GEOLOGICA CARPATHICA
, DECEMBER 2016, 67, 6, 543 – 559
doi: 10.1515/geoca-2016-0034
www.geologicacarpathica.com
Upper Tithonian ammonites (Himalayitidae Spath, 1925 and
Neocomitidae Salfeld, 1921) from Charens (Drôme, France)
CAMILLE FRAU
1
, LUC G. BULOT
2
, WILLIAM A.P. WIMBLEDON
3
and CHRISTINA IFRIM
4
1
Groupement d’Intérêt Paléontologique, Science et Exposition, 60 bd Georges Richard, 83000 Toulon, France; camille_frau@hotmail.fr
2
UM 34 Aix-Marseille Université — CNRS (UMR 7330) — IRD (UMR 161), Centre Saint-Charles, Case 67, 3 Place Victor Hugo,
13331 Marseille Cedex 03, France
3
School of Earth Sciences, University of Bristol, Queens Road, Bristol BS8 1RJ, United Kingdom
4
Institut für Geowissenschaften, Ruprecht-Karls-Universität, Im Neuenheimer Feld 234, 69120 Heidelberg, Germany
(Manuscript received January 28, 2016; accepted in revised form September 22, 2016)
Abstract: This contribution focuses on the Perisphinctoidea ammonite taxa from the Upper Tithonian at Charens (Drôme,
south-east France). Emphasis is laid on five genera that belong to the families Himalayitidae and Neocomitidae.
We document the precise vertical range of the index-species Micracanthoceras microcanthum, and a comparative
ontogenetic-biometric analysis sheds new light on its range of variation and dimorphism as compared to the best-
known Spanish populations. As herein understood, the lower boundary of the M. microcanthum Zone (base of the Upper
Tithonian) is fixed at the FAD of its index species. The faunal assemblages and species distribution of the
P. andreaei Zone are rather similar to those described at the key-section of Le Chouet as confirmed by the co-occurrence
of the genera Protacanthodiscus, Boughdiriella and Pratumidiscus. New palaeontological evidence supports the view
that the basal Neocomitidae Busnardoiceras busnardoi was derived from Protacanthodiscus andreaei in the upper part
of the P. andreaei Zone.
Key words: Ammonoidea, Tithonian, Jurassic, biostratigraphy, Drôme, France.
Introduction
Within the activity of the Berriasian Working Group (BWG) of
the International Subcommission for Cretaceous Stratigraphy
(ISCS/IUGS), Bulot et al. (2014) and Frau et al. (2015)
focused on the systematics of the late Tithonian Perisphinctoidea
(Ammonitina, Ammonoidea) from Le Chouet (Drôme); a sec-
tion that stands as a reference for the definition of the Jurassic/
Cretaceous boundary in south-east France (Wimbledon et al.
2013). However, we failed to characterize the exact range and
intraspecific variation of the late Tithonian index species
Micracanthoceras microcanthum (Oppel, 1865) due to its
sporadic occurrence and the poor preservation of the specimens
collected (Bulot et al. 2014). A new survey of the sections
of the Drôme river valley already documented by Remane
(1970) led us to sample in detail the Charens section where
the lower part of the Upper Tithonian is better exposed than
at Le Chouet and it has yielded rich ammonite faunas. The aim
of the present contribution is to discuss the taxonomy and
illustrate the Himalayitidae and Neocomitidae in this new
collection.
Geological setting
The Charens section is located about three kilometres to the
west of Le Chouet (Drôme, France, Fig. 1). This section is part
of the late Kimmeridgian to early Berriasian “turbiditic”
system of the Subalpine Basin deposited along the northern
Tethys margin (Ferry & Grosheny 2013). The locality lies in
the upper valley of the Drôme River (Long 5°31’08”E,
Lat 44°32’42”N) off the local road (D93) from Die to
Beaurières. It comprises a low mural section, 250 m-long,
along a single-track road (D145) on the east side of the Charens
gorge. The section extends around the slope from an altitude
of 756 m to 768 m and dips 10 degrees to the North.
The studied part of the succession consists of 41 metres of
well-bedded limestone autochthonous intervals intercalated
between conglomeratic, clast or matrix-supported breccias
(Fig. 1). The bottom part of the section corresponds to thin-
bedded biocalcarenites (beds 5 to 12) above a thick conglo-
meratic breccia (bed 1 to 5), that pass into a 12m-long,
ammonite-bearing, biomicrite succession (beds 13 to 67).
The first occurrence of ammonites is reported in bed 17 and
contains Burckhardticeras ponti (Fallot & Termier, 1923) and
Lemencia sp. This assemblage is indicative of the Burckhardti
ceras ponti Zone sensu Enay & Geyssant (1975) that marks
the uppermost Lower Tithonian. The First Appearance Datum
of M. microcanthum is herein reported at the base of the bed 25
in association with Lemencia sp. and poorly-preserved frag-
ments of Cordubiceras cf. cordubae (Olóriz & Tavera, 1979)
(Fig. 1). As herein understood, the FAD of M. microcanthum
marks the lower boundary of the Upper Tithonian (see
discussion in the Conclusion). Above, thick massive, clast-
and matrix supported breccias affect the succession. The bio-
micrite interval (beds 79 to 98) that crops out above the
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FRAU, BULOT, WIMBLEDON and IFRIM
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, 2016, 67, 6, 543 – 559
Fig. 1. Locality map of Le Chouet and Charens sections (Drôme, SE France),
biostratigraphy and vertical range of the late Tithonian Himalayitidae and
Neocomitidae studied in this paper.
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brecciated interval can be correlated with certainty to beds 72
to 89 at Le Chouet section. This is supported by ammonite
occurrences [Moravisphinctes fischeri (Kilian, 1889), Parau
lacosphinctes senoides Tavera, 1985, Protacanthodiscus
andreaei (Kilian, 1889) etc.] and useful lithological markers
such as the intraclastic/microbreccia bed 88 (= bed 80 of
Le Chouet section) and the clast-supported breccia bed 95 – 97
that correspond to the grainstone-conglo merate-grainstone
triplet of Le Chouet (= beds 86 to 88).
The calpionellid assemblages of the Charens section were
previously described by Remane (1970, fig. 6) but the reso-
lution of the drawing prevents us from a bed-by-bed correlation.
A detailed study of the calpionellids and calcareous nanno-
fossils, is currently in progress in the BWG.
Systematic palaeontology
Preservation of our specimens as crushed internal moulds
prevents us from giving other measurements than Dmax = larger
measurable diameter, U = adult width of the umbilicus and
Wh = adult whorl height. The ratios U / D, Wh / D and Wb / D
(umbilical dimension, whorl height and whorl breath as a per-
centage of the adult diameter), are discussed in systematic
descriptions. In synonymies, we distinguished the macroconch
[M] and microconch [m] specimens.
All studied specimens are deposited in the Frau / Bulot
collection at the Musée Paléontologique de Provence (MPP)
of the Aix-Marseille Université, France. All specimens from
Charens were coated with ammonium chloride prior to photo-
graphy. The following abbreviations indicate the repository of
specimens mentioned in the text:
FSL and EM: University of Claude Bernard — Lyon-I.
UJF-ID: University of Grenoble.
BSPM: Bayerische Staatssammlung für Paläontologie und
Geologie, Munich.
Order Ammonoidea Zittel, 1884
Suborder Ammonitina Hyatt, 1889
Superfamily Perisphinctoidea Steinmann, 1890
Family Himalayitidae Spath, 1925
Subfamily Himalayitinae Spath, 1925
Genus Micracanthoceras Spath, 1925
Type species. Ammonites microcanthus Oppel in Zittel,
1868, by original designation of Spath (1925, p. 144).
Remarks. Bulot et al. (2014) recently shed new light on the
content of Micracanthoceras to which the reader is referred.
However, this revision was not supported by palaeontological
evidence due to the poorly-preserved specimens collected at
Le Chouet. New collections from Charens allow the study of
intraspecific variation in Micracanthoceras microcanthum
(Oppel in Zittel, 1868) throughout its range. A biometric com-
parison with the many species introduced by Tavera (1985)
from the Betic Cordillera (Spain) is provided ) (see Table 1 in
Supplementary data).
