Cretaceous Research 26 (2005) 450e459

www.elsevier.com/locate/CretRes

Late Middleeearly Late Albian ammonites from Ecuador

L.G. Bulot a,*, W.J. Kennedy b, E. Jaillard c, E. Robert d
a
UMR CNRS 6019, Centre de Sedimentologie-Paleontologie, Universite de Provence, Place V. Hugo, 13331 Marseille Cedex, France
b
Geological Collections, University Museum of Natural History, Parks Road, Oxford OX1 3PW, UK
c
IRD Equateur, Apartado Postal 17-12-857, Quito, Ecuador
d
EA 3029, Laboratoire de Dynamique des Bassins sedimentaires, Universite de Toulouse III,
39 Allees J. Guesde, 31062 Toulouse Cedex, France
Received 21 September 2004; accepted in revised form 24 January 2005
Available online 21 June 2005

Abstract

Ammonites of the Albian genera Brancoceras, Dipoloceras, Mortoniceratoides and Neophlycticeras are described and illustrated
for the rst time from Ecuador. Precise stratigraphic distribution in the Rio Misahuali eld section allows delineation of the Middle/
Upper Albian boundary in the Oriente of Ecuador through the recognition of the Dipoloceras cristatum Zone.
2005 Published by Elsevier Ltd.

Keywords: Ammonites; Cretaceous; Albian; Ecuador; Subandean Zone

1. Introduction Western Europe and the south-western USA. The

remainder of the fauna (mainly Oxytropidoceras and
The occurrence of Lower Cretaceous ammonites in engonoceratids) is only briey mentioned in the
Ecuador was rst reported by Watson and Sinclair stratigraphic description of the sections studied; they
(1927) and Tschopp (1953) on the basis of faunas will be described elsewhere, together with their Peruvian
collected from the Napo Formation and identied and Colombian counterparts.
respectively by J.B. Reeside Jr., and M. Breistroer.
Very few of these ammonites have been gured, and
none described; some additional material was sub- 2. Geologic and stratigraphic setting
sequently noted by Aspden and Ivimey-Cook (1992).
The aim of the present paper is to revise the The area studied belongs to the Andean active
descriptions and determinations of specimens previously margin that has experienced the subduction of the
identied by Breistroer in Tschopp (1953) and palaeo-Pacic oceanic plate since the Jurassic (Jaillard
document some interesting new nds made by one of et al., 1990). During the Cretaceous Period, various
us (EJ) during the course of a detailed eld study of the palaeogeographic areas can be distinguished on this
Napo Group of the Oriente of Ecuador (Fig. 1). margin (Fig. 1).
Emphasis is laid on cosmopolitan taxa that allow One of these is the East Ecuadorian (or Oriente)
a correlation with the standard ammonite scales of Basin that underlies large areas of eastern Ecuador to
Brazil and extends towards Columbia in the north and
Peru to the south. It was characterised by low sub-
* Corresponding author. sidence rates and marine sedimentation during most of
E-mail address: lucgbulot@aol.com (L.G. Bulot). the Cretaceous (Caneld et al., 1982). Two dierent

0195-6671/$ - see front matter 2005 Published by Elsevier Ltd.

doi:10.1016/j.cretres.2005.01.008
 L.G. Bulot et al. / Cretaceous Research 26 (2005) 450e459 451

