GEOLOGICA CARPATHICA, JUNE 2005, 56, 3, 245254
The oldest (Late Valanginian) Crioceratitinae (heteromorphic
ammonoids) from the Central Western Carpathians
Institute of Geological Engineering, VB Technical University, 17. listopadu 15, 708 33 Ostrava-Poruba, Czech Republic;
(Manuscript received April 16, 2004; accepted in revised form September 29, 2004)
Abstract: An Upper Valanginian ammonoid collection from the Manín Unit (the Butkov Quarry of the Ladce cement
works, middle Váh Valley) obtained recently contains true Crioceratitinae and other related subcriocone ammonoid
shells belonging to six species of Crioceratites, Criosarasinella and Himantoceras. The occurrence of Crioceratites in
the ammonite furcillata Zone represents the first known Valanginian record of this genus from the Central Western
Carpathians. On the basis of the morphology of its early whorls, Criosarasinella is taxonomically assigned to the family
Ancyloceratidae, the same as Crioceratites and Himantoceras.
Key words: Lower Cretaceous, Central Carpathians, taxonomy, heteromorphs.
A large collection of well preserved ammonoids has been ob-
tained from Upper Valanginian strata of the Manín Unit dur-
ing fieldworks in the Butkov Quarry in the period from 2000
until 2003 (Skupien et al. 2003). The research done by a team
of Czech and Slovak paleontologists continued older field
studies by K. Borza, J. Michalík and the author of this article
during 19801995. The newly collected ammonoids enabled
more precise dating of the sequence studied up to the level of
ammonite zones and subzones used at present by Hoedemaek-
er & Reboulet (2003).
The taxonomic and stratigraphic knowledge on Lower Cre-
taceous ammonoids has been enhanced during the last decade.
Primarily, Reboulet (1996) described the French ammonoid
successions of the Vocontian Trough comprising Late Valang-
inian representatives of Crioceratites. Previous findings (be-
fore 1996) of the stratigraphically oldest Crioceratites speci-
mens were only inaccurately dated. This was the reason for
distrust about whether the first Crioceratites could have
evolved prior to the Early Hauterivian. However, Reboulet
(1996) unambiguously documented that this genus already ap-
peared during the Late Valanginian.
The ammonoid association from the Butkov Quarry also
comprises ribbed evolute shells (with whorls in contact) and
shells with slightly gyrocone up to gyrocone, or in older litera-
ture criocone shells (planispirally coiled shells in which the
whorls do not touch one another), including the first Criocera-
tites of Late Valanginian age.
The Manín Unit holds a special position in the geological
structural plan of the Western Carpathians. It is a nappe unit
incorporated into the Paleoalpine Accretionary Belt masking
the suture zone of both the Central and Outer Western Car-
pathians. The complicated geological setting of this nappe unit
resulted in a non-uniform manifold interpretations of its rela-
tion to the units of the Pieniny Klippen Belt and/or to the units
of the Central Carpathians. A brief survey of previous opin-
ions on the position and assignation of the Manín Nappe has
been compiled by Michalík & Vaíèek (1987).
During the Paleoalpine compression, the Manín Nappe was
thrusted along with the West Carpathian superficial nappes
system across the Tatric Superunit. Therefore, it is believed
that the original sedimentation basin of the Manín Unit was
situated along the western Fatric periphery (Vaíèek &
The Butkov Quarry yields the best exposures of the Meso-
zoic sequence of the Manín Unit. The uppermost Jurassic and
Cretaceous carbonates are exploited here as the raw material
for cement production in the Povaské cementárne cement
works, Ladce (Fig. 1). The Upper Jurassic and Lower Creta-
ceous sequence of the Manín Unit at Butkov, its microfauna
and Lower Cretaceous cephalopods have been studied by
Vaíèek & Michalík (1986), Borza et al. (1987), Michalík &
Vaíèek (1987), Michalík et al. (1990), Vaíèek et al. (1994),
Michalík et al. (1995), and Skupien et al. (2003).
The Butkov Lower Cretaceous sequence was divided by
Borza et al. (1987) into several lithostratigraphic units. The
uppermost part of the Ladce Formation (forming the basal
parts of the Lower Cretaceous sequence here) passes upwards
into the overlying Mráznica Formation. The fossils dealt with
in the paper submitted were collected just from these transi-
tional beds. The lithology of the rock sequence mentioned
along with the composition of the Upper Valanginian am-
monoid association collected on the 11
level of the Butkov
Quarry is presented in the Fig. 2.
