoites)
cadenai n. sp., holotype C-5 14 at H : 13
mm (x 1 1/2).
M) Whorl section of Du[renoyia boteroi n.sp.
holotype C-223 x-10, at H:20.5 mm (x 1
1/2).
·
N) Ex ternal suture line of Du[renoyia bo te­
roi n. sp., holotype C·223x-10, at H:13
mm (x 1 1/2).
0) External suture line of Vectisites (Zam­
branoites) zambranoi n. sp., holotype
C-504, at H:10 mm (x 1 1/2).
P) External suture line of Vectisites (Zam·
branoites) cadenai n. sp., holotype C-514,
at H:10 mm (x 1 1/2).
Q) Whorl section of Aoanthohoplit�s seunesi·
forme n. sp., holotype C-127, at H:12
mm (x 1 1/2).
R) External suture line of Acanthohoplites
seunesi{orme n. sp., holotype C-127, at
H: 12 mm (x 1 1/2).
S) Whorl section of A canthohoplites pluri·
costa tum n. sp., holotype C-127, at H:9
mm (x 3).
T) External suture line of Acanthohoplites
pluricostatum n. sp., holotype C-127, at
H: 9 mm (x 3).
U) Whorl section of Acanthohoplites bigou­
reti{orme n. sp., holotype C-127, at H:
9 mm (x 1 1/2).
V) External suture line of Acanthohoplites
bigoureti[orme n. sp., bolo type C-127, at
H: 7 mm (x 3).
X) External suture line of Acanthohoplites
eleganteante n. sp., holotype C-127, at
H:6 mm (x 1 1/2).
Z) Whorl section of Acanthohoplites quita­
suegno n. sp., holotype C-127, at H: 8
mm (x 3).
iJ) External suture line of A can thohoplites
quitasuegno n. sp., holotype C-127, at H:
8 mm (x 3).
W) Whorl section of Acanthohoplites luispe­
rezi n. sp., paratype C-127, at H: 7 mm
(x 3).
Y) External suture line of Acanthohoplites
luisperezi n. sp., paratype C-127, at H: 6
mm (x 3).
a) Whorl section of Acanthohoplites odio­
us n. sp., holotype C-127, at H:6 mm (x 1
1/2); b) ib. at H : 10 mm (x 1 1/2).
u> External suture line of A. odious n.sp.
holotype C-127 at H: 7 mm (x 3).
o) . Whorl section of Acanthohoplites {?) ser­
penti[orme n. sp., holotype C-127 at H:
5 mm (x 5).
a) Whorl section of Acanthohoplltes (?) lep­
tocerati[orme n. sp., holotype ( x 5).
FIGURA 5 FIGURE 5

.
-
�

.A��
\ /� . l-
A

,.
0 I

�
. ' �Q
�
T

�
Q �·
s
X

0 �
v
u

0 z
w

0
C)&
� A
1t:f lr {\ '' )
\ . )
d
¥
il

a a
FIG URA 6
FIGURE 6

,. .

0
E

�1
M

-���
v�:��
: ·-ir
0

R
128 F. ETA YO
FIGURE 7
A) Wh orl section of Neodeshayesites cinf!u/a­
tum n. sp., h olotype C-501·0, at H : 30mm
(x 1 2/10).
B) Wh orl section o f Neodeshayesites contrac­
tu (Riedel), hypotype C-500-3, at H : 22
mm (x 1 1/2).
C) Superposed rib profiles of Neodeshayesi·
tes albertoaluarezi n. sp., at left of each·
fi gu re; N. columbianus (Riedel) at center
of each figure, and N. karsten i (Marcou) , the
profile transecting the other profiles ex­
cept on the first group of figures on the
left; the single profile (third from left to
right) corresponds to N. columbianus
(Riedel). From left to right at D : 23mm,
D : 37mm; D : 4 2mm; H : 21mm; H : 26mm
(al X 1 1/2).
D) External suture line of Neodeshayesiles
contractu (Riedel), hypotype, C-500-4 at
H :25mm (x 1 1 /2).
E) Rib profiles of N. contractu (Riedel ) , hy­
potype C·5 00 (2), from left to right at
H :9mm ; H : l lmm; H : 14mm; H : 1 8mm,
(all X 1 1/2).
F) Rib branching in N. con trac tu {Riedel)
hypotype C-500 (2), at H : l lmm {x 1 1/2).
G) External suture line of Neodeshayesites
cingula tum n. sp., holotypf' C-501-0, at
H : 24mm (x 1 1 /2).
H) External suture line of N. cingula tum n.sp.
paratype C-5.01-23, at H : 14mm {x 1 1/2).
I) Rib profile of N. contracla (Riedel), after
fig; S.' . plate 7 of Riedel's paper ( 1937); at
·.. ...
- SERNA
H : 1 1.5mm; H : 1 3mm; H : 14mm respecti­
vely (X 1 1 / 2).
K) Rib profile of N. cingulatum n. sp., halo­
type C-501-0, at H: 16mm (x 1 1 /2).
L) Rib profile of N. cingula tum n. sp., halo­
type, C-501·0, at H : 22mm (x 1 1/2).
M) Whorl section of Sloyanowiceras lre{fry·
anus (Karsten ), hypotype La Mesa 10 at
H : 20mm (x 3).
N) External suture line of Stoyanowiceras
tref{ryanus {Karsten), hypetype La Mesa
10, at H : 20mm (x 3).
0) Whorl section of Riedelites ('?) que brada­
negra n . sp., C -200 A, hypotype at H : 1 6
m m (x 1 1/2).
P) Wh orl section of Riedelites o bliquum
(Riedel), C-5 14 (A), at H : 16.5 (x 1 1 /2).
Q) Whorl section of Stoyanowiceras lre{{ry­
anus (Karste n ) after the original drawing
in Karsten ( 1 858, pl.4 , fig.1b), (x 7 /9).
R) External suture of Riedelites ('?) quebra­
danegra n. sp., Q·200 A, hypotype at H :
1 1.5mm ( x 3).
S) External suture line of Riedeliles eslher­
sernae n. sp., C-222A,holotype, at H : l l .5
mm (x 3).
T) Whorl section of Riedeliies esthersernae
n. sp., C 222A, hol otype , at H : 1 6 mm
·
(x 1 1/2).
U) u
Wh orl section of Riedelites {ilos m n. sp.
paratype C-225 (2), at H : 1 6.5mm(x 1 1/2).
 F IGURE 7
FIGU RA 7

A
8 )/! �
E

i
K ·+
I
L
I

Q
 130 F. ETA Y O · SERN A

FIGURE 8

A) Whorl section of Platiknemiceras n. sp.A, P) Whorl section of Mammites scu tulatus n.

C-5 10 A, at H : Smm (x 3 ). sp., holotype C-310, at H : 14 mm (x 1
1/2) .
B) Part of external suture line of Engonoce·
ras olgaluciae n. sp., holotype C-5 1 0A, at Q) Whorl section of Codazziceras {ina n. sp.,
H : 14mm (x 3). paratype C-134, at H : _ 4 mm (x 3).