zones can be distinguished: (1) the Subandean Zone

where Mesozoic outcrops allow eld study of the
Cretaceous successions presented herein; (2) the Ama-
zonian Zone where marine Cretaceous rocks are only
known through well data.
The rst survey of the Subandean Zone was carried
out by Watson and Sinclair in the course of 1921 and
their main geological results were published in 1927. The
stratigraphic nomenclature of the area studied is still
very much derived from their early work and subsequent
syntheses by Shell geologists (Tschopp, 1953; see also
discussion in Jaillard et al., 1997).
The ammonites described below were collected
from the lower part of the Napo Group whose nomen-
clature has recently been rened by Jaillard et al. (1997,
pp. 48e54, g. 2). It rests on the sandstone of the Hollin
Formation and is overlain unconformably by the basal
conglomerates of the Tena Formation. As here un-
derstood, the Napo Group is subdivided into the Basal,
Lower, Middle and Upper Napo formations. It ranges
in age from the Late EarlyeEarly Middle Albian to
Early Coniacian (Jaillard et al., 1997).
Our material was collected from the Basal and Lower
Napo formations. Two main areas have been studied in
great detail; the Rio Misahuali and Chinimbimi sections
where ammonites, bivalves, echinoids, microfaunas and
microoras have been collected bed by bed (Fig. 2).
Preliminary results on the biostratigraphy of the
MiddleeUpper Albian of the Chinimbimi section were
published by Robert et al. (1998, 2002). Because most of
the signicant specimens described herein were obtained
at Rio Misahuali, we focus on that section.

3. Field section

The Rio Misahuali section is located south of the

village Tena, and lies on the sides of the Misahuali river
close to its conuence with the Napo River (Fig. 1). The
exposures are good to excellent and quite easily
accessible for the area. It provides a reference section
for the northern part of the Subandean Zone.
No ammonites have been collected from the lower-
most part (basal sandstones and C limestones) of the
Basal Napo Formation. The oldest ammonites (sample
M.94.9/10) were collected from the black laminated
shales rich in crushed ammonites and bivalves that mark
the base of the basal Napo Shales. Besides numerous
Brancoceras (Brancoceras) sp., the fauna also includes
rare Venezoliceras of the venezolianum group and many
buchiids (Aucellina sp.). Comparison with equivalent
Fig. 1. Geological and palaeogeographical map of the South American faunas in Peru (Robert, 2002) and elsewhere in the
Andean margin and location of the area studied. world suggests a middleelate Middle Albian age. This
view is also supported by the occurrence of numerous
Oxytropidoceras (O.) carbonarium in the C limestones
of the nearby Pungaracayu-30 well.
452 L.G. Bulot et al. / Cretaceous Research 26 (2005) 450e459

Fig. 2. Lithologic log and biostratigraphic interpretation of the Misahuali eld section.

The most interesting ammonite fauna was collected such as G. sommermeieri and G. moorei. For the reason
from the nodule beds (M.94.12) at the top of the basal outlined below (see Systematic palaeontology)
Napo shales. It includes Dipoloceras (D.) tarrentense, D. this level is taken to mark the base of the Upper
(D.) a. fredericksburgense and species of Glottoceras Albian (Dipoloceras cristatum Zone). The occurrence of
 L.G. Bulot et al. / Cretaceous Research 26 (2005) 450e459 453