Material preservation, abbreviations and symbols
The ammonoid shells are preserved as sculptural moulds;
however, they (especially the gas chambers of the fragmo-
cone) were strongly flattened. As a rule, the earliest whorls of
the shells corresponding to the shell diameter of about 7
10 mm are not observable.
On the well preserved shells, the following parameters were
D shell diameter; D
preserved maximum shell di-
ameter of the specimen measured (not the maximum of the
given species); Wh whorl height; Uw umbilicus width.
In brackets placed after the values measured in mm the calcu-
lated ratios, Wh/D and Uw/D, follow.
Particular specimens are designated by composite symbols
expressing: the symbol of the locality (BK); then the number
of the quarry level (BK11, or BK7Z), the number denoting
distance in meters on the section line after a hyphen (BK11-
26) and the serial number of the finding after broken line
(BK11-26/6). The specimens illustrated, or referred to in the
text, were deposited in the collections of the Slovak National
Museum in Bratislava (SNM). The Museum assigned deposi-
tory numbers to them (e.g. SNM Z 23555).
Suborder: Ancyloceratina Wiedmann, 1960
Superfamily: Ancyloceratoidea Gill, 1871
Family: Ancyloceratidae Gill, 1871
Subfamily: Crioceratitinae Gill, 1871
Genus: Criosarasinella Thieuloy, 1977
Type species: Criosarasinella furcillata Thieuloy, 1977
From the original generic diagnosis by Thieuloy (1977) it
follows, that the juvenile growth stage should be characterized
Fig. 1. Geographical and geological situations of the Butkov Quarry.
by trituberculate main ribs and the slightly criocone (= subc-
riocone) coiling of whorls. The trituberculate ribs disappear in
the medium stage. A typical feature is represented by frequent
rib bifurcation on the external side. Adult whorls are in contact.
Reboulet (1996), who differentiated microconchs from mac-
roconchs in all the species described by himself, emends par-
tially the original description especially by the fact that the
shells may be evolute in all growth stages.
The Slovak material represented by 3 species is character-
ized by trituberculate ribs on juvenile whorls. C. mandovi
Thieuloy and C. subheterocostata Reboulet have whorls in
contact for the whole time of growth; whereas one of favour-
able preserved specimens of C. furcillata has the last but one
whorl in the shorter part subcriocone. A similar situation may
be seen in one of Thieuloys specimens (1977: Pl. 5, Fig. 4) as
well. Early stages in two of the figured specimens of C. man-
dovi by Reboulet (1996: Pl. 21, Figs. 3, 7) seem to be subcrio-
cone coiled, too.
On the basis of the last mentioned facts, the generic charac-
teristics should be completed by a mention that the whorls of
the Criosarasinella shells are evolute, or may be in early
growth stages subcriocone.
The early whorls of Criosarasinella bearing trituberculate
main ribs differ from the more simply ribbed early whorls
(without trituberculate ribs) of the other subfamily representa-
tives of Neocomitinae Salfeld 1921 (maybe with the exception
of Sarasinella Uhlig, 1905). On the other hand, the style of
ribbing and the tendency towards criocone coiling are closer
to Crioceratites. For the given reasons, Criosarasinella is sup-
posed to be reclassified from the family of Neocomitidae
(Neocomitinae subfamily) to the heteromorphic ammonoid
family of Ancyloceratidae (Crioceratitinae subfamily).
Criosarasinella furcillata Thieuloy, 1977
1977 Criosarasinella furcillata n. sp. Thieuloy, p. 109, Pl. 5,
Figs. 3, 4, ?5
THE OLDEST CRIOCERATITINAE FROM THE WESTERN CARPATHIANS (SLOVAKIA) 247
Fig. 2. Section on the 11
level of the Butkov Quarry with the distribution of ammonoids.
1994 Criosarasinella furcillata Thieuloy Vaíèek et al., p. 59,
Pl. 18, Fig. 2
1996 Criosarasinella furcillata Thieuloy Reboulet, p. 75, Pl. 17,
Fig. 5, Pl. 18, Figs. 18, Pl. 19, Figs. 16, Pl. 20, Figs. 26
Material: Several fragments and two favourably pre-
served shells (field numbers BK11-25/14 = SNM Z 23553 and
BK11-25/5) were added to the shell illustrated in Vaíèek et
al. (1994). In both cases, the most juvenile whorls are not pre-
served and shells end somewhere near to the beginnings of the
body chambers; the latter of the given specimens being pre-
served as an impression.