C) Whorl section of Engonoceras olgaluciae R) Whorl section of Codazziceras scheibei n.

n. sp., holotype C-5 1 0A, at H :9mm sp.,hypotype C-134, at H :13 mm (x 1 1/2).
(x 1 2 / 1 0).
S) Rib profiles of Codazziceras {ina n. sp.,
D) Part of external suture line of Knemiceras p aratype C-134 and C. scheibe (Riedel),
pegnai n. sp., holotype C-1 3 1, at H : 68 hypotype C-134 ; figures to the right re·
mm (lower suture) and H :85mm (upper · ·
p resent the ribs o f latter. H 13 mm (x 1
suture), (x 1). 1/2); long rib on right side at H : 30 mm
(x 1 2 / 1 0). ....,
(�.
..:;J
.
Whorl section of Lyelliceras pseudolyelli·
forme s. sp ., holotype C-255, at H : 15mm T) Whorl sections of Fagesia zanelli n. sp.,
·

(x 2 1 / 10). paratype C-310, a) at H: 6mm ; b ) at H : 15

mm, ( both x 1 1 /2).
F) Whorl section of Knemiceras pegna i n. sp.
holotype C- 1 3 1, at H: 89mm (x 1/4). U) Whorl sections of /mlayiceras (?) ralphim·
layi n. sp., paratype ; a) at H: Groin (x 3);
G) Whorl section of Knemiceras semicosla· b) at H: 24 mm (x 1 2/10).
tum Sommerme ie r, hypotype, VL-Est.
33a. a) at H: 23 mm (x 1 1/2), b) at H : 35 V) Whorl section of Rinconiceras rinconi n.
mm (x 1 2/1 0). sp., paratype C-169, at radius: 13 mm
(x 3).
Whorl section of Lyelliceras carvajalorum
n. sp., holotype C-225, at H : 16 mm (x 3). W) External suture line of Rinconiceras rin­
coni n . sp., holotype C- 169, at H: 8 mm
I) External suture line of L. carvajalorum (x 4 5 /10).
n. sp., paratype C-225, at H : 1 7 mm (x 3).
X) Whorl section of Franciscoiies suarezi ri .
K) External suture line of Pialiknemiceras sp., holotype C-310, at H : 14 mm (x 1
n. sp. A,C-5 10A, at H: 1 1 mm (x 3). 1 / 2).

L) External suture line of Knemiceras semi­ Y) External suture line of Franciscoites sua­
costatum Sommermeier, l e ctotype VL­ rezi n. sp., holotype C-31 0,at H: 1 3 mm
Est 33a, at H: 30 mm (x 1 2/10). (x 3).
M) Whorf section of Codazziceras {ina n. sp., Part of external suture ( L ) of Tegoce·
p aratyp e C-1 34 , at H:12 mm (x 1 1/2). ras benavidescaceresi n. sp., holotype VL.
Est 33a, at H: 23 mm (x 1 2/10).
N) External sutu re line of Codazziceras {ina
n . sp., paratype C-134, H : 12 mm (x 1 1/2). P art of e x ternal su t u re o f Tego ce ras
benavidescaceresi n . sp., h ol o typ e VL.
0) Whorl section of Mammites {ugax n . sp., Est. 3 3a, at H: 23 mm (x 1 2/10).
h olotype C-263, at H : 14 mm (x 1 1/2).
ii) Whorl section of Franciscoites suaresi n.
sp., paratype C-310, at H : 4 m m (x 3).
 FIGU RE 8

FIG U RA 8

J G

\ � ',
K

I
' !
j
\,_
H

0 p

/A\
n IJ

v
0
� t�
X

z
w
132. F. ETA YO · SERNA

FIGURE 9
A) Whorl section of Peroniceras robertogar· L) Last tlXi.emal suture line of Gloriaceras
ciae n. sp., holotype VL 6 Cu-5, at H:13 paulinae n. sp., holotype NRC-CU-5.
mm (x 1 2/10).
M) Whorl section of Prionocycloceras (?)
B) Part of external suture line of Peroniceras cucaitaensen. sp., holotype at H:25mm
robertogarciae n. sp., holotype VL 6 CU· (X 1 2/10).
5, at H : 13 mm (x 1 1/2).
N) Whorl section of Prionocycloceras por·
C) Whorl section of Gloriaceras correai n. sp. tarum n. sp., holotype C-134, at H:20
holotype NRC·CA-1-5, at H:15 mm (x 1 mm (x 1 1/2).
1 /2).
0) Whorl section of Zumpangoceras ospinai
D) Rib profiles of Gloriaceras paulinae n. n. sp., holotype NRC-1, at H:22mm
sp., (on left side), and Gloriaceras correai (x 1 1/7). "
·

n. sp., (on right side), at H:ll mm and H :

2 2 m m respectively (x 1 1/2). P) Whorl section of Zumpangodras ospinai
n. sp., paratype Sora NRC, at H;12mm
.
E) Whorl section of Peroniceras diabloi n.sp. (X 1 2/10).
holotype NRC-3 at H:16 mm (x 1 1/2).
F) Rib profile of P. diabloi n. sp., holotype Q) Whorl section of Protexanites cucaitaense
NRC-3, at H: 16 mm (x 1 1/2). n. sp., holotype VL6-CU3 at H:17mm
(x 1 2/10).
G) External suture line of Peroniceras diabloi R) External suture line of Buenoceras loboi
n. sp., holotype NRC-1-5, at H:22 mm n. sp., holotype NRC-4 (1965), at H:25
9 :H (x 1 1/2). mm (x 1 1/2).
H) External suture line of Gloriaceras correai S) Whorl section of Prionocycloceras ion·
n. sp., holotype NRC - Est. 9, at H. : 22 gispinatum .(Basse), hypotype C-134,
(x 1 2/10). at H:28mm (x 1 2/10).
I) Whorl section ofPeroniceras guerrai n. sp., T) Part of external suture line of P. longis-
holotype NRC-Est. 9, at H: 19 mm (x 1 pinatum (Basse), hypotype C-134, at
1/2). H :23mm (x 1 2/10).
J) External suture line of Peroniceras guerrai U) Whorl section of Buenoceras loboi n. sp.,
n. sp., holotype NRC-Est. 9, at H: 19 mm
· holotype NRC-4 (1965) (x 1 1/2) .
.
·.
(x 1 1/2).
V) External suture line of Hoplitoides her-
Kt Whorl section of Gloriaceras paulinae n. nanmojicae n. sp., holotype C - 3 10, at
sp., paratype NRC-CU-5 ( x f 1/2). H : 26 mm (x 1 1/2).
FIGURA 9 FIGURE ·9

\
j
/ 0
p

M N 0

n
-�l1Jrv?
r"'-
'·
•.
-
Q

j
• I

�"
!/ T
s u
134 F. ETAYO · SERNA

FIGURE 1 0

Scatter diagram of the distribution of spe­ relationships are between the external umbilical
cimens of Carloscaceresiceras carloscaceresi n. width (=Euw) and the height of the whorl
sp. and Carloscaceresiceras monteroi n. sp., in (=H ). Small dots represent sample C-13; large
samples C-13, C-170 according to sizes. The dots represent sample C-170.
FIGURA 10 FIGURE 1 0
.
. -. �...

•
: ...

-·

•
• • •

• •• • •

• • •

• ••

• •
•

•• •

• • • • • •

•
• •
•

'. s '"-------.----"--------H w__________"T"""_____

-r-
10 2 3
136 F. ETAY.O · SERNA

FIGURE 11

Scatter diagram of the relationships be­ type 88Dlple C-223. For the explanation of the
tween Euw and H ,. and the accompanying symbols used to repre11mt types of ribbing eee
variations in type and number of ribs in Chelo· figure 3G. .
nicems (Epicheloniceras) carlosacostai n.. sp.,

NOTE: From fl&ure 11 to 17 the foUowtne •ymbols

wm be u.sed.