Micracanthoceras microcanthum (Oppel in Zittel, 1868)
Figs. 2A– C, 3A,B and 4A,U
Macroconchs [M]
1865 Ammonites Symbolus – Oppel, p. 555
1865 Ammonites Köllikeri – Oppel, p. 555
1868 Ammonites microcanthus – Oppel in Zittel, p. 93, pl. 17,
figs. 1a-b,2
1868 Ammonites Symbolus – Oppel in Zittel, p. 96, pl. 16, figs. 6a,c,
7a,c
1868 Ammonites Köllikeri – Oppel in Zittel, p. 95, pl. 18, fig. 1a,c
non 1868 Ammonites Köllikeri – Oppel in Zittel, p. 95, pl. 18, fig. 2a,c
[= Ardesciella rhodanica (Mazenot)]
non 1886 Perisphinctes Köllikeri (Oppel) – Nicolis & Parona, p. 77,
pl. 4, fig. 6 (= ?Simoceratidae gen. et sp. indet.)
non 1890 Hoplites Koellikeri (Oppel) – Toucas, p. 607, pl. 18,
fig. 11A,B [= Ardesciella rhodanica (Mazenot)]
1897 Hoplites microcanthus (Oppel) – Roman, p. 284, pl. 1, fig. 10a,b
non 1897 Reineckeia Koellikeri (Oppel) – Steuer, p. 31, pl. 32(8),
fig. 5 [= Corongoceras mendozanum (Behrendsen)], 6
non 1900 Reineckeia Koellikeri (Oppel) – Burckhardt, p. 16, pl. 20,
figs. 14, 15, pl. 21, fig. 1 [= Steueria alternans (Gerth)]
non 1922 Reineckeia Koellikeri (Oppel) – Steuer, p. 57 pl. 8, figs. 5,
6 (= Steuer, 1897, pl. 32(8), figs. 5, 6)
1928 Ammonites Köellikeri (Oppel) – Krantz, pl. 28, pl. 3, fig. 1a,b
non 1931 Berriasella Köellikeri (Oppel) – Windhausen, pl. 29,
figs. 4a,b, 7
1931 Micracanthoceras aff. microcanthum (Oppel) – Yin, p. 33, pl. 2,
fig. 1,1a (= Roman, 1897, pl. 1, fig. 10a,b)
1936 Himalayites (Corongoceras) lamberti – Roman, p. 21, pl. 3,
fig. 5,5a
non 1936 Himalayites (Corongoceras) Kollikeri (Oppel) – Roman,
p. 27, pl. 4, figs. 19,19a, 20 (= ?Protacanthodiscus sp. juv.)
1939 Himalayites (Micracanthoceras) microcanthum (Oppel in
Zittel) – Mazenot, p. 233, pl. 37, fig. 12a,b (= Zittel, 1868,
pl. 17, fig. 1a, 2)
non 1960 Corongoceras Lamberti Roman – Collignon, pl. 167,
fig. 756 [= ?Micracanthoceras brightoni (Spath)]
non 1960 Micracanthoceras cf. symbolus (Oppel) – Collignon,
pl. 175, fig. 753 [= ?Micracanthoceras brightoni (Spath)]
1966 Micracanthoceras (Corongoceras) lamberti Roman – Linares &
Vera, pl. 6, fig. 4, pl. 7, figs. 2, 4
non 1976 Himalayites cf. kollikeri (Oppel) – Khimchiashvili, p. 118,
pl. 11, fig. 3 (= Neocomitidae gen. et sp. indet.)
1985 Djurjuriceras mutari – Tavera, p. 150, pl. 19, fig. 2, pl. 20,
fig. 1, text-fig. 11B
1985 Djurjuriceras mediterraneum – Tavera, p. 152, pl. 19, fig. 1,
text-fig. 11A
1985 Djurjuriceras sinuosum – Tavera, p. 154, pl. 20, fig. 2,
text-fig. 11C
1985 Micracanthoceras (Corongoceras) symbolum (Oppel) – Tavera,
p. 189, pl. 24, fig. 1a,b, text-fig. 14A
1985 Micracanthoceras (Corongoceras) ornatum –Tavera, p. 191,
pl. 25, fig. 8, text-fig. 14E
non 1985 “Corongoceras” köllikeri (Oppel) – Tavera, p. 194, pl. 28,
fig. 2a,b, text-fig. 17G [= Protacanthodiscus hexagonum
(Tavera)]
non 1986 Corongoceras symbolum (Oppel) – De Wever et al., p. 166,
pl. 3, fig. 4 (= Himalayitidae gen. et sp. indet.)
1990 Micracanthoceras microcanthum (Oppel) – Fözy, p. 329, pl. 3,
fig. 4
1990 Corongoceras symbolum (Oppel) – Fözy, p. 328, pl. 5, fig. 2
non 1997 “Corongoceras” kollikeri (Oppel) – Benzaggagh & Atrops,
p. 158, pl. 7, fig. 3 [= Dalmasiceras spiticeroides (Djanélidzé)]
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Fig. 2. Micracanthoceras microcanthum (Oppel in Zittel, 1868): A1–2 — plaster cast FSL.136039 of the lectotype of Ammonites microcanthus
Oppel in Zittel, 1868 [m]; B1–2 — plaster cast FSL.13049 of Ammonites symbolus Oppel in Zittel, 1868 [M]; C1–3 — MPP–CHR.37/3 [M]
bed 37. Scale bar is 10 mm.
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Fig. 3. Micracanthoceras microcanthum (Oppel in Zittel, 1868): A1– 4 — plaster cast FSL. FSL. 13086 of the paratype of Ammonites
microcanthus Oppel in Zittel, 1868 [M]; B1–2 — re-illustration of the holotype of Ammonites koellikeri Oppel in Zittel, 1868
(BSPG–AS/III/468) from Parent et al. (2011, fig. 32) [M]. Scale bar is 10 mm.
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1997 Micracanthoceras (Corongoceras) rhodanicum (Mazenot) –
Benzaggagh & Atrops, p. pl. 6, fig. 2
2000 Micracanthoceras (Corongoceras) rhodanicum (Mazenot) –
Benzaggagh, pl. 4, fig. 3 (= Benzaggagh & Atrops 1997, pl. 6,
fig. 2)
non 2010 Corongoceras cf. symbolum (Oppel) – Benzaggagh et al.,
figs. 5j,k, l,m [= [m] Protacanthodiscus andreaei (Kilian)]
2013 ?Djurjuriceras sp. – Bujtor et al., Fig. 4F
non 2013 Micracanthoceras (Corongoceras) koellikeri (Oppel) –
Szives & Fözy, p. 300, pl. 4, fig. 1 [= [m] Protacanthodiscus
andreaei (Kilian)], pl. 6, fig. 7 (= Protacanthodiscus juv.), pl. 8,
figs. 3 [= [m] Protacanthodiscus andreaei (Kilian)], 4a,b
(= Perisphinctoidea indet. juv.), 9 (= Perisphinctoidea indet.)
2016 Micracanthoceras (Corongoceras) sp. – Bahrouni et al.,
figs. 5.7, 5.12
Microconchs [m]
1865 Ammonites microcanthus – Oppel, p. 155
1868 Ammonites microcanthus – Oppel in Zittel, p. 93, pl. 17,
figs. 3a,b, 4, 5a,c
1868 Ammonites fraudator – Zittel, p. 110, pl. 21, figs. 1a,b, 2a,b, 3
1890 Hoplites microcanthus (Oppel) – Toucas, p. 608, pl. 18, fig. 12
non 1897 Reineckeia microcantha (Oppel) – Steuer, p. 156(30),
pl. 31(7), fig. 3,5 [= ?Corongoceras steinmanni (Krantz)]
non 1900 Reineckeia microcantha (Oppel) – Burckhardt, p. 16, pl. 20,
fig. 16,17 (Perisphinctoidea indet.)
non 1903 Hoplites microcanthus Péron sp. non Oppel– Burckhardt,
p. 58, pl. 10, fig. 12,16 [= ?Corongoceras mendozanum
(Behrendsen)]
non 1921 Hoplites aff. microcanthus (Oppel) – Burckhardt, pl. 18,
fig. 5,9 (= Perisphinctoidea gen et sp. nov.)
non 1922 Reineckeia microcantha (Oppel) – Steuer, p. 56, pl. 7,
fig. 3,5 (= Steuer, 1897, pl. 31(7), fig. 3-5)
non 1931 Micracanthoceras aff. microcanthum (Oppel) – Spath,
p. 543, pl. 92, fig. 3a,b [= Micracanthoceras brightoni
(Spath)]
non 1931 Micracanthoceras sp. nov. aff. fraudator (Zittel) – Spath,
p. 545, pl. 109, fig. 15 [= Micracanthoceras brightoni (Spath)]
1934 Himalayites (Micracanthoceras) microcanthus (Oppel) –
Dacqué, pl. 46, fig. 6, 6a (= Zittel, 1868, pl. 17, fig. 3a,b)
1936 Himalayites (Micracanthoceras) microcanthus (Oppel) –
Roman, p. 22, pl. 4, fig. 6,6a
1936 Himalayites (Micracanthoceras) microcanthus (Oppel) var.
marocana nov. var. – Roman, p. 24, pl. 4, fig. 8,8a
non 1936 Himalayites (Micracanthoceras) microcanthus (Oppel)
var. marocana nov. var. – Roman, p. 24, pl. 4, fig. 9,9a
(= Burckhardticeras sp.)