Description: Evolute microconchs; some shells subcrio-
cone in the early growth stage. Trituberculate ribs of the speci-
men 25/14 alternate with simple ribs without tubercles up to
the diameter of about 18 mm. The former ribs are usually ac-
companied by thinner subsidiary ribs running from a common
umbilical tubercle. In the next stage, thin and dense ribs fol-
low without differentiation between the main and the subsid-
iary ones. At first, they are straight, slightly proverse, then
they become slightly S-shaped. Ribs run partly in pairs from
the umbilical tubercles at the umbilicus, partly in single, and
they are not accompanied by any tubercles. On the periphery
of the shell, short ribs are inserted between most of the ribs, or
are split from them, and their number is higher.
Remarks: In addition to shells coiled evolutely in the
whole course of growth, specimens of Criosarasinella furcil-
lata in non-adult stages may also occur with temporarily subc-
Fig. 3. 1 Criosarasinella furcillata Thieuloy; Spec. SNM Z 23553, microconch (m); 11
level, layer No. 25. 2 Criosarasinella man-
dovi Thieuloy; Spec. SNM Z 23554, (m); 11
level, layer No. 26. 3 Criosarasinella mandovi Thieuloy; Spec. SNM Z 23555, (m); 7
el, layer No. 51. 4 Criosarasinella subheterocostata Reboulet; Spec. SNM Z 23556, (m); 11
level, P1 horizon. 5 Criosarasinella sub-
heterocostata Reboulet; Spec. SNM Z 23557, (m); 11
level, layer No. 25. 6 Crioceratites heterocostatus Mandov; Spec. SNM Z 23558,
level, P2 horizon. Ladce Formation, lower part of the furcillata Ammonite Zone (figures 15); Figure 6 Mráznica Formation,
furcillata Zone. All specimens are figured in natural size.
THE OLDEST CRIOCERATITINAE FROM THE WESTERN CARPATHIANS (SLOVAKIA) 249
kanides) see Mandov (1976), and in the High Tatras in Po-
land in the Kocieliska Formation (Lefeld 1974).
Occurrence: The findings come from the Ladce Forma-
tion on the 11
level (26 m above the base of the section
Fig. 2) and on the 7
level (at 51 m).
Criosarasinella subheterocostata Reboulet, 1996
1996 Criosarasinella subheterocostata n. sp. Reboulet, p. 81, Pl.
22, Figs. 19 (cum syn.)
Material: Two incomplete sculpture moulds (BK11-P1/1 =
SNM Z 23556, BK11-25/4 = SNM Z 23557).
Description: Evolute shells of medium size (preserved
by D of about 60 mm). Whorls are not high, approximately
correspond to the width of the umbilicus. The whorls overlap
one another little. The sculpture of juvenile whorls is clear
from the diameter of about 10 mm. They consist of tritubercu-
late main ribs with subsidiary ribs in singles, sometimes occur
in pairs. Lateral and marginal tubercles are strong, umbilical
ones are weak. Subsequently, the conspicuousness of the lat-
eral ribs is suppressed and at the diameter of about 20 mm, the
lateral tubercles fade out. The umbilical tubercles become
rather emphasized. From the umbilical tubercles three thin ribs
usually run out. Equally strong subsidiary ribs are embedded
between bundled ribs.
After the part with imperfectly preserved ribbing, the body
chamber follows according to markedly stronger, rather heter-
ogeneous ribbing. From the umbilical tubercles, S-shaped ribs
run out in singles, or in pairs. With few exceptions, the ribs bi-
furcate approximately at the half of the whorl height. On the
external side, the ribs are inclined towards the mouth. Over the
external side they pass without any interruption.
Measurement: On spec. 25/4, Wh=19.1 (0.38) and
Uw=20.6 (0.41) were measured at D=50 mm. The maximum
preserved diameter was about 60 mm.
Remarks: C. subheterocostata differs from C. furcillata
and C. mandovi by bundles of three ribs running from the um-
bilical tubercles after the early trituberculate stage and by the
lesser number of short inserted ribs on the periphery. The rib-
bing of the body chamber of C. subheterocostata is rather un-
levelled, S-shaped. The prevailing majority of the ribs bifur-
cate at about half the height of the whorl, i.e. far lower than
the ribs of the remaining species are inserted (which is the
dominating variant), or split. The last whorl has a greater
height in comparison with the former species. The height of
the whorl and the character of the sculpture of the last whorl
remind us somewhat of Teschenites subpachydicranus Rebou-
let, the juvenile whorls of which, however, lack the tritubercu-
C. subheterocostata should be, according to its name, close
to C. heterocostata Mandov sensu Thieuloy (1977). With ref-
erence to the fact that shells of C. heterocostata are distinctly
criocone, C. heterocostata does not belong to Criosarasinella
but to Crioceratites.