SYMBOLS No. OF RIBS

• 46 - 50

• 41 · 45

• 36 . 40

• 31 - 35

26 - 30

21 · 25

0 16 . 20

+ 11 - 15
 .
· FIGURA I I !'. FIGURE I I

---

,., . .

0D

O D'
25 OD

OD'

on
00 D

OD
15
OD OD OD

OD

Hw

10 20 30
138 F. ETAYO · SERNA

FIGURE 12

Scatter diagram of the relationships be­ durhamiceras jUtZndurhllrni n.sp., type �ple
tween Euw and H, and the accompanying Santa Ana, A.P.G. .
variation in type and number or ribs in Juan·

v
 · :;:· . .,.i.'·:· · .
�- �..;..
..

FIGURA 1 2 FIGURE 1 2

·K'

0K '

0K
25
0K ·K

0K
0G
.
K . °K
0G ·K
00f( • 0K
·K
15 0 0K ·K
0K0G

5 �--------�--------
H_w
--------�----------
1 0 20
140 F. ETA Y O · SERNA

FIGURE 1 3
Scatter diagram o f the relationships be­ lleiceras (?) tequendamai n. sp., type sample
tween Euw and . H, and the accompanying va­ C-223.
riation in type and numher of ribs in Eadouvi-
FIGURA 1 3 FIGURE . l 3

OD'

15

t n•

tn• t o•

Hw
10 20
142 F. ETA Y O - SERNA

FIGURE 14

Scatter diagram of the relationships be· rum Biirgl (=S, Sm), and Dufrenoyia hansbuer·
tween Euw and H and the accompanying va· gli n. sp., (=Th). Data from Blirgl 1956b, Table I.
riation in number of ribs in Du{renoyia sancto·
FIGURA 14 FIGURE 14

' Sm �·. I

jl: ·s
::J
....

· Sm
.s
' Sm ·Tb

' Sm ·S ·Sm
·s
·s ·s
·Th
0 S · Sm ·Sm
·s · s · sm ·Th
·s •Tb
· Sm • · Th ·Th · Th
0s · S · Tb• Th

· sm

· sm

Hw

20 30 40
144 F, E T A Y O · SE R N A

FIGURE 15

Scatter diagram of the relationships be­ hayesites columbianus (Riedel), reference sam­
tween Euw and H and the accompanying va­ ple C-12-C.
riation in type and number of ribs in Neodes-
FIGURA 1 5 FIGURE 15

•A
•F
A • • •F'
•F
E • •F •A
•F
•D •A
•A
• n •F •F
•F' •F
A• •F
•F • A
FeF
1 .n ••E
•F' •F •F
•F •F
• •F

Hw
5 �------,---r---�
10 20 30
146 F. ETAYO · SERNA

FIGURE 16
Scatter. diagram of the relationships be­ hayesites albertoaluarezi n. sp., type sample
tween Euw and H and the accompanying va­ G-219.
riation in type and number of ribs in Neodes·
FIGURA 1 6 FIGURE 1 6

Hw

1 0 2 0 30
148 F. ETA YO · SERNA

FIGURE 17

Scatter diagram of the relationships be· yesites karsteni (Marcou), Reference sample
tween Euw and H and the accompanying va· C-501.
riation in type and number of ribs in Neodesha·
FIGURA 1 7 FIGURE 1 7

35

es en
ec

25

8B
ec
•c eA
•n •n
Ee ec
A(B•) 8 A

15

eB

HW
5 �------�---.r--.--
10 20 30
P L AT E S 1 - 1 5
152 F. ETA Y O · SERN A

PLATE 1

Fig. 1 ) Du{rcnoyia codazziana (Karsten), L a Fig. 5 ) Gargasiceras interiectum (Riedel), La

Ye, ( x 1 1/1 0). Ye-5 bis (x 1).

Fig. 2) Du{renoyia sanclorum (Burgi), hypo· Fig. 6) Du(renoyia boleroi n. sp., holotype
type La Ye-7 (x 1 2/100). C-223-x-10, (x 1 ).

Fig. 3) Du{renoyia sanctorum (Burgi), morph Fig. 7) Du{renoyia codazziana (Karsten), La

1, La Ye (x 1 3/1 00). Ye -2 (x 1 ).

Fig. 4) Du{renoyia hansbuergli n. sp., hypoty· Fig. 8) Du{renoyia codazziana �KarstenJ,

pe C-223-x-28 (x 1). morph II, La Cumbre x 1 1/1 ).

......

Plate 1. Ammoni �� from the Du(renoyia sanclorum·Stoyanowlceras tre{{ryanus Assembla-

ge zone . F1gs. 1·8. Upper Aptian. . ·.
154 F. ETA Y O · SERNA

PLATE 2

Fig. 1) Cheloniceras (Epicheloniceras) barreroi Fig. 6) Eodouvilleiceras (?) lequendamai n. sp.,

n. sp., holotype C-514 (x 1 7 / 10). holotype C-223-21, (x 1 12/100).

Fig. 2) Cheloniceras (Epicheloniceras) pardoi Fig. 7) Cheloniceras (Epicheloniceras) (?)

n. sp., holotype C-504 (x 1 12/100).
Fig. 3) Cheloniceras (Epicheloniceras cama·
choi n. sp., holotype VL-10-17 (x 1).
amazonarum (Burckhardt) , hypotype
C-5 14 (x 1).
Fig. 8) Cheloniceras (Epicheloniceras) jimenoi
n. sp., holotype C-514 (x 1).

Fig. 4) Cheloniceras (Epicheloniceras) huerta­ Fig. 9) Cheloniceras (Epicheloniceras) carlos­

si n. sp., holotype VL-7-Sa-11 (x 1). acostai n.sp., holotype C-223-16 (x 1).

Fig. 5) Cheloniceras (Epicheloniceras) dogna­ Fig. 10) Cheloniceras (Epicheloniceras) carlos­

elvirae n. sp., holotype VL-12-2 (x 1 acoslai n. sp., holotype C-223-16 (x 1
5 / 1 00). 16/ 100).

-·

Plate 2. Ammonites from the Du{renoyia sanctoru m-Stoyanowiceras tre{(ryanus Assembla­

ge zone. Figs. 1-10. Upper Aptian.
156 F. ETA Y O · SERNA

PLATE 3

Fig. 1) Pseudosaynella ralphimlayi n. sp., ho­ Fig. 8) Stoyanowiceras treffryanus Karsten,

lotype SV-28 (x 1 2/10). morph I (x 1 3{100).

Fig. 2) Corteziceras cortezi n. sp., holotype Fig. 9) Melchiorites media Riedel, hypotype C·
Santa Ana (x 1). 157 (x 1 72/100).

Fig. 3) Acanthohoplites (?) serpentiforme n. Fig. 10) Valdedorsella (?) colombiana n. sp.,
sp., holotype C-157 (x 1 77/100). holotype Santa Ana (x 89/100).

Fig. 4) Pseudohaploceras leivaense n. sp., ho­

lotype VL-10-17 (x 1).
Fig. 5 ) Juandurhamiceras juandurhami n. sp.,
paratype Santa Ana 31 (x 1 4/ 100).
Fig. 6) Corteziceras (!) n.sp., ind., C-5 13 (x 1).
Fig. 11) Hypacanthoplites (?) dognaliciae n. sp.,
holotype T.P.I. Guaduas (x 1 2/100).
Fig. 12) Stoyanowiceras treffryanus Karsten,
morph II, Penn. 7 (x 1 8/100).