Actinoceramus concentricus (Parkinson) (determined by 1999 Dipoloceras (Dipoloceras) tarrantense Scott, 1928;
A. Dhont) at that level is also noteworthy. Kennedy in Kennedy et al., p. 1107, gs. 4.12e
Higher up the succession ammonites are scarce. A 4.14, 7.1e7.7, 7.10e7.12, 10.7.
large fragment of Oxytropidoceras (Benavidesites) kar-
steni Stelier was collected from a hardground surface Type. The holotype, by monotypy, is the original of
(M.94.15) in the middle of the T limestones. Accord- Scott, 1928, p. 112, pl. 16, gs, 1, 2; text-g. A3, from
ing to Renz (1968, 1982), this species is fairly common in the lower Upper Albian Goodland Limestone, 6.1 m
the uppermost part of the La Puya Member of the Penas (20 ft) below the top, 3.5 km (2 miles) north of a point
Altas Formation of Western Venezuela, where it co- where the White Settlement road crosses the Tarrant/
occurs with ammonites characteristic of the Mortoni- Parker County line.
ceras inatum Zone (Hysteroceras orbignyi Subzone
according to Renz, 1982; Hysteroceras varicosum Sub- Material. FSM 94.12.1 is from bed M.94.12 (upper-
zone according to Owen, 1999). most part of the Basal Napo Formation). Another less
The youngest Albian ammonites found at Misahuali well-preserved specimen (FSM 94.12.3) was collected
are from lower part of the B limestones (Lower Napo from the same bed.
Formation). Both specimens (M.94.16 and M.94.19a)
are poorly preserved mortoniceratids, one of them Dimensions D Wb Wh Wh:Wh U
showing close anities with Deiradoceras. The lower FSM 94.12.1 126.0 46.9 43.6 1.08 56.0
(100) (37.2) (34.6) (44.4)
part of the Lower Napo Formation therefore still Costal dimensions 78.9 65.3 28.9 2.26 32.7
belongs to the Upper Albian Mortoniceras inatum (100) (82.7) (36.6) (50)
Zone.
Description. FSM 94.12.1 is a complete adult. Orna-
4. Systematic palaeontology ment of the adapertural 120  sector of the shell and the
previous whorl are utterly distinctive. Prior to adult
Repositories. The specimens mentioned below are modication, the coiling is very evolute, with a wide, deep
housed in the following repositories: FSM, Faculte umbilicus. The umbilical wall is notched to accommodate
des Sciences, Marseille; BMNH, The Natural History the ared ribs of the previous whorl. The whorl section is
Museum, London; MNHP, Museum National dHis- very depressed oval in intercostal section; the costal whorl
toire Naturelle, Paris. section is even more depressed, with a whorl breadth to
height ratio of 2.26 through the ared ribs. There are nine
Dimensions. All dimensions are given in millimetres. massive ared horns per whorl at this stage of de-
D, diameter; Wb, whorl breadth; Wh, whorl height; velopment. Each horn bears a strong narrow rib at the
U, umbilicus. Figures in parentheses are dimensions as adapertural and adapical edge on the anks and venter,
a percentage of diameter. which is deeply sunken between the ares. Between the
ared horns are one or two low, narrow prorsiradiate
Order: Ammonoidea Zittel, 1884 primary ribs that sweep forwards across the ventrolateral
Suborder: Ammonitina Hyatt, 1889 shoulder to dene a very obtuse chevron, interrupted by
Superfamily: Acanthocerataceae de Grossouvre, 1894 a coarse, blunt siphonal keel and anking grooves. The
Family: Brancoceratidae Spath, 1934 last two ared horns are signicantly weaker than the
Subfamily: Mojsisovicziinae Hyatt, 1903 others on the outer whorl, and are separated by two coarse
Genus and Subgenus Dipoloceras Hyatt, 1900 single ribs, with a third linked to the nal are.
Adapertural of the last are there are 13 strong, coarse,
Type species. Ammonites cristatus Brongniart, 1822, narrow ribs of variable strength and elevation. They are
p. 395, pl. 7, g. 9, by original designation of Hyatt, strongly prorsiradiate and markedly concave on the
1900, p. 589. anks, and directed backwards on the venter in a very
obtuse chevron divided adapically; there is a coarse
Dipoloceras (Dipoloceras) tarrantense Scott, 1928 siphonal keel. A pair of ribs arise from the umbilical
Figs. 3, 4 shoulder immediately adapical of the aperture. The
sutures are not seen.
1928 Dipoloceras tarrantense Scott, p. 112, pl. 16, gs. 1,
2; text-g. A3. Discussion. The remarkable development of ared
1928 Dipoloceras tarrantense Scott; Adkins, p. 224. horns, occupying all of the anks and rising above the
1931 Dipoloceras tarrantense Scott; Spath, p. 363; text- venter, characterises the species. At this growth stage
g. 118aec. the specimen closely resembles in gross morphology
1994 Dipoloceras tarrantense; Emerson et al., table 48 the Texas example gured by Kennedy in Kennedy
on p. 117. et al. (1999, g. 4.12e4.14), diering only in the more
454 L.G. Bulot et al. / Cretaceous Research 26 (2005) 450e459