Distribution: Reboulet (1996) states the uppermost part
of the furcillata Subzone (Subheterocostata Horizon) from
riocone whorls. This can be documented primarily by the
Thieuloys specimen (1977: Pl. 5, Fig. 4), or by the specimen
25/14 from the Butkov locality. Reboulet (1996) in his synon-
ymy designated Thieuloys above mentioned specimen of C.
furcillata (Pl. 5, Fig. 4) on the basis of criocone coiling as Cri-
oceratites n. sp. 1 (= Crioceratites coniferus in Busnardo et al.
2003). The assignation to Cr. coniferus is inconsistent with
the fact that the given species bears periodical trituberculate
main ribs on all whorls. Numerous bifurcate ribs characteristic
just of Criosarasinella completely exclude any resemblance to
Measurement: At D=53.5 mm (almost D
), spec. 25/
14 has Wh=18.5 mm (0.345) and Uw=21.4 (0.40). At D=
59 mm, spec. 25/5 has Wh=20.8 (0.35) and Uw=25.0 (0.42).
Distribution: According to Reboulet (1996), the species
given occurs in the Vocontian Trough in the ammonite furcilla-
ta Zone (Late Valanginian, Reboulet & Atrops 1999).
Occurrence: The new findings come from the Ladce
Formation on the 11
level of the Butkov Quarry, 25 m above
the base of the section (Fig. 2); fragments of whorls also occur
in the horizon of 26 m.
Criosarasinella mandovi Thieuloy, 1977
1977 Criosarasinella mandovi n. sp. Thieuloy, p. 110, Pl. 5,
Figs. 6, 7 (cum syn.)
1996 Criosarasinella mandovi Thieuloy Reboulet, p. 78, Pl. 16,
Fig. 4, Pl. 20, Figs. 1, 7, Pl. 21, Figs. 1, 37 (cum syn.)
Material: 4 incomplete shells and 4 relatively well pre-
served, partly deformed shells (BK7Z-51/8 = SNM Z 23555;
BK11-26/21, 24 and 23 = SNM Z 23554, the last one with a
more complete counter-impression).
Description: Early whorls of evolute microconchs are
well preserved (in spec. 26/23 the shell diameter 8 to 25 mm).
Whorls remain in contact on the whole shell. Main tritubercu-
late ribs are somewhat stronger than simple subsidiary ribs. A
subsidiary thinner simple rib runs simultaneously from each
umbilical tubercle of the main ribs. The subsidiary ribs (usual-
ly two ribs between the last trituberculate ribs) are simple.
Further, rather thin ribs follow that bear sporadically umbilical
tubercles from which ribs in pairs run. The other equally
strong ribs are simple. On the external side, rather short ribs
are more and more frequently inserted between the ribs. Rib-
bing changes in the zone of passing the phragmocone into the
body chamber. Ribs are stronger and conspicuously sparse.
Sometimes, umbilical tubercles are only indicated on them
there; but sometimes, they are quite conspicuous. Only rarely,
two ribs run from the umbilical tubercles; simple ribs occur
most frequently. On the periphery, the majority of ribs bifur-
cate, or the bifurcation is replaced by short inserted ribs.
Measurement: Microconchs with the average diameter
of 7077 mm. On shell 51/8, at D=76 mm, Wh=25.0 (0.33)
and Uw=32.7 (0.43).
Distribution: Reboulet (1996) states C. mandovi from
Upper Valanginian in the Vocontian Trough (ammonite furcil-
lata Zone, Reboulet & Atrops 1999). According to Thieuloy
(1977) the species also occurs in Bulgaria (Western Bal-
Fig. 4. 1 Crioceratites coniferus Busnardo, Charollais, Weidmann et Clavel; Spec. SNM Z 23560, macroconch (M); 11
level, layer No.