Fig. 7) Corteziceras cortezi n. sp., morph I, Fig. 13) Juandurhamiceras giraldoi n. sp., bolo­
Santa Ana (x 1). type Santa Ana 42 (x 1 8/100).

. .....
..
Plate 3. Ammonites from the Dufrenoyia sanctorum-Stoyanowiceras tre((ryanus Assembla·
ge zone .. Figs. 1-2, 4-8, 10·13. Ammonites from the Parahoplites (?) hubachi-Acan­
thohoplrtes (?) leptocerati(orme Assemblage zone. Figs. 3, 9. Upper Aptian.
158 F. E T AY O · SERN A

PLATE 4

Fig. l) Stoyanowiceros tre{fryanus Karsten, Fig. 5) Juandurhamiceras joepecki n. sp., San-

Pensilvania (x 1 2/10),hypotype. ta Ana 9 (x 1 2/100).

Fig. 2) Juandurhamiceras juandurhami n. sp., Fig. 6) Corteziceras latecosta ta (Riedel), C-255

holotype Santa Ana 30 (x 1). (x 1).

Fig. 3) Riedelites obliquum (Riedel), bypoty- Fig. 7 ) Stoyanowiceras lreffryanus (Karsten)?

pe C-200 D (x 1 2/10). Penn. 13 (x 1 25/1 00).

Fig. 4) Acanthohoplites odiosus n. sp., bolo· Fig. 8) Heminautilus etheringtoni Durham,

type C-127 (x1. 4{100). hypotype (x 1 2/10).

Plate 4. Ammonites from the Du{renoyia sanctorum-Stoyanowiceras treffryanus Assemblll

ge zone. Figs. 1-3, 5, 7-8. Ammonites from the Prahoplites ('?) hubachi-Acantho
hoplites ('?) leptocerati{onne Assemblage zone. Figs. 4, 6. Upper Aptian.
160 F. E T A Y O · S E R N A

PLATE S
Fig. 1) Parahoplites (?) hubachi n.sp., holotype Fig. 6) Parahoplites(!) hubachi n.sp., morph I,
C-127 (x 1 2/10). C-127 (x 1).
Fig. 2) A canthohoplites bigoureti{orme n. sp., Fig. 7) Gargasiceras (?) juanwyatti n.sp., bolo-
holotype C-127 (x 1 4/100). type C·151 (x 1).
Fig. 3) Stoyanowiceras treffryanus (Karsten), Fig. 8) Acanthohoplites luisperezi n. sp., bolo-
Penn. (x 1), hypotype. type C-157 (x 82/100).
Fig. 4) A canthohoplites eleganteante n. sp., Fig. 9) Heminautilus e theringtony Durham,
holotype C-127 ( x 1 4/100). hypotype (x 1 3/10).
Fig. 5) Acanthohoplites triston n. sp., bolo-
type C-127 (x 1).
v

Plate 5 . Ammonites from the Dufrenoyia sanctorum-Stoyano�eras treffryanus Assembla·

ge zone. Figs. 3, 9 . Ammonites from the Parahoplites (?) hubachi-Acanthohoplites
(?) leptocerati(orme Assemblage zone. Figs. 1·2, 4-8 Upper Aptian.
162 F. ETAY O · SERNA

PLATE 6

Fig. 1 ) Vectisites (Zambranoites) zambranoi Fig.ll) Eogaudryceras (Eotetragonites) caba­

n. sp., holotype C-504 (x 1).
Fig. 2) Vectisites (Zambranoites) duquesi n.
sp., holotype C-514 (x 1).
Fig. 3) Cheloniceras (Epichetoniceras) leon­
hardtriedeli n. sp.,hypotype (x 1).
Fig. 4) Vectisites (Zambranoites) cadenai n.
sp., holotype C-514 (x 1).
Fig. ·5) Colombiceras (?) sarmientoi n. sp.,
holotype C-5 13 (x 1).
Fig. 6) Melchiorites palmeri n. sp., holotype
C-223 (x 1).
ricoi n. sp., holotype C-5 1 3 (x 87/10).
Fig.12) Riedelites esthersernae n. sp., holotype
C-222A (x 1).
Fig.13) Colomblceras {oreroi n. sp., holotype
C-200-7 (x 1 1/10).
Fig.14) Eogaudry ceras (Eogaudrycerus) helmsi
n. sp., holotype C-223 (x 1).
Fig.15) Aconeceras gutierrezpalmae n. sp., ho­
Jotype C-223 Jx 1 6/10).
Fig.16) Riede lites (?) quebradanegra nsp., hypo·
type C-200 A (x 1 26/100).

Fig. 7) Riedelites obliquum (Riedel), morph I, Fig.17) Pseudoptychoceras gilberti n. sp.,

C-5 14 A (x 1). Hypotype.
Fig. 8) Gargasiceras in teriectum ( Riedel),
morph I, La Ye 4 h is (x 1 2/10).
Fig. 9) Vectisites (Zambranoites) cruzi n. sp.,
holotype (x 1).
morph I C-223 (x 1 42/100).
Fig.18) Pseudoptychoceras gilberti n. sp., ho·
Jotype C-223 (x 1 44/100).
Fig.19) Toxoceratoides (Colomboceraloides)
renzoni n. sp., holotype LB·10 (x 1).

Fig.10) Melchiorites palmeri n. sp., paratype Fig.20) Vectisites (Zambranoites) mateusi n.

C-200 (x 1). sp., holotype VI- 11-3 (x 1 33/ 100).

Plate 6. Ammonites from the Du{renoyia sanciorum-Stoyano;Jceras tre{fryanus Assembla·

ge zone. Figs. 1·20. Upper Aptian.
164 F . E T AYO · SERNA

PLATE 7

Fig. 1) Riedelites filosum n. sp., holotype . Fig. 6) Douvilleiceras solitae (D'Orbigny),hy·

C-255-6 (x 1 ).
Fig. 2) Acanthohop/ites quitasuegno
holotype C- 127 (x 1).
Fig. 3) Acanthohoplites pluricostatum n. sp.,
h olotype C-127 (x 1).
Fig. 4) Acanthohoplites leptocerati{orme
n. sp., holotype C-127 (x 1).
Fig. 5) Douvilleiceras tarapacaense
paratype C-218 (x 1).
potype C- 160A (x 1).
n. sp.,
Fig. 7) Douvilleiceras solitae (D'Orbigny),hy·
potype C-160A (x 1).
Fig. 8 ) Acanthohop/ites seunesi{orme n. sp.,
holotype C-127 (x 1 1/10).
Fig. 9) Stoyanowiceras cintatum n. sp., bolo·
n. sp., type T. P. Guaduas (x 1 1/10).
v

Plate 7 . Ammonites from the Du{renoyia sanctorum-Stoyanowiceras tre{{ryanus Assembla·

ge zone. Fig. 9. Ammonites from the Parahoplites (?) hubachi-Acanthohoplites (?)
leptocera ti{orme Assemblage zone. Figs. 1·4, 8. Upper Aptian. Ammonites from
the Douvilleiceras solitae-Neodeshayesites columbianus Assemblage zone. Figs. 5-7.
Lower Albian.
166 F. ETAY O · SERNA

PLATE 8
Fig. 1) Eodouuilleiceras pedrocaruaja/i n.sp., Fig. 5) Holcophylloceras hernandezi n. sp.,
holotype C-170 (x 1). holotype C-13 (x 1 8/10).