Fig. 3. A, B, Dipoloceras (Dipoloceras) tarrantense Scott, 1928. Side and ventral views of FSM M94.12.1; both !1.

numerous (three versus one or two) ribs between ared cf. 1931 Rhytidoceras elegans van Hoepen, p. 43, text-
horns. The dramatic change in ornament on the nal gs. 4e7.
120  sector of the shell before the adult aperture is cf. 1941 Rhytidoceras elegans van Hoepen; van Hoepen,
highly distinctive and shows the previously undescribed p. 64, gs. 14e18.
morphology of the adult shell. cf. 1941 Rhytidoceras crassicostatum van Hoepen, p. 67,
text-gs. 19, 20; pls. 9, 10.
Occurrence. Dipoloceras tarrantense was previously
known only from the lower Upper Albian upper part of Type. The holotype, by original designation, is the
the Goodland/Comanche Peak Limestone in Tarrant and original of van Hoepen, 1931, text-g. 4; 1941, text-g.
Parker Counties, Texas, where it co-occurs with Dipolo- 17, from the lower Upper Albian of the Mzinene River,
ceras cristatum. In Ecuador, both specimens have been Zululand, South Africa.
collected from the uppermost part (nodule beds) of the
Basal Napo Shales (Napo Formation sensu Jaillard Material. FSM KT.6.427, from Rio Longota (Mis-
et al., 1997). ahuali) (Basal Napo Formation, exact horizon unknown).
Subgenus Rhytidoceras van Hoepen, 1931 Dimensions D Wb Wh Wb:Wh U
FSM KT.6.427 160.0 43.2 50.0 0.84 67.0
Type species. Rhytidoceras elegans van Hoepen, 1931, (100) (27.0) (31.3) (41.0)
p. 43, text-gs. 4e7, by original designation of van
Hoepen, 1931, p. 43. Description. Coiling is very evolute; the broad
shallow umbilicus comprises 41.9% of the diameter.
Dipoloceras (Rhytidoceras) a. elegans (van Hoepen, The low umbilical wall is feebly convex, the ventrolat-
1931) eral shoulders broadly rounded. The whorl section is
Fig. 5 compressed oval, with a whorl breadth to height ratio
 L.G. Bulot et al. / Cretaceous Research 26 (2005) 450e459 455

ventral rib-endings of D. (R.) elegans are not so-

dierentiated. In this respect, the specimen is transi-
tional to early Mortoniceratinae.

Occurrence. The most closely allied forms of Dipolo-

ceras (Rhytidoceras) occur in the lower Upper Albian of
Zululand, South Africa, where they rst appear together
with Dipoloceras (D.) cristatum, but range above. In
Ecuador the one and only specimen was collected by
Shell geologists and we lack detailed data on its precise
stratigraphic position. Nevertheless, the preservation
diers very much from most of the ammonites described
herein, with the exception of O. (B.) karsteni from the
T limestones (the worm surface is encrusted by
serpulids and the matrix includes glauconite and phos-
phatic pebbles). We accordingly suggest that the specimen
originates from one of the hardground surfaces that
characterise the T limestones.

Subfamily: Brancoceratidae Spath, 1934

Genus and subgenus Brancoceras Steinmann, 1881

Type species. Ammonites senequieri dOrbigny, 1841,

p. 292, pl. 86, gs. 3e5, by subsequent designation of
Hyatt, 1900, p. 590.

Brancoceras (Brancoceras) sp.

Fig. 6F

Material. FSM M.94.10.4, from bed M94-10 of the

Mishahuali section (lowermost part of the Basal Napo
Formation).