26. 2 Crioceratites coniferus Busnardo, Charollais, Weidmann et Clavel; Spec. SNM Z 23561, microconch (m); 11
level, debris between
P1 and P2 horizons. 3 Himantoceras trinodosum Thieuloy; Spec. SNM Z 23562; 11
level, layer No. 26. 4 Himantoceras trinodosum
Thieuloy; Spec. SNM Z 23563; 11
level, horizon P2. 5 Crioceratites heterocostatus Mandov; Spec. SNM Z 23559, (m); 11
bris below P1 horizon. Ladce (figures 1, 3) and Mráznica Formations, the ammonite furcillata Zone (Upper Valanginian). All specimens are
figured in natural size. Photos taken by K. Mezihoráková, Ostrava. Before photographing, all specimens were bleached by ammonium chlo-
ride. All shells figured will be deposited in the Slovak National Museum in Bratislava under inventory numbers SNM Z 2355323563.
THE OLDEST CRIOCERATITINAE FROM THE WESTERN CARPATHIANS (SLOVAKIA) 251
Occurrence: The Ladce Formation (25 m above the base
of the section) and the lower part of the Mráznica Formation
(horizon P1) on the 11
level (Fig. 2).
Genus: Crioceratites Léveillé, 1837
Type species: Crioceratites duvali Léveillé, 1837
Crioceratites heterocostatus Mandov, 1976
Figs. 3.6, 4.5
1976 Crioceratites (Crioceratites) majoricensis heterocostatus n.
subspec. Mandov, p. 57, Pl. 5, Figs. 1, 3
?1977 Criosarasinella heterocostata (Mandov) Thieuloy, p. 111,
Pl. 5, Fig. 8
1983 Criosarasinella heterocostata Mandov Vaíèek et al., Pl. 1,
1986 Criosarasinella heterecostata (Mandov) Vaíèek et
Michalík, p. 469, Pl. 4, Fig. 3
1996 Crioceratites primitivus n. sp. Reboulet, p. 175, Pl. 23,
Fig. 3, Pl.24, Figs. 15
Material: Three rather well preserved sculpture moulds,
in which the earliest whorls have not been preserved (BK11-
P1/14, BK11-P2/5 = SNM Z 23558, BK11-P1/1s = SNM Z
Description: Criocone shells. The sculpture changes in
the three basic stages. In the earliest stage, which is merely un-
clearly preserved and ends at the shell diameter of about eight
mm, thin ribs are clear. They bear slight lateral and marginal
tubercles from time to time. Any umbilical tubercles are not
The second stage is characterized by main ribs with three
rows of tubercles. At the beginning, strongly developed lateral
tubercles are conspicuous with the wide base of a circular out-
line. Umbilical and marginal tubercles are weaker. Between
the main ribs, subsidiary ribs are inserted merely in singles at
first. Soon the number of ribs inserted increases slowly and the
markedness of lateral tubercles fades out. Sporadically, some
subsidiary ribs run from the umbilicus in pairs. When about 45
subsidiary ribs are found in the interval between two main ribs,
some of them bifurcate on the periphery or short inserted ribs
are inserted between them on the same level. Before the end of
the phragmocone, when the number of subsidiary ribs between
the main ribs is up to 7 (somewhere about the diameter of
30 mm), the trituberculate main ribs disappear. On the rear side,
these ribs are marked by a shallow, rather wide constriction.
The third stage represents the body chamber. The main fea-
ture is represented by rather strong and sparse, straight and
slightly S-shaped ribs separated by occasional shallow con-
strictions. Some of the ribs can run out in pairs from the umbi-
licus, others bifurcate on the external side, or the bifurcation
can be replaced by short inserted ribs. On the other hand,
equal, simple, rather sparse ribs seem to prevail on the body
chamber. Marginal tubercles are indicated on all ribs, especial-
ly at the beginning of the body chamber.
Measurement: The preservation of the shells does not
allow desirable measurement accuracy of the parameters re-
quired. The largest shell (P1/1s) attains the maximum diame-
ter only slightly over 72 mm. At D=72.0 mm, H=19.8
(0.275), U is about 39 mm (0.54).
Remarks: As it follows from the synonymy, I regard C.
primitivus as a younger synonym of the name C. heterocosta-
tus Mandov. As Thieuloy (1977) states, from the stratigraphi-
cal point of view, Mandovs material represents a heteroge-
neous association of ammonoids, the components of which are
of Late Valanginian and partly of Early Hauterivian age. Man-
dovs holotype of C. heterocostatus comes most likely from
The specimens with criocone shells found at Mt Butkov,
which are deformed in various degrees, correspond in size, to
the category of microconchs, similarly to the type material of
Mandov (1976). Reboulets material (1996) designated as Cri-
oceratites primitivus corresponds partly to incomplete, but
usually better preserved macroconchs. With the Slovak speci-
mens, the trituberculate stage disappears probably somewhat
later (at D of about 35 to 45 mm) than with the French materi-
al (at D of about 30 mm). The Slovak specimens correspond
perfectly to Mandovs description (1976), including the inter-
ruption of ribs on the body chamber in the siphonal area.