1938 Himalayites (Micracanthoceras) microcanthus (Oppel) –
Roman, p. 321, pl. 31, fig. 302,302a (= Zittel, 1868, pl. 17,
fig. 3a,b), text-fig. 302 (= Zittel, 1868, pl. 17, fig. 1b)
non 1939 Micracanthoceras n. sp. aff. koellikeri (Oppel) – Imlay,
p. 44, pl. 17, figs. 2, 3, 4 [= Parrasiella astillerense (Imlay)
nom. correct.]
1939 Himalayites (Micracanthoceras) microcanthum (Oppel in
Zittel) – Mazenot, p. 233, pl. 37, fig. 2 a,b (= Toucas, 1890,
pl. 18, fig. 12), 3
non 1956 Himalayites (Micracanthoceras) cf. microcanthus? (Oppel)
– Ksiazkiewicz, p. 214, pl. 25, fig. 3 (= Perisphinctoidea indet.)
1957 Micracanthoceras microcanthus (Oppel) – Arkell et al. ,
p. L356, fig. 468.1a,b (= Zittel, 1868, pl. 17, fig. 3a,b)
non 1960 Micracanthoceras microcanthus (Oppel) – Collignon,
pl. 65, figs. 754, 755 [= Micracanthoceras aff. brightoni
(Spath)]
1966 Himalayites (Micracanthoceras) microcanthus (Oppel) –
Linares & Vera, pl. 5, fig. 2a,b, pl. 6, fig. 5, pl. 7, fig. 3
1977 Himalayites (Micracanthoceras) microcanthus (Oppel) –
Sapunov, pl. 5, fig. 3
1977 Himalayites (Micracanthoceras) fraudator (Oppel) – Sapunov,
pl. 5, fig. 5
1977 Aulacosphinctes linoptychus (Uhlig) – Sapunov, pl. 6, fig. 1
1979 Himalayites (Micracanthoceras) microcanthus (Oppel) –
Sapunov, p. 193, pl. 58, fig. 4 (= Sapunov, 1977, pl. 5, fig. 3)
1979 Himalayites (Micracanthoceras) fraudator (Zittel) – Sapunov,
p. 194, pl. 58, fig. 5
1979 Aulacosphinctes linoptychus (Uhlig) – Sapunov, p. 195, pl. 59,
fig. 1 (= Sapunov, 1977, pl. 6, fig. 1)
1979 Aulacosphinctes venustus Collignon – Sapunov, p. 195, pl. 59,
fig. 2a,b
1982 Himalayites (Micracanthoceras) microcanthus (Oppel) –
Nikolov, p. 213, pl. 77, fig. 1 (= Sapunov, 1977, pl. 5, fig. 3)
1982 Himalayites (Micracanthoceras) fraudator (Zittel) – Nikolov,
p. 213, pl. 77, fig. 2 (= Zittel, 1868, pl. 21, fig. 2a)
1982 Berriasella (Picteticeras) subeudichotoma – Nikolov, p. 64,
pl. 13, fig. 8
1982 Aulacosphinctes linoptychus (Uhlig) – Nikolov, p. 218, pl. 78,
fig. 5
non 1982 Aulacosphinctes linoptychus (Uhlig) – Nikolov, p. 218,
pl. 78, fig. 4a, b, c [= Aulacosphinctes linoptychus (Uhlig)]
1985 Himalayites (Micracanthoceras) microcanthus (Oppel) – Haas
et al., pl. 20, fig. 7
1984 Micracanthoceras (Micracanthoceras) microcanthus (Oppel)
densecostatus nov. ssp. – Vigh, p. 76, pl. 3, fig. 3a,b
1985 Micracanthoceras (Micracanthoceras) microcanthum (Oppel)
– Tavera, p. 169, pl. 21, figs. 1a,b, 2, 3, 4, text-figs. 13A, E, G
1985 Micracanthoceras (Micracanthoceras) cf. brightoni (Spath) –
Tavera, p. 175, pl. 21, fig. 5, text-fig. 13F
1985 Micracanthoceras (Corongoceras) rhodanicum Mazenot –
Tavera, p. 180, pl. 22, figs. 7, 8a,b, 9a,b, text-fig. 14D
1985 Micracanthoceras (Corongoceras) flexuosum – Tavera, p. 187,
pl. 23, figs. 5a,b, 6, 7, text-fig. 14C
1985 Micracanthoceras (Corongoceras) radians – Tavera, p. 178,
pl. 23, figs. 1, 2, 3a,b, 4a,b, text-fig. 14I
1985 Micracanthoceras (Corongoceras) lotenoense (Spath) – Tavera,
p. 176, pl. 23, figs. 11, 12, 13a,b, text-figs. 14J, K
1985 Micracanthoceras (Corongoceras) minor – Tavera, p. 182,
pl. 23, figs. 8a,b, 9, 10, text-fig. 14H
1985 Micracanthoceras (Corongoceras) mendozanum (Behrendsen)
– Tavera, p. 186, pl. 25, fig. 3a,b, text-fig. 14G
1985 Micracanthoceras (Corongoceras) leanzai – Tavera, p. 192,
pl. 25, figs. 4, 5, text-fig. 14F
1985 Himalayitidae gen. y sp. indeterminados – Tavera, p. 233, pl. 28,
figs. 5, 6, text-fig. 17F
1985 Aulacosphinctes parvulus (Uhlig) – Tavera, p. 144, pl. 17, fig. 4,
text-fig. 10G
1985 Aulacosphinctes macer (Collignon) – Tavera, p. 146, pl. 17,
fig. 3
1985 Aulacosphinctes sulcatus – Tavera, p. 146, pl. 17, figs. 1a,b,
2a,b, text-figs. 10E, H
1985 Aulacosphinctes venustus Collignon – Tavera, p. 147, pl. 17,
fig. 5, text-fig. 10F
1985 Micracanthoceras (Corongoceras) symbolum (Oppel) – Tavera,
p. 189, pl. 24, figs. 2, 3, pl. 25, figs. 1a,b, 2, 3, text-fig. 14B
1985 Corongoceras symbolum (Oppel) – Cecca, p. 143, pl. 1, fig. 3
1989 Micracanthoceras microcanthum (Oppel) – Cecca et al., p. 65,
pl. 1, figs. 3, 4a,b
1989 Micracanthoceras aff. microcanthum (Oppel) – Kaiser-Weidich
& Schaier, p. 363, pl. 4, fig. 3a,b
1995 Micracanthoceras microcanthum (Oppel) – Fözy, p. 138, pl. 21,
fig. 7
1995 Micracanthoceras microcanthum (Oppel) – Eliáš & Vašíček,
pl. 1, fig. 4
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Fig. 4. Micracanthoceras microcanthum (Oppel in Zittel, 1868): A1–2 — MPP–CHR.54/1a [?m] bed 25 (top); B — MPP–CHR.34/2 [m]
bed 25 (top); C1–2 — plaster cast of MPP–CHR.29/1 [m] bed 29; D1–2 — MPP–CHR.50/3 [m] bed 25 (base); E — MPP–CHR.34/1 [?M]
bed 25 (top); F — MPP–CHR.60/6 [m] bed 28 (base); G — MPP–CHR.54/1b [m] bed 25 (top); H — MPP–CHR.50/1 [m] bed 25 (base);
I — MPP–CHR.53/2 [m] bed 31; J1–2 — MPP–CHR.60/7 [m] bed 25 (base); K1–3 — MPP–CHR.67/4 [m] bed 35 (base); L — MPP–
CHR.51/2 [m] bed 29 (top); M — MPP–CHR.67/9 [m] bed 35; N1–2 — MPP–CHR.82/1 [m] bed 53; O — MPP–CHR.45/1 [M] bed 45;
P — MPP–CHR.92/11 [m] bed 64; Q — MPP–CHR.92/28 [m] bed 64; R — plaster cast of MPP–CHR.87/1 [m] bed 66; S — MPP–CHR.92/13
[m] bed 64; T — MPP–CHR.92/10 [m] bed 64; U1–2 — MPP–CHR.98/1 [m] bed 76. Scale bar is 10 mm.