Fig. 4. Dipoloceras (Dipoloceras) tarrantense Scott, 1928. Apertural Description. The specimen is preserved as a crushed
view of FSM M94.12.1; !1 (see also Fig. 3). lm, lacking shell, deformed into an ellipse with a major
diameter of 26 mm. Coiling is very evolute, serpenticone,
the wide umbilicus comprising an estimated 38% of the
of 0.84, the greatest breadth just outside the umbilical diameter. The whorls expand very slowly. Ribs arise
shoulder. The inner and middle anks are broadly either singly or in pairs from the umbilical shoulder, and
convex, the outer anks convergent, ventrolateral appear to have been recti- to feebly rursiradiate, exing
shoulders and venter broadly rounded. Thirty-six backwards on the outer ank and thickening on the
strong, narrow, crowded primary ribs arise on the ventrolateral shoulder. There are an estimated 16 ribs on
umbilical wall and strengthen into feeble bullae, perched the outer half whorl of the specimen.
on the umbilical shoulder. The bullae give rise to ribs
either singly or in pairs. The ribs are feebly prorsiradiate Discussion. Although poor, the specimen can be
and weakly sinuous, concave on the inner to mid-ank, assigned to Brancoceras (Brancoceras), notably the type
slightly concave on the outer ank and ventrolateral species B. (B.) senequieri, as represented by a suite of
shoulder, and near-transverse on the venter. Although well-preserved specimens from the condensed Albian
the surface of the mould is corroded, there are traces of of Gourdon, Alpes-Maritimes, France, in the collections
weakly developed ventral bullae. of the Universite de Grenoble.

Discussion. The whorl proportions, coiling, density Occurrence. Brancoceras (Brancoceras) is a typical
and style of ribbing are those of Dipoloceras Early and Middle Albian genus. The present specimen
(Rhytidoceras), particularly the type species, D. (R.) and many other unlabelled juvenile specimens were
elegans (van Hoepen, 1931, p. 43, text-g. 47; 1941, collected from the lowermost part (black shales) of the
gs. 14e18). The specimen diers, however, in the Basal Napo Shale together with rare Venezoliceras gr.
very clearly developed ventral tubercles, whereas the venezolanum Stieler. Because of its stratigraphic position
 456
L.G. Bulot et al. / Cretaceous Research 26 (2005) 450e459
Fig. 5. AeC. Dipoloceras (Rhytidoceras) a. exuosum (van Hoepen, 1931). Apertural, side, and ventral views of FSM KT.6.427; !1.
 L.G. Bulot et al. / Cretaceous Research 26 (2005) 450e459 457

Fig. 6. A, B, Neophlycticeras brottianum (dOrbigny, 1841), side and ventral views of FSM G350. CeE, GeI, Mortoniceratoides rigidum (Spath,
1933); C, E, side views, D, venter of FSM KTG 463; GeI, apertural, side and ventral views of the holotype, BMNH C. 34879. F, Brancoceras
(Brancoceras) sp., side view of FSM M.9410.4. All !1.

well below the level with Dipoloceras and association Material. FSM KTG 463 from the upper Rio
with V. gr. venezolanum, a middleelate Middle Albian Bueno (Basal Napo Formation, exact horizon un-
age is indicated. known).