Distribution: Reboulet (1996) placed his findings to the
uppermost part of the ammonite trinodosum Zone, which cor-
responds to the furcillata Subzone according to Reboulet &
Atrops (1999) and to Hoedemaeker & Reboulet (2003). The
Bulgarian specimens come from the Western Balkanides.
Occurrence: All the specimens come from the lower part
of the Mráznica Formation on the 11
level (horizons P1, P2)
from the upper part of the furcillata Subzone (Fig. 2).
Crioceratites coniferus Busnardo, Charollais, Weidmann et
1996 Crioceratites n. sp. 1 Reboulet, p. 174, Pl. 23, Figs. 1, 2, 4,
?Pl. 24, Fig. 7
2003 Crioceratites coniferus n. sp. Busnardo et al., p. 61, Pl. 4,
Figs. 3, 6
Material: Two imperfectly preserved sculpture moulds,
with which juvenile whorls are not preserved. The lesser shell
represents most likely a microconch (BK11-P2/2s = SNM Z
23561), the larger one a macroconch (BK11-26/10 = SNM Z
Description: The microconch with preserved one and a
half of the whorl. The adult whorl is of medium size. The shell
is coiled in a free spiral. On the partly preserved phragmo-
cone, on which the sculpture is preserved, there are tritubercu-
late main ribs and thin simple subsidiary ribs, about four in
number. The body chamber, indicated by a different mode of
fossilization, which forms approximately a half of the whorl,
is covered by trituberculate main ribs separated by a bundle of
five thinner simple subsidiary ribs. Umbilical tubercles on the
main ribs seem to be the weakest tubercles. Lateral tubercles
situated considerably high at about 2/3 of the whorl height are
conspicuous. Marginal tubercles protrude into nine mm long
spines. All the simple subsidiary ribs have indicated marginal
tubercles on the last half of the whorl. Subsidiary ribs may
sporadically bifurcate at about the height where lateral tuber-
cles are developed. A narrow smooth zone in the siphonal area
cannot be excluded.
A criocone macroconch begins with a not high last but one
whorl belonging to the phragmocone. The height of the earli-
est preserved whorl is about 5 mm. Trituberculate ribs are em-
bedded here with 45 subsidiary ribs. On the last relatively
high half a whorl that probably pertains to the body chamber,
trituberculate main ribs can be seen, between which 67 sub-
sidiary ribs occur. All the tubercles on the main ribs are rough-
ly equivalent. One to two subsidiary ribs may bifurcate at the
lateral tubercles. It cannot be excluded, that all the subsidiary
ribs bear weak marginal tubercles and that the ribs in the siph-
onal area are interrupted, or at least weakened. Shallow con-
strictions have been observed on the frontal side of the three to
four last trituberculate ribs of the final whorl.
Measurement: The microconch attains the shell diame-
ter of a little more than 60 mm, the macroconch about
115 mm. The other parameters could not be measured with the
required accuracy. In both the cases, six main ribs fall per half
Remarks: Of the specimens given in the synonymy, it is
merely Reboulets juvenile shell (1996) illustrated in
Plate 24, Fig. 7 that differs from the Slovak material by
coarser ribbing. As it follows from Busnardos et al. (2003)
holotype macroconch morphology, just the other shells illus-
trated by Reboulet in Plate 23 from Criosarasinella furcillta
Zone fully correspond to the newly determined C. coniferus.
Small marginal tubercles on the subsidiary ribs of the body
chamber and also the rare bifurcation of inserted ribs on the
macroconch of the Slovak criocone material are remarkable.
The features presented indicate a link between C. coniferus
and C. heterocostatus.
Busnardo et al. (2003) consider that Crioceratites (C. co-
niferus) appear at the base of the Hauterivian stage only. The
Valanginian Crioceratites primitivus Reboulet is either re-
garded as Himantoceras Thieuloy, 1964 or (at least) phyleti-
cally related to it. The coiling type of one of the specimens il-
lustrated by Reboulet (1996: Pl. 24, Fig. 1) indicates the latter
possibility. However, the shell formation of other specimens
of Cr. heterocostatus (= Cr. primitivus) and shells of Himan-
toceras is anyhow different (also from the point of view of a
growth in whorl height), and thus the opinion presented may
be disputed. On the other hand, the Crioceratites coniferus,
whose generic assignation is undoubted, lacks himantocera-
toid coiling. According to the findings described here and to
Reboulets findings (1996) as well, both the above mentioned
species and genus appeared as early as in the Late Valanginian
Criosarasinella furcillata Zone.