Fig. 2) Douuilleiccras larapacaense n. sp., Fig. 6) Douvilleiceras tarapacaense n. sp.,

holotype c. 218-10 (x 1). holotype C-218-10 (x 1).

Fig. 3) Neodeshayesites albertoaluarezi n.sp., Fig. 7 ) Douuil/eiceras abozagloi n. sp., holo-

holotype C-129-1 (x 1). type C·215 (X 1 6/100).

Fig. 4) Phylloceras buchianum (Forbe� ,hypo­

type C-217 (x 2).

......
Plate 8. Ammonites from the Douuilleiceras solitae-Neodeshayesites columbianus Assem
blage zone. Figs. 1-7. Lower Albian.
168 F. ETA YO · SERNA

PLATE 9

Fig. 1 ) Trochleiceras juliverti n. sp., holotype Fig. 5 ) Lyelliceras pseudolyelli{orme n. sp.,

•
C-215·1 (x 2 64/100). holotype C-255 (x 1).

Fig. 2) Neodeshayesites karste11i (Marcou),hY· Fig. 6) Ralphimlayites apuloense n. sp., bolo·

pctype C-501·1 (x 1 8/100). type C·510A (x 1 29/100).

Fig. 3) Neodeshayesites albertoalvarezi n. sp., Fig. 7) Neodeshayesites contracta (Riedel),hy·

paratype C-219 (x 1). potype C-500·6 (x 1 9/100).

Fig. 4) Rinconiceras rinconi s. sp., holotype Fig. 8) Neodeshayesites columbianus (Riedel),

C-169-3 (x 1). hypotype C-12-C·S (x 1 4/100).

Plate 9. Ammonites fr�m the Douvilleiceras solitae-Neodeshayesites columbia

nus Assem·
blage z !lne. �tgs. �-3, 7-8: Lower Albian. Ammonites from the Platiknem
iceras
c?lombwna:Rmcomc�ras nnconi-Lyelliceras pseudolyelliforme Assemblage zone.
F1gs. 4-6. Mtddle Albtan.
170 F . E T A YO - SERNA

PLATE 10

Fig. 1) Carloscaceresiceras caceresi n .sp . , halo· Fig. 6) Neodeshayesites karsteni (Marcou)

type C-170·20 (x 1 8/ 100). n. sp., morph I, C-501-14 (x 1).
Fig. 2) Parahoplites (?) inconstans Riedel ?, Fig. 7 ) Neodeshayesites columbianus ( Riedel),
C-501-13 (x 1). paratype C-12C-25 (x 1 12/100).
Fig. 3) Hypophylloceras wiedmanni n. sp., ho· Fig. 8) Neodeshayesites cingula tum n. sp., pa­
lotype C-2fi!'i (x 1). ratype C-5 01-21 (x 1).
Fig. 4) Trochleiceras (Jaumaportaiceras) hoffs· Fig. 9) Neodeshayesites karsteni (Marcou) ,
tetteri n. sp., holotype C-215 (x2 2/10). m orph I C-501 (x 1).

Fig. 5) Carloscaceresiceras mon teroi n. sp., Fig. 10) Neodeshayesites cingulatum n. sp., ho·
holotype C-13-33 (x 2/10). lotype C-501·0 (x 1 8/100).
)

Plate 10. Ammonites from the Parahoplites ('?) hubachi·Acantliohoplites ('?) leptocerati{or­
me Assemblage zone. Figs. 2·3. Upper Aptian. Ammonites from the Douvilleiceras
solitae-Neorl.eshayesites colu mbianus Assemblage zone . Figs. 1, 3-10. Lower AJbian .
174 F. ETAYO · SERNA

PLATE 12

Fig. 1) Engonoceras (?) duartei n. sp., holoty­ Fig. 3) Platiknemiceras sp. ind. B., VL. Est.
pe VL Est. 33a (x 9/10). 33a (x 1).
Fig. 2) Knemiceras pegnai n. sp., holotype C- Fig. 4 ) Enl(onoceras (?) duartei n. sp., holoty­
131 (x 6/10). pe VL- Est. 33a (x 81/100).

Plate 1 2. Ammonites from the Platiknemiceras colombiana-Rinconiceras rinconi-Lyelliceras

pseudolyelliforme Assemblage zone. Figs. 1-4. Middle Albian.
176 F. E T AY O · SERNA

PLATE 13

Fig. 1) Mammites nodosoidesappelatus n. sp. Fig. 10) Mammites scutulatus n. sp., holotype
holotype C.310 (x 1). C·310 (x 1).
Fig. 11) Fagesia zanellai n. sp., holotype C-310
Fig. 2) Franciscoites suarezi n. sp., holotype (x 1).
C-310 (x 1 37/100).
Fig. 12) Prionocycloceras portarum n. sp., ho·
lotypE' C-134 ( x 1).
Fig. 3) Imlayiceras (?) ralphimlayi n. sp., ho·
lotype C-310 (x-1). Fig. 13) Paramammites colombianus n. sp.,
bolotype LVZ-7 (x 1).
Fig. 4) Hoplitoides hernanmojicae n. sp., ho·
lotype C.310 (x 1). Fig. 14) Hoplitoides lagiraldae n. sp., holotype
C-264 (x 1).
Fig. 5) Paramammites colombianus n. sp., bo-
lotype LVZ-7 (x 1). Fig. 15) Hoplitoides (?) sp. ind., C·310 (x 1).
Fig. 6) Ma mmites fugax n. sp., holotype c- Fig.16) Prionocyclaceras longispinatum(Basse ) ,
263 (x 1). hypotype C-134 (x 1).
Fig. 7) Lewesiceras (?) ulloai n. sp., holotype Fig. 17) Codazziceras {ina n. sp., holotype
C-310 (x 1). C-134 (X 1 15/100).
Fig. 8) Hoplitoides hernanmojicae n. sp., para- Fig. 18) Gloriaceras correai n. sp., holotype
type C-310 (x 1). NRC-Ca-1-5 (x 1 26/100).
Fig. 9) Hoplitoides lacabagnae n. sp., holoty· Fig. 19) Gloriaceras paulinae n. sp., holotype
pe (x 1). NRC·CU-5 (x 1).

Plate 1 3 . Ammonites from the Mammites nodosoidesappelatus-Franciscoites suarezi Assem­

blage zone. Figs. 1-11, 13-15. Lower 'furonian. Ammonites from the Gloriaceras
correai-Pro texanites cucaitaense -Codazziceras scheibei Assemblage zone. Figs.
12,
16-19. Lower Coniacian.
178 F. ETA YO · SERNA

PLATE 14

Fig. 1) · Peroniceras diabloi n. sp., holotype Fig; 6) Zumpangoceras (?) sorae n. sp., holo-
NRC-= ( x 1). type NRD-Sora (x 1).
Fig. 2) Buenoceras loboi n. sp., holotype Fig. 7) Nice{oroceras boyacaense n. sp., bolo·
NRC-4 (x 1). type NRC-9 (x 1).
Fig. 3) Peroniceras guerrai n. sp., holotype Fig. 8) Zumpangoceras ospinai n. sp., holoty-
NRC-Est. 9 (x 1). pe NRC-1 ( x 1).
Fig. 4) Prionocycloceras portarum n. sp., ho· Fig. 9) Prionocycloceras (?) cucaitaense n. sp.,
lotype C-134 ( x 1). holotype NRC-5, (x 1 12/100).
Fig. 5) Fagesia zanellai n. sp., holotype C-310 Fig.10) Gloriaceras correai n. sp., holotype
(x 1). NRC-Ca -l 5 (x 1). ·

Plate 1 4. Ammonites from the Mammites nodosoidesappelatus-Franciscoites suarezi Assem­

blage zone. Fi� 5. Lower �ronian. Ammonites fro"fu the Gloriaceras correai-Pro­
_
.

texamte � c.uca1taense-Codazzzceras scheibei Assemblage zone. Figs. 1-4 ' 7-10. Lo-
wer Comactan.
180 F. ETAYO · SERNA

PLATE 15

Fig.1) Codaz�iceras scheibei (Riedel), hypo­ Fig. 3) n. sp., holoty·

Protexanites cucaitaense
type C-134 (x 1 6/100). pP VL6·CU·3 (x 1 12/100).