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1997 Micracanthoceras microcanthum (Oppel) – Benzaggagh &
Atrops, pl. 5, fig. 4
1997 Micracanthoceras microcanthum (Oppel) – Geyssant in Cariou
& Hantzpergue, pl. 26, fig. 1 (= Zittel, 1868, pl. 17, fig. 3a,b)
non 1998 Micracanthoceras fraudator (Zittel) – Howarth, p. 79,
pl. 15, fig. 2a,b (= Perisphinctoidea indet.)
non 1999 Micracanthoceras fraudator (Zittel) – Fatmi & Zeiss, p. 97,
pl. 45, fig. 3a,b, pl. 55, fig. 5a,b [= Micracanthoceras brightoni
(Spath)]
2000 Micracanthoceras microcanthum (Oppel) – Benzaggagh, pl. 4,
fig. 4 (= Benzaggagh & Atrops, 1997, pl. 5, fig. 4)
non 2009 Micracanthoceras cf. microcanthum (Oppel) – Shome &
Bardhan, p. 199, pl. 4, figs. a,g [= Micracanthoceras brightoni
(Spath)]
2004 Micracanthoceras microcanthum (Oppel) – Marino et al., pl. 2,
fig. 7
2005 Micracanthoceras sp. gr. microcanthum (Oppel) – Boughdiri et
al., pl. 2, fig. 9, 10
2005 Aulacosphinctes sp. gr. sulcatus Tavera – Boughdiri et al., pl. 2,
fig. 7
non 2012 Micracanthoceras microcanthum (Oppel) – Salazar, p. 90,
figs. 4.13a,m (= Steueria sp.)
2010 Aulacosphinctes sp. – Benzaggagh et al., fig. 5d,e
non 2015 Micracanthoceras microcanthum (Oppel) – Salazar &
Stinnesbeck, p. 19, figs. 15a,m (= Salazar, 2012, figs. 13a,m)
Types: The lectotype is the specimen drawn by Zittel (1868,
pl. 17, fig. 3a,b). A plaster cast of the lectotype (FSL.136039)
and the large (FSL. 13086) Zittel’s paratype (see Zittel, pl. 17,
fig. 1a,2) are herein illustrated in Fig. 2A
and Fig. 3A. For
comparison, we re-illustrate the lectotype BSPM-AS/III/468
of Ammonites köllikeri Zittel, 1868 in Fig. 3B, that was desig-
nated and illustrated by Parent et al. (2011, fig. 32A1). A plaster
cast (FSL.13049) of the lectotype of Ammonites symbolus
illustrated by Zittel (1868, pl. 16, fig. 6a,c) is herein illustrated
for the first time on Fig. 2B.
Emended diagnosis: Small to large, widely umbilicate,
dimorphic Himalayitidae. Small, lappeted microconchs with
two ornamental stages composed of (i) a perisphinctid stage in
juvenile whorls with rigid, straight to prorsiradiate, bifurcate
ribs — subrounded whorl section with convex flanks — ribs
cross the venter but a ventral groove progressively appears and
form a shallow or deep sulcus — (ii) a microcanthum stage
in inter mediate and adult whorls characterized by regular and/
or irregular, rigid or dense, alternation of variable number of
single, bifurcate and sporadically trifurcate, ribs — greater or
lesser thickenings, punctiform tubercles at the furcation point
— narrow venter with a ventral groove or deep sulcus bor-
dered by more or less thickenings on latero-ventral ends of
ribs — subrectangular to laterally compressed subhexagonal
whorl section. Macroconchs are large and develop (iii) an
adult ornamental stage. Two extreme morphotypes are recog-
nized (i) koellikeri morphotype characterized by compressed,
subrectangular, higher than wide, whorl section — flat venter
covered by a more or less attenuated ventral band — micro
canthum stage with enlarged ribs on the umbilical shoulder
combined with more or less numerous trifurcate primary ribs
— (ii) symbolum morphotype characterized by a depressed
subhexagonal whorl section — rigid microcanthum stage with
strong lateral tubercles and thickenings on the ventral shoulder
delimiting a narrow sulcus. Himalayitid suture line marked by
large trifid lateral
lobe
.
Material: Forty-one specimens MPP-CHR.29/1, MPP-
CHR.34/1, MPP-CHR.34/2, MPP-CHR.37/1, MPP-
CHR.37/2, MPP-CHR.37/3, MPP-CHR.45/1, MPP-CHR.50/1,
MPP-CHR.50/2, MPP-CHR.50/3, MPP-CHR.51/1, MPP-
CHR.51/2, MPP-CHR.53/2, MPP-CHR.53/4a and b, MPP-
CHR.54/1a and b, MPP-CHR.60/3, MPP-CHR.60/4,
MPP-CHR.60/6, MPP-CHR.60/7, MPP-CHR.60/1, MPP-
CHR.60/2, MPP-CHR.62/1, MPP-CHR.63/1, MPP-
CHR.67/4, MPP-CHR.67/7, MPP-CHR.67/9, MPP-CHR.68/1,
MPP-CHR.68/2, MPP-CHR.82/1, MPP-CHR.92/2, MPP-
CHR.92/4, MPP-CHR.92/5, MPP-CHR.92/10, MPP-
CHR.92/11, MPP-CHR.92/13, MPP-CHR.92/28, MPP-
CHR .87/1, MPP-CHR.98/1, MPP-CHR.R100/6.
Measurements (mm): See Table 2 in Supplementary data.
Description: Specimens MPP-CHR.67/4 (Fig. 4K), MPP-
CHR.82/1 (Fig. 4N), MPP-CHR.87/1 (Fig. 4R), MPP-
CHR.92/11, MPP-CHR.92/28 (Fig. 4Q), MPP-CHR.98/1
(Fig. 4U) and MPP-CHR.R100/6 perfectly match the morpho-
logical and ornamental features of the lectotype of
M. microcanthum.
MPP-CHR.29/1 (Fig. 4C) and MPP-CHR.50/3 (Fig. 4D)
differ from the lectotype by their smaller size and less evolute
coiling. These specimens show a long perisphinctid stage that
extends over the phragmocone with a ventral groove or sulcus.
The microcanthum stage is attenuated and limited to the end of
the body chamber.
Specimens MPP-CHR.34/1 and MPP-CHR.45/1 are frag-
ments of medium size body chambers characterized by a rigid
microcanthum stage.
Specimen MPP-CHR.37/3 is a large size, subcomplete
specimen characterized by a suboval whorl section. The orna-
mentation of the inner whorls matches the morphological and
ornamental features of the lectotype. The ornamentation of the
body chamber is composed of spaced, sharp, single, some-
times bifurcate ribs with smooth interspaces. Lateral tubercles
occur at the furcation point.
Bivariate diagrams of the dimensional growth parameters of
the shell (U, Wh in function of D) of the well-preserved
specimens show homogeneous scatters around the mean curve
with R² still high > 0.9 in every case (see Figs. S1– S4 in Sup-
plementary data). The growth of those parameters is isometric
and harmonic, and corresponds to the relationship Y = bD.
Preservation of our specimens as crushed internal moulds
prevent us from studying Wb/D and Wb/Wh ratios.
Dimorphism (Fig. 5): Bulot et al. (2014) outlined that the
understanding of M. microcanthum was limited to the mor-
phology of the lectotype and its accepted variability illustrated
by Spanish material. In this regard, M. fraudator (Zittel) as
well as Berriasella (Picteticeras) subeudichotoma Nikolov
perfectly match the lectotype of M. microcanthum. Bulot et al.
(2014) also assumed that the great majority of specimens
referred to Corongoceras from the Betic Cordillera by Tavera
(1985) belongs to Micracanthoceras. The biometric analysis
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investigated on Tavera’s measurements strengthen the conten-
tion that the many typological Corongoceras species from the
Betic Cordillera fit into a homogeneous sample similar to that
of M. microcanthum (see Supplementary data). Bivariate
diagrams of the dimensional growth parameters of the shell
(D, U, Wh and Wb in function of D) show homogeneous
scatters around the mean curve and the growth of those para-
meters is isometric and harmonic. We therefore consider that
M. microcanthum is the senior subjective synonym of M. (C.)
flexuosum Tavera, M. (C.) radians Tavera, M. (C.) minor
Tavera and M. (C.) leanzai Tavera. Since several specimens
illustrated by Tavera (1985), such as the one on pl. 22, fig. 2,
Fig. 5. Supposed dimorphism and intraspecific variations observed in Micracanthoceras micro canthum (Oppel, 1865). Note that microconchs
show different rates of shell morphogenesis (tachymorphic versus bradymorphic variants), while macroconchs range between two extreme
(robust versus slender) morphologies related to the Buckman’s first law of covariation.