Distribution: With the exception of the specimen in his
Plate 24, Reboulet (1996) reported C. coniferus from the up-
per part of the Late Valanginian furcillata Subzone in the Vo-
contian Trough; Busnardo et al. (2003) indicated the occur-
rence of the holotype specimen in the basal Hauterivian
radiatus Zone in the External Prealps in Switzerland.
Occurrence: The macroconch comes from the upper part
of the Ladce Formation on the 11
level of the Butkov Quar-
ry, from horizon 26, where it occurs together with Criosa-
rasinella furcillata and C. mandovi; the microconch was
found in the debris between horizons P1 and P2 on the 11
level (Fig. 2) in the topmost Valanginian rocks.
Genus: Himantoceras Thieuloy, 1964
Type species: Himantoceras trinodosum Thieuloy, 1964
Himantoceras trinodosum Thieuloy, 1964
1996 Himantoceras trinodosum Thieuloy Reboulet, p. 171, Pl. 25,
Figs. 611 (cum syn.)
The material consists of two fragments of the last whorl and
of another two rather complete sculptural moulds. One of
them preserved the last but one whorl (BK11-26/19 = SNM Z
23562) and the other one belongs to the adult part of the shell
that pertains to the phragmocone in part and to the body cham-
ber in the remaining part (BK11-P2/1 = SNM Z 23563).
Description: The best preserved shell begins with crio-
cone whorls that pass into free, long, arc/like arm in maturity.
Even at the preserved most juvenile part, trituberculate ribs
can be seen. Umbilical tubercles are the weakest there; mar-
ginal tubercles are the strongest. Between the main ribs, vari-
able numbers of thin ribs are inserted (03). Further, all the tu-
bercles are as strong as the marginal tubercles and the number
of subsidiary ribs increases to 46. On the body chamber, sub-
sidiary ribs are markedly stronger and more sparsely distribut-
ed. The number of them varies from six to seven.
Measurement: The arch of the last half-whorl of spec. 26/
19 has a span (D) of 72 mm. The whorl height of the earliest
preserved part is four mm, at the end of the shell it reaches
about 15 mm.
Remarks: The Slovak specimens are very close to the
material illustrated by Thieuloy (1979: Pl. 3, Figs. 10, 11)
from the La Charce section. The coiling of shells of represen-
tatives of Himantoceras (including H. lessinianum Faraoni,
Flore, Marini, Pallini et Pezzoni, 1997 described recently) is
different from the coiling of typical Crioceratites representa-
Distribution: H. trinodosum served as the Late Valang-
inian zonal species as late as a short time ago (Reboulet & At-
rops 1999). In addition to French localities (Vocontian
Trough), it is known from Spain (Betic Cordillera), from the
Western Carpathians (imperfectly preserved fragments from
the Stráovská hornatina Upland and from the Pieniny Klip-
pen Belt) and from Bulgaria.
Occurrence: Two specimens come from the upper part
of the Ladce Formation and two from the lower part of the
Mráznica Formation on the 11
level (17.5 and 26 m above
the base of the section and horizons P4 and P2 see Fig. 2);
the last one comes from the debris at the prospecting gallery
No. 3 on the 6
Recently, the oldest occurrences of true Crioceratites were
usually attributed to the upper part of the Lower Hauterivian.
However, in 1996, Reboulet proved that the first Crioceratiti-
nae appeared as early as in the Late Valanginian in the Vocon-
tian Trough. On the other hand, Busnardo et al. (2003) have
THE OLDEST CRIOCERATITINAE FROM THE WESTERN CARPATHIANS (SLOVAKIA) 253
made some critical remarks on Reboulets taxonomic assigna-
tion of his Late Valanginian species.
Criosarasinella, occupying a rather unclear position, is one
of three ancyloceratid ammonite genera which occur at Mt
Butkov. According to Thieuloy (1977), adult whorls of Cri-
osarasinella shells remain in contact. On the other hand, (sel-
domly preserved) juvenile whorls can be slightly criocone
coiled and poses trituberculate ribs. In spite of this fact, Thieu-
loy (1977) and also subsequent authors (including Wright et
al. 1996 new Treatise) regard Criosarasinella as the rep-
resentative of Neocomitinae. However, the trituberculation of
main ribs on internal whorls, or the subcriocone shells of C.
furcillata and C. mandovi contradicts the rules of phylogene-
sis to such a degree, that Criosarasinella cannot be assigned
to the neocomitids. Anyway, Thieuloy himself (1977) has stat-
ed subsequently that Criosarasinella represents a new element
in the conception of the ancyloceratid origin. According to
him, juvenile whorls may be interpreted as a new phenomenon
passing from the internal whorls to the body chamber.