Fig. 2) n. sp., ho·

Peroniceras robertogarcille Fig. 4) n. sp., holoty·
Pro texanites cucaitaense
loty� VL6, CU-5 (x 1 13/100). pe VL6-CU·3 (x 1 16/100).

Plate 1 5 . Ammonites from · the Gloriaceras correai-Pro texanite&· cucaitaense-Codazziceras

Assemblage zone. Figs. 1·4. Lower Coniacian.
scheibei
 183

INDEX O F GENERA AND SPECIES

Acan thoceras, 84. fig. 3, p. 120).

?ospinae (Karsten), 84. (�orph 1), 21 (Text. fig. 3, p.120).
A can thohoplites, 44. ._
Cortezzceras, n . g., 27.
abichi Anthula, 51. .-- cortezi, n . sp., 27, (Plate 3, p. 156), (Text.
aschiltaensis (Anthula), 44. fi g. 4, p. 1 22).
bigoti (Seunes), 4 7. · (morph 1), 28, (Plate 3, p. 156).
bigoure ti (Seunes), 4� la tecostata (Riedel), 28; (Plate 4, p. 158),
bigouretiforme, n . sp., 44, (Plate 5, p . 160), �Text. fig. 4, p. 122).
(Text. fig. 5, p.124). . n. sp., 2 9 , {Plate 3 , p.156), (Tex t . fig. 3,
p . 120) . ;
'derognati Roch', 46. .
eleganteante, n. sp ., 47, (Plate 5, p. 160), Cheloniceras, 32.
basseae Breistroffer, 38.
(Text. fig. 5, p. 124).
)
·
leptocerati(orme, n.sp., 47 ,(Plate 7, p . 164) clansayesense (Jacob , 36.
(Text. fig. 5 , p. 124).
·
aff. comuelianum (d Orbigny), 32.
luisperezi, n. sp., 4 7 , - (Plate 5, p. 160), . stoliczkanum (Gabb), 33.
(Text. fig. 5, p. 124). Desmoceras, 24.
odiosus, n. sp., 46, (Plate 4 , p.158), (Text. 'akuschaense An thula' 24.
fig. 5, p. 124). 'hyattum (Gabb)', 25.
pluricostatum, n . sp., 46, (Plate 7 , p.164), 'sparsicosta Boese', 25.
(Text. fig. 5, p.124). Discoceras, 9 1 .
quitasuegno, n. sp., 45, (Plate 7. p. 164), cf. largilliertianus (d'Orbigny), 9 1 .
(Text. fig. 5, p. 124). Donjuaniceras, 94.
rochi Breistroffer, 45. Douuilleiceras, 53.
serpen tiforme, n. sp., 49, (Plate 3, p. a bozagloi, n . sp., 55, (Plate 8, p. 166).
156), (Text. fig. 5, p. 124). · albense Spath, 56.
seunesi (Jacob), 45. (
mammilla tum Schlotheim), 54, 57.
'martini d'Orb. , 35 .
·

seunesi(orme, n . sp., 45, (Plate 7 , .164),

·

(Text. fig. 5 , p.l24). monile (J. Sowerby) , 54, 56.

A coneceras, 22. multinodosum Rhein, 54.
gu tierrezpalmae, n.sp., 22, (Plate 6 , p.162), solitae (d'Orbigny), 54, (Pl ate 7, p.164),
(Text. fig.3, p.120). (Text. fig. 6 , p.1 26). . ·
nisoides (Sarasin), 23. tarapacaense, n. sp., 54 (Plate 7, p. 164;
nisus (d'Orbigny), 23. Plate 8, p. 166), (Text. fig. 6, p . 126).
Barroisiceras, 97, 99. Dufrenoyia, 58.
a lstadenense Solger, 101. bo teroi, n.sp., 58, (Plate 1, p.152), (Text.
fig. 5, p. 124). .
subtubercula tum (Gerhardt), 1 02.
Benueites, 86. codazziana (Karsten), 58, (Plate 1, p.152).
spinosus Reyment, 86. discoidalis Casey , 60.
Brancoceras, 7 9 . durangensis Humphrey, 60.
? carinatum Collignon, 7 9 . formosa insculp ta Casey , 59.
Buenoceras, n . g., 101. furcata (J. de C. Sowerby) , 58.
loboi, n. sp., 101, (Plate 14, p.178), hansbuergli, n.sp., 60, 61, (Plate 1, p . 152),
(Text. fig. 9 , p. 132). (Text. fig. 14, p. 142).
Burckhardites, 67. justinae (Hill), 59.
pa/umbes Humphrey, 67. media Buergl , 60.
Carloscaceresiceras, n.g., 25. · mulatoensis Humphrey, 59.
caceresi, n. sp., 25, 26, (Plate 10, p.170), no tha Casey, 60.
(Text. figs. 4, p. 122; 10, p. 134). sanctorum Buergl, 59, (Plate 1, p . 152),
? chimuense (Be navides-Caceres), 27, (Pla­ (Text. fig. 14, p . 142).
(morph 1), 60, (Plate I, p.152), (Text. fig.
te 1 1 , p. 172), (Text. fig. 4, p. 122).
mon teroi, n . sp ., 26, (Plate 10, p. 170), 14, p . 142).
(Tex t. fig. 4, p. 122; 10, p.134). scalata Casey, 58.
Codazziceras, n .g., 83. texana Burckhardt, 61.
(ina, n. sp., 84, (Plate 13, p. 176), (Text. Eodouuilleiceras, 56.
fig. 8, p. 1 30). horridu m (Riedel), 57.
scheibei (Riedel), 83, (Plate 15, p. 180), pedrocarvaja/i, n. sp., 57, (Plate 8, p . 166),
(Text. fig . 8, p. 130). (Text. fig. 6, p . 1 26).
stutzeri (Riedel), 85. tequendamai, n. sp., 57, (Plate 2, .p. 154),
,
Colombiceras, 49 69, 70. (Text. fig. 6, p. 126; 13, p. 140).
caucasicum Luppov, 49, 70. Eogaudryceras, 19 .
crassicostatus (d'Orbigny), 49. helmsi, n. sp., 19, (Plate 6, p. 162), (Text.
(oreroi, n. sp., 5 0, (Plate 6, p.162), (Text. fig. 3, p. 120).
fig. 6, p.126). numidum var. in termedia (Fallot), 20.
sarmien toi, n. sp., 50, (Plate 6, p. 162), vocon tian um (Fallot), 20.
(Text. fig. 6, p. 126). Eote tragonites, 20.
tobleri (Jacob), 50, 5 1 . cabaricoi, n. sp., 20, (Plate 6, p . 162),
Colombocera toides, ri. sg., 20. (Text. fig. 3, p. 120).
renzom,. n. sp., 20 (Plate 6, p.162), (Text. jacobi am bigua Breistroffer, 20.
184