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bear short lateral lappets, it seems obvious that the morpho-
logy of the lectotype of M. microcanthum corresponds to
a microconch form.
From a re-examination of the literature and the material
collected at Charens (see for example MPP-CHR.37/3 on
Fig. 2C), we consider that the macroconchs of M. micro
canthum match the morphology of Zittel’s paratype of
M. microcanthum and the lectotype of M. koellikeri. Both
forms show similar inner whorls to that of microconchs of
M. microcanthum, but differ by their large size and the addi-
tion of an adult ornamental stage. We support the supposition
that both morphotypes range between two extreme morpholo-
gies, namely robust and slender poles, related to the Buck-
man’s first law of covariation between shell shape and
ornamen tation. The macroconch koellikeri morphotype is
characte rized by large size, moderately evolute coiling, com-
pressed, subrectangular, higher than wide, whorl section —
with a flat venter covered by a more or less attenuated ventral
band — and develops a robust microcanthum stage in the
adult. The other macro conch morphotype corresponds to the
typological species Micra cantho ceras symbolum and
M. lamberti. This morphotype is characterized by a smaller
size than the other one, evolute coiling, robust ornamen tation,
depressed subhexagonal whorl section and spaced, rigid
microcanthum stage in the adult. As herein understood, the
species Djurjuriceras mutari Tavera, Djurjuriceras mediter
raneum Tavera and Djurjuriceras sinuosum Tavera link the
two macroconch poles of M. microcanthum by their eleva ted,
sub-hexagonal adult whorl section with simplified micro
canthum stage (see for example specimen MPP-CHR.37/3
on Fig. 2C).
The co-occurrence of the two micro- and macroconch
morpho logies on the northern (Czech Republic, Hungary,
Austria, south-east France) and southern (Morocco, Algeria,
Sicily) margins of the Mediterranean-Caucasian Subrealm sup-
port the dimorphism observed in M. micro
canthum.
Remarks: A close exami na tion of the literature convinced
us that great confusions exist between M. microcanthum and
the many Mediterranean-Caucasian forms referred to Aulaco
sphinctes Uhlig, 1910 by Sapunov (1979), Nikolov (1982),
Tavera (1985), Boughdiri et al. (2005) and Benzaggagh et al.
(2010). In these works, the specimens referred to Aulaco
sphinctes either belong to:
• juveniles of M. microcanthum (compare Tavera 1985, pl.
17, fig. 3, 4a,b and Benzaggagh et al. 2010, fig. 5d,e with
MPP-CHR.50/3 on Fig. 4D) or;
• gracile microconchs of M. microcanthum (compare
Nikolov 1982, pl. 78, fig. 5 and Sapunov 1979, pl. 59, fig. 1,
2a,b with MPP-CHT.92/13 and MPP-CHR.92/10 on Fig. 4S
and 4T).
The type specimens of Aulacosphinctes sulcatus from the
Betic Cordillera differs from the lectotype of M. microcanthum
by its long perisphinctid stage with deep ventral sulcus, and
a limited Microcanthum stage at the end of the adult whorl.
The general features of A. sulcatus therefore coincide with
a delayed shift of the ontogenetic sequence of M. micro canthum.
This form co-occurs with the typical M. micro canthum at the
base of the Microcanthum Zone in southern Spain as well as at
Charens where similar specimens were collected (see for
example MPP-CHT.29/1 on Fig. 4C). We therefore assume
that A. sulcatus is a bradymorphic variant of M. microcanthum
in the sense of Beznosov & Mitta (1995) (see also fig. 5). In
this regard, the morphological and ornamental features of the
Spanish Aulacosphinctes specimens described by Tavera
(1985) fall well into the range of variation of the Spanish
M. microcanthum (see Supplementary data).
Olóriz (1978) previously introduced new Aulacosphinctes
species from the Betic Cordillera that are A. trifidus, A. quadri
and A. berriaselliformis. As pointed out by that author, these
species co-occur in the uppermost Lower Tithonian, and
closely resemble Burckhardticeras peroni (Roman, 1936), but
they could be distinguished by the presence of a ventral groove
at the sub-adult and adult stages. Olóriz (1978) pointed out
that B. peroni is a microconch form that develops a thin ven-
tral groove at the end of the adult whorl. Olóriz noticed that
the species shows a wide range of intraspecific variation, but
that it had, unfortunately, never been evaluated. One may won-
der if the presence of a ventral groove might not result from
extreme intraspecific variation of B. peroni or reflect a phy-
letic link with M. microcanthum. In any case, it is clear that the
genus Aulacosphinctes should be limited to its type species
Ammonites mörikeanus Oppel, 1863 and those closely allied
Indian and Malagasy taxa described by Oppel (1863), Uhlig
(1910), Besairie (1936), Collignon (1960) and Fatmi (1973).
Occurrence: M. microcanthum occurs between beds 25 and
76, M. microcanthum Zone, Upper Tithonian. The species has
been reported from the Czech Republic, Austria, Hungary,
Germany, Bulgaria, Italy (Umbria, Sicily), South East France,
southern Spain, Morocco, Tunisia and Algeria.
Genus Protacanthodiscus Spath, 1923
Type species: Hoplites andreaei Kilian, 1889, by original
designation.
Remarks: A detailed revision of the ontogeny, variability,
dimorphism and age of the type species Protacanthodiscus
andreaei was recently given by the authors of the present
paper and the reader is referred to it (see Frau et al. 2015).
Protacanthodiscus andreaei (Kilian, 1889)
Fig. 6A,C
Macroconchs [M]
1889 Hoplites Andreaei – Kilian, p. 670, pl. 32, fig. la,b
1907 Hoplites (Acanthodiscus) Andreaei var. punica – Pervinquière,
p. 38, pl. 2, fig. l2a,b
1939 Berriasella Andreaei (Kilian) – Mazenot, p. 96, pl. 12, fig. la,b
(= Kilian, 1889, pl. 32, fig. la,b)
non 1939 Berriasella Andreaei (Kilian) – Mazenot, p. 96, pl. 13,
fig. 4a,b [= Jabronella (Erdenella) isare (Pomel) nom. correct.]
non 1960 Berriasella (Protacanthodiscus) aff. andreaei (Kilian) –
Collignon, pl. 165, fig. 665 (= Himalayitidae gen. et sp. indet. juv.)
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Fig. 6. Protacanthodiscus andreaei (Kilian, 1889): A1–2 — Plaster cast of the holotype (FSL.13056); B — MPP–CHR.79/4 [M] bed 89;
C — MPP–CHR.21/25 [m] bed 92. Boughdiriella chouetense Frau, Bulot & Wimbledon, 2015 nom. correct.: D — MPP–CHR.21/24 [m]
bed 92; E — plaster cast of MPP–CHR.21/33 [?M] bed 92; F — MPP–CHR.19/1 [m] bed 90; G — MPP–CHR.21/35 [m] bed 92. Pratumi discus
elsae Bulot, Frau & Wimbledon, 2014: H1–2 — MPP–CHR.21/43 [?m] bed 92. Busnardoiceras busnardoi (Le Hégarat, 1973):
I — MPP–CHR.19/1 [m] bed 90; J — MPP–CHR.21/2 [m] bed 92; K — MPP–CHR.21/16 [m] bed 92; L — MPP–CHR.21/14 [m];
M — MPP–CHR.19/3 [m] bed 90; N — MPP–CHR.21/15 [m] bed 92; O1–2 — MPP–CHR.21/42 [m] bed 92; P1–2 — MPP–CHR.19/4 [M]
bed 90; Q — plaster cast of MPP–CHR.21/38 [M] bed 92; R — plaster cast of MPP–CHR.21/12 [M] bed 92; S — MPP–CHR.21/36 [M]
bed 92; T1–2 — MPP–CHR.21/26 [M] bed 92; U — MPP–CHR.21/18 [M] bed 92; V — MPP–CHR.21/7 [M] bed 92. Scale bar is 10 mm.
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non 1960 Berriasella (Protacanthodiscus) aff. andreaei var. varia
bilis – Collignon, pl. 165, figs. 666, 667 (= Himalayitidae gen.
et sp. indet. juv.)