This peculiar new morphology is distinctly shown by Cri-
osarasinella mandovi and by the somewhat younger Criocer-
atites heterocostatus. The shell of the latter species is already
conspicuously criocone in the whole range. However, Rebou-
let (1996) believed that the character of Criosarasinella rib-
bing does not reputedly permit (?) the primitive crioceratids to
be derived from it.
In addition to the Late Valanginian C. heterocostatus, an-
other species of Crioceratitinae, i.e. Crioceratites coniferus
Busnardo occurs at Butkov as well. They occur in the same
horizon as Criosarasinella mandovi (then even somewhat ear-
lier than C. heterocostatus). Like C. heterocostatus, the shell
of C. coniferus is also distinctly criocone. In contrast to C. het-
erocostatus, it bears trituberculate main ribs even on the body
According to Busnardo et al. (2003), C. coniferus safely oc-
curs in the Early Hauterivian radiatus Zone. Thus, the Swiss
findings bridge the gap between Upper Valanginian and Low-
er Hauterivian Crioceratites occurrences, mentioned by Re-
The stratigraphic position of criocone shells and the compo-
sition and age of the accompanying association of ammonoids
from the Upper Valanginian Ladce and Mráznica Formations
at Butkov are very close to those of the Late Valanginian am-
monoid association of the same age from the Vocontian
Trough (see Reboulet 1996).
In accordance with Ceccas conception (1997), it is sup-
posed that the heteromorphic ammonoids (bochianitids, pro-
tancyloceratids, etc.) represented a special Tithonian to Early
Valanginian evolutionary branch that was not followed by the
ancyloceratid branch. The character of shell coiling of the old-
est ancyloceratids considerably varied, similarly to other
groups at the beginning of their evolution. The ancyloceratids
appeared during the upper Late Valanginian, probably about
the end of the ammonite verrucosum Zone. A possibly solu-
tion was indicated by Reboulet (1996: Figs. 18 and 19) in the
phylletic appearance of Himantoceras from Neocomites neo-
comiensis (dOrbigny) via N. beaumugnensis Sayn and
In addition to planispirally coiled shells with overlapping
whorls, the rich collection of Upper Valanginian ammonoids
from the Western Carpathian Mt Butkov locality contains het-
eromorphic ammonoids with trituberculate ribs. The whorls of
Crioceratites and Himantoceras are mostly not in contact at
all, but in other forms (like Criosarasinella), they touch one
another and may have a tendency towards uncoiling. Of three
species of this genus that occur at Butkov, the C. mandovi and
also C. furcillata zonal species document that on some shells,
whorls may lose contact with the following whorls, i.e. subc-
riocone shells appear.
Trituberculate ribs on early whorls of Criosarasinella and
their sporadic subcriocone coiling indicate a closer relation-
ship to Crioceratites than to neocomitids, to which Criosa-
rasinella has been assigned so far. In the contribution submit-
ted, the Criosarasinella is classified as an ancyloceratid
(Crioceratitinae subfamily) and not as neocomitid (Neocomiti-
nae subfamily). As for stratigraphy, the ammonoid association
studied belongs to the Criosarasinella furcillata Zone, as
shown by the occurrence of the zonal species. It also follows
that Crioceratites heterocostatus and C. coniferus represent
the oldest representatives of the Crioceratites, showing that
this genus evolved as early as in the Late Valanginian.
Acknowledgments: The greatest thanks belong to the Grant
Agency of the Czech Republic for giving the postdoctoral
grant to P. Skupien, in the framework of which the significant
ammonite collection was obtained. However, the authors
thanks go especially to those who participated with the author
in field sampling done in the quarry for the last three years.
They are, in addition to the grant holder, J. Michalík (SAV
Bratislava), D. Reháková (UK Bratislava) and L. Kratoch-
vílová (VB Ostrava). Mrs K. Mezihoráková is also thanked
for taking photographs of ammonites. Critical and construc-
tive reviews by F. Cecca (University of Paris) and especially
S. Reboulet (University of Lyon) contributed to the consider-
able improvement of this paper.
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