Engonoceras, 74. lagiraldae, n . sp . , 92, (Plate 13, p. 1 76) .

duartci, n . sp., 75, (Plate 12, p. 174). l.ltesellatus von Koenen , 89, 92.
gibbosum Hyatt, 75. munieri Pervinquhhe, 89.
o /galuciae, n. sp., 74, (Plate 1 1, p. 172), sp. ind., 92, (Plate 13, p. 176).
(Text. fig. 8, p. 130). Holcophylloccras, 17.
pierdenalis (von Buch), 7 4. gue t tardi (Raspail ), 18.
s tolleyi Bohm, 75. hernandezi, n.sp., 17, 18, (Plate 8, p .166),
sp., 75, (Plate 1 1, p. 172). (Text. fig. 3, p. 120).
uddeni (Cragin), 75. Hypacanlhoplites, 4 1 .
Epicheloniceras, 32. dognaliciae, n. sp., 4 1 , (Plate 3, p. 156).
a mazonaru m (Burckhardt), 33, (Plate 2, plesiotypicus (Fri ttel ), 4 1.
p. 154), (Text. fig. 4, p. 122). Hypophylloceras, 18.
barreroi, n. sp., 36, 58, (Pl ate 2, p. 154), su bserecitense Wiedmann, 19.
(Text. fig . 4, p. 122). wiedmanni, n . sp., 18, (Plate 10, p. 170),
carlosacostai, n . sp., 34, (Plate 2, p. 154), (Text. fig. 3, p. 120).
(Text. fig . 4, p. 122; 1 1 , p. 136), Hysteroceras, 79.
camachoi, n. sp . , 33, (Plate 2, p. 154), d 'orbigny Spath, 79.
(Text. fig. 4, p. 122). Imlayiceras, 88.
debile Casey , 33. ralphimlayi, n. sp., 88, (Plate 13, p. 176),
dognaelvirae, n . sp., 32, (Plate 2, p. 154 ), (Text. fig. 8, p. 130).
(Text. fig. 4, p. 122). washbournei, 88.
h uertasi n. sp . , 37, (Plate 2, p. 154), Jaumaportaiceras, n. sg., 31.
. (Text. fig. 4 , p. 122). ho{{stetteri, n. sp., 3 1 , (Plate 10, p. 170),
jimenoi, n. sp., 35, (Plate 2, p. 154), (Text. fig. 4, p. 122).
(Text. fig. 4, p. 122). Juandurhamiceras, n. g. 42.
leonhardtriedeli, n . sp., 37, 58, (Pl. 6, p. giraldoi, n. sp ., 43 (Plate 3, p. 156).
1 62), (Text. fig. 4, p. 122). joepechi, n. sp ., 43, (Plate 4 , p. 158).
pardoi, n. sp., 37 , (Plate 2, p. 154), (Text. juandurhami, n.sp., 42, (Plate 3, p.156;
fig. 4, p. 1 22). Plate 4, p. 158), (Text. fig. 5 , p. 124;
subnodosoc()sla tum ( Sinzow), 35. Text. fig. 1 2 , p. 138).
Eubrancoceras, 79. Knemiceras, 77.
aegocera toides (Steinmann), 79. pegnai, n . sp., 77, (Plate 12, 174), (Text.
Fagesia, 89. fig. 8, p. 138). .
peroni colombiana Fritsche, 90. raimondi Lisson, 78. .
superstes (Kossmat), 89. raimondi tardum Benavides-Caceres, ·78.
lhe vestensis (Peron ) , 90. semicostalum Sommermeier, 77, (Plate
zanellai, n. sp., 89, (Plate 13, p. 17 6 ; 14, 1 1 , p. 172), (Text. fig. 8, p. 130).
p. 1 78), (Tex t. fig. 8, p. 1 30). syriacus (von Buch ), 77.
Forresteria, 99. _
Lewesiceras, 30 .
. ,, cf. allaudi (Boule, Leomoine, Thevenine), ubalense Leanza, 88.
.. .. . 99 . ulloai, n. sp ., 30 (Plate 13, p . 176), (Tex t .
.
.-· (orresten· Reeside, 100. fi� 4 , p . 122 �
Frariciscoites, n. g., 87. Lyelliceras, 79.
· suarezi, n . sp., 87, (Plate 13, p. 176), carvajalorum, n.sp., 79, (Plate 1 1, p. 172),
· (Text. fig. 8, p.130). (Text. fig. 8, p. 130).
toroanus (Karsten), 87. isaacleai, n. sp., 80, (Plate 1 1 , p. 172).
Gargasiceras, 5 1. lyelli (Deshayes ), 79 .
gargasensis (d'Orbigny), 51. pseudo!yelli (Parona & Bomarelli), 80.
in teriectum (Riedel), 51, (Plate 1 , p.152), pseudolyelli{orme, n.sp., 80, (Plate 9, p.
· (Text. fig. 6, p. 126). 168l,. (Text. fig. 8, p. 1 30).
.· (morph 1), 52, (Plate 6, p. 162), (Text. Macroscaphiles, 49 .
fig. 6. p . 1 26). Magne ticeras, 31.
juanwyalti, n. sp., 53, (Plate 5, p. 160), magne ti Collignon, 31.
(Text. fig. 5, p. 124). Mammites, 85.
·
pulcher (Riedel), 5 3. {ugax, n.sp., 86, (Plate 13; p. 176), (Text.
Glonaceras, n . g., 96. fig. 8, p. 130).
. ,.,_.
.
correai, n. sp. , 96, 97, (Plate 13t p. 176; nodosoides (Schlotheim), 85. .
�- ·· . Pia � 14, p. 178), (Text . . fig. 9 ,:'P· 132). nodosoides a{ra Pervinquh�re, 86.• .·
··
paulmae, n. sp., 98, (Plate 13, p. 176), nodosoidesappelatus, n.sp., 85, (Plate 13,
··

(Text. fig. 9, p. 132). p. 140). .

Hem inau tilus, 1 04. revelierianum Coustillier, 87.
etheringtoni Durham, 104 , (Plate 4, p. scutulatus, n. sp ..t 87, (Plate 13, p. 1 76);
158; Plate 5, p. 160). (Text. fig. 8, p. 1010 ) .
saxbii (Morris), 104. Melchiorites, 23.
Hoplitoides, 89, 90. media Riedel , 23, 24, {Plate 3, p. 156),
inca Benavides-Caceres, 89. (Text. fig. 3, p. 120).
ingens (von Koene), 9 1 . melchioris (Tietze), 23.
ingens laeuis (Solger), 91. palmeri, 23, (Plate 6, p. 162), (Text. fig.
hernanmojicae, n .sp., 90, (Plate 13, p.176). 3, p. 120). -., .
" lacabagnae, n.sp., 91, (Plate 13, p. 176). Neodeshayesites, 62, 64 .
 1 85

albertoalvarezi, n. sp., 63, 64, (Plate 8, acu tospinatum (Basse ), 96.

p . 166; Plate 9 , p. 168), (Text. fig. 6,
p. 126; Text. fig. 16� p. 146).
cingula tum, n. sp., 66, (Plate 10, p.170),
(Text. fig. 7, p. 128).
colu mb ianus (Riedel), 62, (Plate 9, p. 168;
Plate 10, p. 170);''"(Text. fig; 6, p. 126;
Text. fig. 15, p. 144).
con tracta (Riedel), 66, (Plate 9, p. 168),
(Text. fig. 7, p. 128).
·
cucailaense, n. sp., 96, (Plate 14, p. 178),
(Text. fig. 9, p. 132).
guyabanus (Steinmann), 94.
lenti (Gerhardt), 96.
longispinalum (Basse), 94 , (Plate 13,
p. 176), (Text. fig. 9, p. 132).
medio tubercula tus (Gerhardt), 95.
portarum, n. sp., 95, (Plate 13, p. 176;
Plate 14, p. 178), (Text. fig. 9, p. 132).

karsleni (Marcou), 64, (Plate 9, p. 168; Prodeshayesites, 62.