? 1960 Protacanthodiscus andreaei (Kilian) – Christ, p. 125, pl. 9,
fig. 5
1979 Protacanthodiscus andreaei (Kilian) – Sapunov, pl. 57, fig. 6
1982 Protacanthodiscus andreaei (Kilian) – Nikolov, p. 211, pl. 73,
fig. 2a,b (= Kilian, 1889, pl. 32, fig. 1a,b), pl. 74, fig. 2
(= Sapunov, 1973, pl. 57, fig. 6)
1985 Neoperisphinctes falloti (Kilian) – Tavera, p. 132, pl. 16,
fig. 10a,b
1985 Durangites gigantis – Tavera, p. 162, pl. 18, figs. 8, 9,
text-fig. 12A, B
1985 Protacanthodiscus andreaei (Kilian) – Tavera, p. 198, pl. 26,
figs. 5a,b, 6, text-fig. 15H
1985 Protacanthodiscus sp. 1 gr. andreaei (Kilian) – Tavera, p. 199,
pl. 26, figs. 1a,b, 2, text-figs. 12H, 15E, I
1985 Protacanthodiscus berriasensis – Tavera, p. 206, pl. 28,
fig. 1a,b, text-fig. 15D
1985 Protacanthodiscus coronatus – Tavera, p. 200, pl. 27, fig. 1a,b,
text-fig. 15A
1985 Protacanthodiscus darwini – Tavera, p. 203, pl. 26, figs. 3, 4,
text-fig. 15B
1985 Protacanthodiscus nodusus – Tavera, p. 202, pl. 27, fig. 2,
text-fig. 15C
1994 Protacanthodiscus (Protacanthodiscus) andreaei (Kilian) –
Boughdiri, p. 147, pl. 3, figs. 1a,c (= Kilian, 1889, pl. 32,
fig. la,b), 3a,b, pl. 4, fig. 2
1994 Protacanthodiscus (Protacanthodiscus) n. sp. A – Boughdiri,
p. 208, pl. 3, fig. 5
1994 Protacanthodiscus (Protacanthodiscus) n. sp. 7 – Boughdiri,
p. 176, pl. 3, fig. 2
1994 Protacanthodiscus (Protacanthodiscus) n. sp. 11 – Boughdiri,
p. 186, pl. 3, fig. 6
1994 Protacanthodiscus (Protacanthodiscus) n. sp. 12 – Boughdiri,
p. 188, pl. 4, fig. 1
1994 Protacanthodiscus (Protacanthodiscus) coronatus (Tavera) –
Boughdiri, p. 151, pl. 4, figs. 6, 7
1998 Durangites (Protacanthodiscus) andreaei (Kilian) – Enay et al.,
figs. 2.16a,b (= Kilian, 1889, pl. 32, fig. la,b)
1998 Durangites (Protacanthodiscus) apertus (Tavera) – Enay et al.,
figs. 2.13, 14
2010 Protacanthodiscus apertus (Tavera) – Benzaggagh et al., fig. 5i
2010 Durangites humboldti Burckhardt – Benzaggagh et al., fig. 5n
non 2013 Protacanthodiscus coronatus (Tavera) – Szives & Fözy,
p. 301, pl. 1, figs. 2, 4 ( = Himalayitidae gen. nov. et sp. nov.)
non 2013 Protacanthodiscus andreaei (Kilian) – Szives & Fözy,
p. 300, pl. 5, fig. 4a,b, pl. 6, fig. 5a,b, pl. 8, fig. 10a,b
(= Himalayitidae gen. nov. et sp. nov.)
2015 Protacanthodiscus andreaei (Kilian, 1889) – Frau et al., p. 125,
figs. 7A,C
Microconchs [m]
1983 Durangites acanthicus Burckhardt – Cecca et al., p. 128, pl. 5,
fig. 1a,b
1985 Durangites singularis – Tavera, p. 168, pl. 17, figs. 6, 7, 8a,b,
9a,b, text-fig. 12E
1985 Durangites sutneroides – Tavera, p. 168, pl. 18, figs. 1, 2, 3, 5
non 1985 Durangites sutneroides – Tavera, p. 168, pl. 18, figs. 4
(= Spiticeratinae indet. juv.)
1985 Durangites heilprini (Aguilera) – Tavera, p. 164, pl. 18, figs. 6,
7, text-fig. 12G
1985 Durangites acanthicus Burckhardt – Tavera, p. 156, pl. 18,
figs. 10a,b, 11, text-fig. 12C
1985 Durangites vulgaris Burckhardt – Tavera, p. 157, pl. 18,
figs. 12, 13a,b, text-fig. 12I
1985 Durangites apertus –Tavera, p. 161, pl. 18, figs. 14, 15a,b,
text-fig. 12F
1985 Durangites cf. astillerensis Imlay – Tavera, p. 166, pl. 18,
fig. 16, text-fig. 12D
1994 Protacanthodiscus (Protacanthodiscus) n. sp. 3 – Boughdiri,
p. 168, pl. 3, fig. 7
1994 Protacanthodiscus (Protacanthodiscus) n. sp. 4 – Boughdiri,
p. 170, pl. 2, fig. 17
1994 Durangites (Durangites) humboldti (Burckhardt) – Boughdiri,
p. 99, pl. 1, figs. 33a,b, 34
1994 Durangites (Durangites) juanense (Cantú-Chapa) – Boughdiri,
p. 106, pl. 1, fig. 35
1998 Durangites (Durangites) juanensis Cantú-Chapa – Enay et al.,
figs. 2-6, 7 ( = Boughdiri, 1994, pl. 1, fig. 35)
1998 Durangites (Durangites) humboldti Burckhardt – Enay et al.,
figs. 2.8, 9, 10a,b
1998 Durangites (Protacanthodiscus) sp. – Enay et al., figs. 2.11
2010 Protacanthodiscus cf. andreaei (Kilian) – Ivanov et al., pl. 3,
fig. 3
2010 Durangites aff. fusicostatus Burckhardt – Ivanov et al., pl. 3,
fig. 1a,b
2010 Durangites cf. vulgaris Burckhardt – Ivanov et al., pl. 3, fig. 2a,b
2010 Durangites singularis Tavera – Ivanov et al., pl. 3, fig. 4
2010 Corongoceras cf. symbolum (Oppel) – Benzaggagh et al., Figs.
5j,m
2013 Micracanthoceras (Corongoceras) koellikeri (Oppel) – Szives
& Fözy, p. 300, pl. 4, fig. 1, ? pl. 6, fig. 7, pl. 8, fig. 3
2015 Protacanthodiscus andreaei (Kilian, 1889) – Frau et al., p. 125,
figs. 5A,L, 6A,E
Type: The holotype is specimen no. EM.1930 from the
De Verneuil collection, originally drawned by Kilian (1889,
pl. 32, fig. la,b). A plaster cast (FSL.13056) is herein illus-
trated in Fig. 6A.
Material: Eleven specimens MPP-CHR.79/4, MPP-
CHR.19/1, MPP-CHR.19/5, MPP-CHR.21/2, MPP-CHR.21/3,
MPP-CHR.21/6, MPP-CHR.21/19, MPP-CHR.21/29, MPP-
CHR.21/32, MPP-CHR.21/33, MPP-CHR.21/39.
Measurements (mm): See Table 3 in Supplementary data.
Description: The material collected at Charens is identical
to specimens described at Le Chouet (Frau et al. 2015).
P. andreaei corresponds to medium to large size Himalayitidae
with a moderately open and deep umbilicus. The whorl section
is subcircular with convex flanks on juveniles that become
progressively subrectangular to subhexagonal on the adults.
Ornamentation is composed of bifurcate and trifurcate ribs
with a tubercle at the furcation point. Some ribs can be looped
on the ventral shoulder. On the body chamber, bifurcate ribs
end on the ventral shoulder with a latero-ventral tubercle on
the first, or occasionally the second, branch. The venter is
marked by a flattened band or a shallow sulcus depending on
the shape of the whorl section.
Remarks: The koellikeri macroconch morphotype of
M. microcanthum share great affinities with macroconchs of
Protacanthodiscus andreaei (Kilian, 1889) (see for example
Frau et al. 2015, fig. 7a,c). P. andreaei can be distinguished by
its more involute coiling and the occurrence of strong tuber-
cles at the furcation point and on the end of the posterior
branches of the bifurcate ribs. Following the revision of
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P. andreaei by Frau et al. (2015), it seems obvious that
P. andreaei was derived from M. microcanthum in the upper
part of the M. microcanthum Zone (= M. fischeri Subzone
sensu Wimbledon et al. 2013).
Occurrence: The first occurrence of P. andreaei is found in
bed 89 at Charens. This occurrence is identical to that docu-
mented at Le Chouet (Wimbledon et al. 2013; Frau et al.
2015).
P. andreaei has been reported from south-east
Spain
and France, Italy, Bulgaria, Tunisia and Morocco.
Genus Boughdiriella Frau, Bulot & Wimbledon, 2015
Type species: Boughdiriella chouetensis Frau, Bulot &
Wimbledon, 2015, by original designation
Remarks: The genus was recently introduced by the authors
of the present contribution to distinguish small Himalayitidae
forms which had previously been referred to the “Medi-
terranean Durangites” (see discussion in Frau et al. 2015).