Genus Mortoniceratoides Cooper, 1982 Dimensions D Wb Wh Wb:Wh U

47.2 17.4 20.0 0.87 14.6
(100) (36.9) (42.4) (30.9)
Type species. Mortoniceras (Pervinquieria) rigidum
Spath, 1933, p. 413, text-gs. 142, 144f, by original
designation of Cooper, 1982, p. 296. Description. FSM KTG 463 is a half whorl of body
chamber with a maximum preserved diameter of
Mortoniceratoides rigidum (Spath, 1933) 47.2 mm. Coiling is very evolute. The umbilicus
Fig. 6CeE, GeI comprises 30.9% of the diameter, and is of moderate
depth, with a attened, feebly convex wall. The
1933 Mortoniceras (Pervinquieria) rigidum Spath, p. 413, umbilical shoulder is quite narrowly rounded. The
text-gs. 142, 144f. whorl section is compressed trapezoidal in costal
1982 Mortoniceratoides rigidus (Spath); Cooper, p. 296, section, with the greatest breadth at the umbilical
text-g. 13e, f. shoulder. Fourteen ribs arise at the umbilical seam,
and strengthen across the umbilical wall and shoulder.
Types. Holotype, BMNH C. 34879; paratype, They are strong, narrow, and prorsiradiate on the
BMNH C. 35929, from Bed 8, the cristatum nodule anks. Single ribs are very feebly convex. Other ribs
bed of the lower Middle Albian Gault Clay at Folke- branch below mid-ank, with the adapical of the two
stone, Kent (Fig. 6GeI). secondary ribs either straight or with an initial
458 L.G. Bulot et al. / Cretaceous Research 26 (2005) 450e459

sinuosity, beyond which the two ribs of the pair are umbilicus. The whorl section is compressed, with the
subparallel. The ribs strengthen across the ventrolat- greatest breadth outside the umbilical shoulder, the
eral shoulder, and are transverse on the venter, inner anks broadly convex, the outer anks attened
terminating in an incipient ventral bulla. There is and convergent, the venter fastigiate. At a diameter of
a strong, continuous siphonal keel anked by prom- 30 mm there are 17 low, broad, exuous, prorsiradiate
inent grooves. ribs per half whorl. Primary ribs arise at the umbilical
seam, and strengthen across umbilical wall and
Discussion. The highly distinctive ank and ventral shoulder. They are straight on the inner ank, feebly
rib development of the fragment, notably the branching convex at mid-ank, exed back and concave on the
pattern and transverse ventral development, are identi- outer ank, and projected forwards and thickened
cal to those of the holotype (Fig. 6GeI). This is rather across ventrolateral shoulders and venter, where they
more coarsely ribbed than the present specimen, which are near-transverse, with small, sharp siphonal clavi.
can be matched with topotypes. Single intercalated ribs arise around mid-ank, and
have comparable ventral and ventrolateral develop-
Occurrence. Lower Upper Albian of southern Eng-
ment to the primary ribs. The outer half whorl is
land, associated with Dipoloceras (D.) cristatum. The
crushed, but shows the ribs coarsening and broadening
precise horizon of the specimen described herein is
markedly.
unknown but its preservation suggests that it originates
from nodule beds that occur at the top of the Basal Discussion. The present specimen diers in no
Napo Shales sensu Jaillard et al. (1997). As a conse- signicant respects from similarly compressed specimens
quence, this suggests that, as in England, the Ecuador- of N. (N.) brottianum discussed by Kennedy and
ian Mortinoceratoides occurs in the Dipoloceras Delamette (1994) and Matrion et al. (1998).
cristatum Zone together with other Dipoloceras species.
Occurrence. Where well-dated, N. (N.) brottianum is
Family: Lyelliceratidae Spath, 1921 early Late Albian in age, occurring in the Dipoloceras
Subfamily: Stoliczkaiinae Breistroer, 1953 cristatum to low Hysteroceras varicosum subzones of the
Genus and subgenus Neophlycticeras Spath, 1922 western European Mortoniceras inatum Zone. The
precise position of the specimen described herein is
Type species. Ammonites brottianus dOrbigny, 1841,
unknown but its preservation suggests that it originates
p. 290, pl. 89, gs. 8e10, by original designation of
from the nodule beds that occur at the top of the Basal
Spath, 1922, p. 107.
Napo Shales sensu Jaillard et al. (1997). The geographic
range is southern England, France, Switzerland, Tuni-
Neophlycticeras (Neophlycticeras) brottianum (dOrbigny,
sia, Morocco, Zululand (South Africa), Madagascar,
1841)
and the present record from Ecuador.
Fig. 6A, B