Plate 10, p. 170), (Text.: fig. 6, p. 126; Prolyelliceras, 81.
Text. fig. 17, p. 148). geureyi (Jacob, 82.
(morph 1), 66, (Plate 10, p. 170)� (Text. loba tum Riedel, 8 1 .
fig. 1 7 , p. 148).
Neoptychites, 87. peru uianum Spath, 81.
Nice{oroceras, 9 2. Pro lacanthoceros, 83.
boyacaense, n. sp., 92, (Plate 14, p. 178). Pro tacan thoplites,5 1.
colu mbianus Basse, 92. Pro tengonocera.�. 76.
umbulazi{orme Basse, 92. gabbi (Bohm), 76.
Parahoplites, 72, (Plate 5, p. 160). Protexanites,97, 102.
hubachi, n. sp., 72 (Plate 5, p. 160). bourgeoisi (d'Orbigny), 102.
(morph 1), 73, (Plate 5, p. 160), (Text. cagnaense (Gerhardt), 102.
fig. 5 , p. l24).
cucaitaense,n. sp., 102, (Plate 15, p. 180);
incons lans Riedel, 74, (Plate 10, p. 170),
(Text. fig. 5, p. 124). (Text. fig. 9, p. 1 32).
melchioris Anthula, 72. Pse udohaploceras, 25.
mullicostatus Sinzow, 73. leiuaense, n.sp., 25, (Plate 3, p. 1 56),
nicholsoni Benavides-Caceres, 70. (Text. fig. 3, p. 120).
numidicus Somay, 74.
Pseudoptycho.ceras, n. g., 21.
tristan, n. sp., 73, (Plate 5 , p. 160),
gilberti, n. sp., 21, (Plate 6, p. 162), (Text.
(Text. fig. 5, p. 124).
Paramammites, 90. fig. 3, p.120).
colo mbianus, n. sp., 90, (Plate 13, p. 176). (morph I), 22, {Plate 6, p. 162).
gracilis (Pervinquiere), 90. Pseudosaynella, 29 .
po/ymorphus (Pervinquil�re), 90. bicurvata (Michelin), 30.
Parengonoceras, 75. fim briata Imlay, 30.
ebrayi (de Lorio!), 78. ralphimlayi, n.sp., 29 (Plate 3, ·p : 156), ·
elegans Renz, 75. (Text. fig. 4, p.122).
Paroxy tropidoceras, 79. walco tti (Hill), 30.
Pedioceras, 5 3.
Puzosia, 24.
Peroniceras, 92.
angladei Sayn, 24.
czornigi de Grossouvre , 97.
diabloi, n. sp., 93, (Plate 14, p. 178), Ralphimlayites, n.g. 8 1.
- apuloense, n. sp., 81, 82.
(Text. fig. 9, p. 132).
dravidica Kossmat, 97. prorsocurua tum (Gerhardt), 81, &2.
guerrai, n.sp., 94, (Plate 14, p.178). (Text. Reesidites, 102.
fig. 9, p. 1 32). minimus (Hayasaka and Fukada), 102.
leei Reeside, 97. .'?.iedelites, n. g., 70.
moureti de Grossouvre, 93. esthersernae, n . sp., 70 (Plate 6, p. 162),
robertogarciae, n. sp., 93, (Plate 15, (Text. fig. 7, p. 128).
p. 1 80), (Text. fig. 9, p . 1 32). filosum, n. sp., 7 2, (Plate 7, p. 164),
aff. rousseauxi de Grossouvre, 93. (Text. fig. 7, p. 128).
sub tricarinatum, Sturm,98, 99.
westfallicum de Grossouvre, 94. obliquum (Riedel), 70, 71.
Pegnaceras, 5 3. quebradanegra, n.sp., 70, (Plate 6, p.162);
rursiradiatus (Humphrey), 53. (Text. fig. 128).
Phylluceras, 17. Rinconiceras, n. g, 78.
buchianum Forbes ?, 17, (Plate 8, p.166), Rinconi, n. sp., 78, (Plate 9, p. 168),
(Text. fig. 3, p. 120). (Text. fig. 8, p. 130).
moriezense Sayn, 17 . Schloenbachia, 101.
serecitensis (Pervinquiere), 18.
flexuosa Gerhardt, 101.
velledae (Michelin), 17.
Stoyanowiceras, 67 .
Pla tiknemiceras, 76.
bassei Bataller, 76. cin tatum, n. sp., 69, (Plate 7, p. 164).
colo mbiana, n. sp., 76, (Plate 1 1 , p. 172). colo mbiana (Scott), 7 0.
gracile (Douville), 76. tref{ryanus (Karten), 67, (Plate 4, p.158 ;
sp. ind. A, 76, (Text. fig. 8, p. 130). Plate 5, p. 160), (Text-fig. 7, p. 128).
sp. ind. B, 76, (Plate 12, p. 174). (morph I), 69, (Plate 3, p.156).
Prionocycloceras, 94. (morph II), 69, (Plate 3, p. 156).
186

Tegoceras, 82. colombiana, n. sp., 24, (Plate 3, p. 156),

benavidescaceresi, n . sp., 82, (Plate 1 1, (Text. fig. 3, p. 120).
p. 172), (Text. fig. 8, p. 130). Vascoceras, 89.
cama tteanum (d'Orbigny), 82. Ve ctisites, 38.
seunesi (Parana & Bonarelli), 83. Zambranoites, n. sg., 38.
Texanites, 83. cadenai, n . sp., 40, (Plate 6, p. 162), (Text.
aff. serratomaginatus (Redtenbacher), 83. fig. 5 , p. 124).
caprotinus Casey, 38, 40.
Thomasites, 87. ·
cruz� n . sp., 39, (Plate 6, p. 162), (Text.
Toxoceratoides, 20.
fig., 5, p. 124).
royerianum (d'Orbigny), 20. duquesi, n. sp., 40, (Plate 6, p. 162), (Text.
Trochleiceras, 31. fig. 5, p. 124).
ambanjabense Collignon, 32. mateusi, n. sp., 39, (Plate 6, p. 162).
balearense Fallot & Tennier, 31. zambranoi, n. sp., M, (Plate 6, p. 162),
juliverti, n. sp., 31. (Text. fig. 5, p. 124).
Uhligella, 27. Zumpangoceras, 99.
latecostata Riedel, 28. burckhardti Basse , 99.
reesidei Humphrey , 28. ospinai, n. sp ., 99, 100, (Plate 14, p. 178),
(Text. fig. 9, p. 132).
zurcheri Jacob & Tobler, 27. sorae, n. sp., 101, (Plate 14, p. 178).
Valdedorsel/a, 24.

* * *