Boughdiriella chouetense Frau, Bulot & Wimbledon, 2015
nom. correct.
Figs. 6D – G
2015 Boughdiriella chouetensis – Frau, Bulot & Wimbledon, p. 123,
figs. 4A,I [cum. syn.].
Type: The holotype is specimen no. MPP-CHT.21/65 from
the Frau/Bulot collection. It was illustrated by Frau et al.
(2015, fig. 4A).
Material: Seven specimens MPP-CHR.21/4, MPP-
CHR.21/8, MPP-CHR.21/11, MPP-CHR.21/24, MPP-
CHR.21/27, MPP-CHR.21/35, MPP-CHR.21/40.
Measurements (mm): See Table 4 in Supplementary data.
Description: The material collected at Charens is identical
to that described from Le Chouet (Frau et al. 2015). B. choue
tense is composed of small-sized Himalayitidae with a dis-
coidal and moderately evolute shell. The whorl section is
sub-circular in inner whorls and compressed, sub-oval whorl
section in adult. The umbilical wall is low. The ornamentation
is composed of rigid, straight, simple ribs in the inner whorls.
From D~15 mm to the end of the body chamber, ribs are
flexuous, prominent and bifurcated on the upper third of the
flanks. Sporadic trifurcate and intercalated ribs occur. Ribs
delimit a shallow furrow on the venter that is progressively
attenuated at the end of the adult whorl.
Remarks: As discussed by Frau et al. (2015), Boughdiriella
and Protacanthodiscus share strong affinities but the latter
genus can be distinguished by its subrectan gular to subhexa-
gonal whorl section combined with tubercu lation at almost all
ontogenetic stages in both micro- and macroconch forms.
R. Enay (comm. pers. 2015) expressed doubt about the inter-
pretation of the peristome in Boughdiriella. He suggested that
the genus may group only microconchs. Therefore, one may
wonder if B. chouetense might not cor respond to an extreme
microconch morphology of P. andreaei. A detailed biometric
analysis is currently in progress to decipher the range of
variability of both species.
Occurrence: B. chouetense only occurs in bed 92 at Charens,
topmost Andreaei Zone, Upper Tithonian. The vertical range
of the species is shorter than at Le Chouet. B. chouetense is
reported from
south-east France
and Bulgaria
but its occur-
rence in Tunisia, Turkey, Italy and Morocco cannot be
excluded (see discussion in Frau et al. 2015).
Genus Pratumidiscus Bulot, Frau & Wimbledon, 2014
Type species: Pratumidiscus elsae Bulot, Frau &
Wimbledon, 2014, by original designation.
Remarks: The genus Pratumidiscus was introduced by
Bulot et al. (2014) based on a single specimen from the upper-
most Tithonian of Le Chouet. The genus was considered to be
endemic in South East France but its general features closely
resemble those of the Boreal lineage Riasanella Mitta, 2011 —
Riasanites Spath, 1923 and therefore it was considered as the
probable Tethyan rootstock.
On the other hand, P. elsae shares strong affinities with the
gracile morphotype of P. andreaei that was recently described
by Frau et al. (2015). P. elsae can be distinguished by its more
evolute coiling, irregular ribbing and loss of the lateral tubercles
on the body chamber. In this regard, we assume that P. elsae
was derived from the gracile morphotype of P. andreaei such
as those illustrated by Frau et al. (2015, Fig. 6C and D).
Pratumidiscus elsae Bulot, Frau & Wimbledon, 2014
Fig. 6H
2014 Pratumidiscus elsae – Bulot, Frau & Wimbledon, p. 122, fig. 6
Type: The holotype is specimen no. MPP-CHT.19/5 from
the Frau/Bulot collection. It was illustrated by Bulot et al.
(2014, fig. 6).
Material: A single specimen MPP-CHR.21/43.
Measurements (mm): See Table 5 in Supplementary data.
Description: Small fragment of a body chamber with a sub-
rectangular whorl section and tabulate venter. Ribbing com-
posed of irregular single, bifurcate and intercalate ribs. All ribs
thicken on the ventral shoulder into radially elongated bullae
that delimit a ventral groove.
Remarks: The specimen at our disposal matches well the
adult features of P. elsae. It differs in its smaller size and
slightly rursiradiate ribs. These features suggest that this
specimen corresponds to a microconch form. New specimens
are urgently needed to document the range of variation and
dimorphism of P. elsae.
Occurrence: P. elsae is found in bed 92 at Charens, topmost
P. andreaei Zone, Upper Tithonian. This matches the occur-
rence of the holotype found at Le Chouet (Bulot et al. 2014).
P. elsae is endemic to south-east France.
Family Neocomitidae Salfeld, 1921
Genus Busnardoiceras Tavera, 1985
Type species: Parapallasiceras busnardoi Le Hégarat,
1973, by original designation.
556
FRAU, BULOT, WIMBLEDON and IFRIM
GEOLOGICA CARPATHICA
, 2016, 67, 6, 543 – 559
Remarks: A revised account of the genus Busnardoiceras
and its type species was recently provided by the authors of
the present contribution to which the reader is referred (Bulot
et al. 2014).
Busnardoiceras busnardoi (Le Hégarat, 1973)
Figs. 6 I –V
Macroconchs [M]
1939 Berriasella ciliata Schneid – Mazenot, p. 37, pl. 1, fig. 1a,b
1973 Parapallasiceras busnardoi – Le Hégarat, p. 47, pl. 3, figs. 4, 5
(= Mazenot, 1939, pl. 1, fig. 1a,b), ? pl. 38, fig. 2
2014 Busnardoiceras busnardoi (Le Hégarat) – Bulot et al., p. 124,
fig. 8F,F’
Microconchs [m]
1982 Fauriella shipkovensis (Nikolov & Mandov) – Nikolov, p. 109,
pl. 31, fig. 1
1985 Berriasella (Berriasella) tithonica – Tavera, p. 237, pl. 33,
figs. 4a,b, 5, 6
? 2001 Parapallasiceras busnardoi Le Hégarat – Wippich, p. 78,
pl. 20, fig. 2
2014 Busnardoiceras busnardoi (Le Hégarat) – Bulot et al., p. 124,
figs. 8A,E
Type: The holotype is specimen no. UJF-ID.563 from the
Mazenot collection. It was illustrated by Mazenot (1939, pl. 1,
fig. 1a,b).
Material: Nineteen specimens MPP-CHR.19/1, MPP-
CHR.19/3, MPP-CHR.19/4, MPP-CHR.21/2, MPP-
CHR.21/7, MPP-CHR.21/10, MPP-CHR.21/12,
MPP - CHR.21/13,
MPP-CHR.21/14,
MPP-CHR.21/15,
MPP-CHR.21/16, MPP-CHR.21/17, MPP-CHR.21/18,
MPP-CHR.21/26, MPP-CHR.21/28, MPP-CHR.21/36,
MPP-CHR.21/37, MPP-CHR.21/38, MPP-CHR.21/42.
Measurements (mm): See Table 6 in Supplementary data.
Description: The material collected at Charens is similar to
that from Le Chouet recently described by Bulot et al. (2014).
B. busnardoi comprises small to medium size, planulate neo-
comitid forms with high and compressed, subrectangular
whorl section and moderately evolute shallow umbilicus.
The ornamentation is composed of straight to slightly prorsi-
radiate, bifurcate ribs on the phragmocone. Ribs bifurcate on
the upper third of the flanks. The ornamentation of the body
chamber is composed of dense slightly sinuous, irregular,
bifurcate and intercalate rib. Sporadic virgatotome and simple
ribs occur. Ribs are interrupted on the ventral shoulder by
small punctiform tubercles delimiting a shallow groove on
the inner whorls. In macroconchs, ribs cross the venter.
Remarks: The newly collected material from Charens
shows that the innermost whorls of B. busnardoi match well
the juvenile and intermediate ornamental stages of microconchs
of P. andreaei as described by Frau et al. (2015).
B. busnardoi can easily be distinguished since the species
never develops lateral tubercle at the furcation point.
B. busnardoi is further characterized by its adult neocomitid
morphology. We therefore believe that B. busnardoi
was derived from P. andreaei since both species co-occur in
the topmost P. andreaei Zone in the Le Chouet and Charens
sections.