1841 Ammonites brottianus dOrbigny, p. 290, pl. 85, 5. Conclusions

gs. 8e10.
1994 Neophlycticeras brottianum (dOrbigny, 1841); The study of the Middle and Late Albian ammonite
Kennedy and Delamette, p. 7, text-gs. 3bed, fauna of the lower part of the Napo Group of the
5aec, gei, 6aes; 7aef; 8y, z (with full synonymy). Oriente Basin of Ecuador (Subandean Zone) sheds
1998 Neophlycticeras (Neophlycticeras) brottianum new light on the biostratigraphy and palaeobiogeogra-
(dOrbigny, 1841): Matrion, Dubus and Touch, phy of the area. New eld data show that the base of the
p. 19, pl. 1, gs. 1e3; pl. 2, gs. 1e5; text- Upper Albian lies at the top of the Basal Napo Shales
gs. 3, 4. sensu Jaillard et al. (1997) (Basal Napo Formation).
The Upper Albian ranges up to the top of the B
Type. Holotype, by monotypy, MNHP5757, dOr- Limestones of the Lower Napo Formation. Neverthe-
bigny Collection, the original of dOrbigny, 1841, pl. 85, less, our new collections do not provide any evidence for
gs. 8e10, from the condensed Albian of Perte du the uppermost part of the Albian (Stoliczkaia dispar
Rhone, Ain, France. Zone).
The fauna combines Andean (Glottoceras of the
Material. FSM G350-1 from Rio Coca (Basal Napo sommermeieri and moorei plexus) and cosmopolitan
Shales, exact horizon unknown). elements (mainly Mojsisovicziinae). Among the cosmo-
politan taxa, some genera and species are reported and/
Description. FSM G350-1 is a partially crushed or conrmed for the rst time. These include Neo-
individual with a maximum preserved diameter of phlycticeras brottianum, whose geographic range in
47 mm. Coiling is very involute, with a small deep the New World was hitherto restricted to Western
 L.G. Bulot et al. / Cretaceous Research 26 (2005) 450e459 459

Venezuela. Noteworthy also is the occurrence of Jaillard, E., Caron, M., Dhondt, A., Ordonez, M., et al., (10 other
Mortoniceratoides and Dipoloceras (D.) tarrantense, authors), 1997. Datos nuevos y discusion. In: Jaillard, E. (Ed.),
Sintesis Estratigraca y Sedimentologica del Cretaceo y Paleogeno
previously only known, respectively, from southern de la Cuenca Oriental del Ecuador. Informe Final del Convenio
England and Texas. Similarly, the subgenus Rhytido- Orstom-Petroproduccion, Orstom, Paris, 164 pp.
ceras, mainly known from Madagascar and South Kennedy, W.J., Delamette, M., 1994. Neophlycticeras Spath, 1922
Africa, is reported from the rst time in South America. (Ammonoidea) from the Upper Albian of Ain, France. Neues
Jahrbuch fur Geologie und Palaontologie, Abhandlungen 191, 1e24.
Kennedy, W.J., Gale, A.S., Hancock, J.M., Crampton, J.S.,
Cobban, W.A., 1999. Ammonites and inoceramid bivalves from
Acknowledgement close to the Middle-Upper Albian boundary around Fort Worth,
Texas. Journal of Paleontology 73, 1101e1125.
Matrion, B., Dubus, B., Touch, R., 1998. Le genre Neophlycticeras
For one of us (LGB) the research work developed (Spath, 1922) dans lAlbien de lAube et du Boulonnais. Bulletin
herein was supported by a grant from the Societe de Annuel de lAssociation Geologique Auboise 19, 15e28.
Secours des Amis des Sciences. dOrbigny, A., 1840e1842. Paleontologie francaise: Terrains cretaces.
1. Cephalopodes. Masson, Paris. 1e120 (1840), 121e430 (1841),
431e662 (1842).
Owen, H.G., 1999. Correlation of Albian European and Tethyan
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