Occurrence: B. busnardoi occurs between beds 90 and 92,
topmost P. andreaei Zone, Upper Tithonian. This occurrence
matches well the vertical range documented at Le Chouet by
Wimbledon et al. (2013) and Bulot et al. (2014). The revised
synonymy list of B. busnardoi shows that representatives
of the species are more widespread than it was previously
thought. Examination of the type specimen of Berriasella
tithonica Tavera, 1985 shows that it is a junior subjective
synonym of B. busnardoi. Therefore, the species occurs in
Bulgaria, SE France, southern Spain and probably in Morocco.
Conclusion
Based on bed-by-bed sampling, we document the precise
distribution of Micracanthoceras microcanthum at Charens.
This confirms its high stratigraphic value for defining the base
of the Upper Tithonian. As herein understood, the lower
boundary of the M. microcanthum Zone is fixed at the FAD of
the index species at Charens (i.e. bed 25). According to the
preliminary calpionellid zonation from Charens (Wimbledon
& Reháková unpublished data), the FAD of M. microcanthum
falls within the upper part of the Chitinoidella Standard Zone,
thus confirming similar result achieved in Spain (Pruner et al.
2010) and Morocco (Benzaggagh et al. 2010). A comparative
analysis of the ammonite and calpionellid zonation is still
ongoing and will shed new light on this problem. In this
regard, the occurrence of ammonites below the FAD of
M. microcanthum is confirmed at Charens which indicates the
B. ponti Zone sensu Enay & Geyssant (1975). This fauna will
be described elsewhere (Frau et al. in prep.).
From our detailed ontogenetic and biometric studies, we
assume that M. microcanthum shows a wide range of variation
that represents sexual and non-sexual polymorphism. Hetero-
chronic shifts in development are documented in microconchs
while macroconchs range between two extreme morpho-
logical poles related to the Buckman’s first law of covariation
(Fig. 5).
The vertical range of the age-diagnostic species Protacantho
discus andreaei is rather similar to that described at Le Chouet
(Frau et al. 2015). The FAD of this species typifies the lower
boundary of the P. andreaei Zone sensu Wimbledon et al.
(2013). The radiation of the Himalayitidae in the P. andreaei
Zone is confirmed by the co-occurrence of the genera
Protacanthodiscus, Boughdiriella and Pratumidiscus. As
a result, the M. microcanthum Zone and the P. andreaei Zone
are two biochronozones in the sense of Callomon (1985) since
they are defined by the FAD of their index-species and charac-
terized by their faunal assemblages. The two zones could
be retained as workable on a larger geographical scale
since both index species are widespread across the
557
UPPER TITHONIAN AMMONITES FROM CHARENS (DRÔME, FRANCE)
GEOLOGICA CARPATHICA
, 2016, 67, 6, 543 – 559
Medi terranean- Caucasian Subrealm of the Tethyan Realm
sensu Westermann (2000) (Fig. 7).
A close examination of a new collection of the basal
Neocomitidae Busnardoiceras busnardoi shows that its inner
whorls closely resemble those described by Frau et al. (2015)
in P. andreaei. An ongoing revision of the early Berriasian
Perisphinctoidea taxa originally reported at Le Chouet by
Wimbledon et al. (2013) will shed new light on the relation-
ships between B. busnardoi and other basal Neocomitidae.
Nevertheless, there is no doubt that B. busnardoi represents an
important element that links the radiation of the Neocomitidae
from the Himalayitidae at the base of the B. jacobi Zone
auctorum.
Acknowledgements: This work is a contribution of the
Berriasian Working Group (BWG) of the International Sub-
commission on Cretaceous Stratigraphy (I.S.C.S.). Jean-Marc
Chabal, mayor of Charens, is gratefully acknowledged for
providing access to the Charens section. Cyprien Lanteaume
(Aix-Marseille Université) and Emile Hourqueig (Toulouse)
are acknowledged for their help in collecting ammonites from
the Charens section and for their continuous support during
the fieldwork seasons of the first author (C.F.). The authors are
indebted to Daniela Reháková (Comenius University) for
providing the preliminary results on the calpionellid assem-
blages for the Charens section and Emmanuel Robert (Univer-
sity of Lyon I) for providing access to the collections under his
care. Horacio Parent (Universidad Nacional de Rosario) is
gratefully acknowledged for valuable discussion, photographs
and measures of Ammonites Koellikeri. We thank the two
anonymous reviewers for their useful comments. Warm thanks
are due to the Jean-Pierre Lopez, Martine Broin and their
families (Le Chouet and Les Près, France) for their hospitality
and friendship during our fieldwork seasons, and to Elsa
Schnebelen-Bulot (Saint-Privat-de-Champclos) for her
continuous support during this study.
References
Arkell W.J., Kummel B. & Wright C.W. 1957: Mesozoic Ammonoidea.
In: Moore R.C. (Ed.): Treatise of Invertebrate Paleontology, Part
L, Mollusca 4, Cephalopoda, Ammonoidea. The Geological
Society of America & The University of Kansas Press, New York
& Lawrence, 80–437.
Bahrouni N., Houla Y., Soussi M., Boughdiri M., Ben Ali W., Nasri A.
& Bouaziz S. 2016: Discovery of Jurassic ammonite bearing
series in Jebel Bou Hedma (South-Central Tunisian Atlas).
Implications for stratigraphic correlations and paleogeographic
reconstruction. J. Afr. Earth Sci. 113, 101–113.
Benzaggagh M. 2000: Le Malm supérieur et le Berriasien dans le
Prérif interne et le Mésorif (Rif, Maroc) — biostratigraphie,
lithostratigraphie, paléogéographie et évolution tectono-
sédimentaire. Doc Lab. Géol. Fac. Sci. Lyon 152, 1–347.
Benzaggagh M. & Atrops F. 1997: Stratigraphie et associations de
faune d’ammonites des zones du Kimméridgien, Tithonien et
Berriasien basal dans le Prérif interne (Rif, Maroc). Newsl.
Stratigr. 35, 127–163.
Benzaggagh M., Cecca F. & Rouget I. 2010: Biostratigraphy distri-
bution of ammonites and calpionellids in the Tithonian of the
internal Prerif (Msila area, Morocco). Pal. Zeitschrift 84,
301–315.
Besairie H. 1936: Recherche géologiques à Madagascar. 1. La géologie
du Nord-Ouest. Mém. Acad. Malgache 21, 1–259.
Beznosov N.V. & Mitta V.V. 1995: Polymorphism in the Jurassic
ammonoids. Paleontol. J. 29, 46–57.
Boughdiri M. 1994: Les genres d’ammonites Durangites et Protacan
thodiscus (Tithonien supérieur) dans la Téthys occidentale (SE
Espagne, SE France, Algérie et Tunisie). Stratigraphie,
Paléontologie et Biogéographie. PhD thesis, Université Claude
Fig. 7. Palaeo-biogeographical distribution of the two index species Micracanthoceras microcanthum (Oppel in Zittel, 1868) and
Protacanthodiscus andreaei (Kilian, 1889) during the upper Tithonian (palaeogeographical map of the northern hemisphere modified
after Cecca 1999).
558
FRAU, BULOT, WIMBLEDON and IFRIM
GEOLOGICA CARPATHICA
, 2016, 67, 6, 543 – 559
Bernard Lyon I, 1–268.
Boughdiri M., Olóriz F., Lopez Marques B., Layeb M., De Matos E.
& Sallouhi H. 2005: Upper Kimmeridgian and Tithonian Ammo-
nites from the Tunisian “Dorsale” (NE Tunisia), updated biostra-
tigraphy from J. Oust. Riv. Ital. Paleontol. S. 111, 305–316.
Bujtor L., Krische O. & Gawlick H.-J. 2013: Late Berriasian ammo-
nite assemblage and biostratigraphy of the Leube quarry near
Salzburg (Northern Calcareous Alps, Austria). N. Jb. Geol.
Palaont. Abh. 267, 273–295.
Bulot L.G., Frau C. & Wimbledon W.A.P. 2014: New and poorly
known Perisphinctoidea (Ammonitina) from the Upper Tithonian
of Le Chouet (Les Près, Drôme, SE France). Volumina Jurassica
12, 113–128.
Burckhardt C. 1900: Profils géologiques transversaux de la Cordillère
Argentino-Chilienne. Museo La Plata Sección Geológica y
Mineralógica, Anales II, La Plata, 1–136.
Burckhardt C. 1903: Beiträge zur Kenntnis der Jura, und Kreide-
formation der Cordillera. Palaeontographica 50, 1–144.
Burckhardt C. 1921: Faunas jurasicas de Symon (Zacatecas) (Atlas).
Bol. Inst. Geol. Mexico 33, 32 pl.
Callomon J.H. 1985: Biostratigraphy, chronostratigraphy and all that
— again! Proceedings of the International Symposium on
Jurassic Stratigraphy, Erlangen 1984 3, 611–624.
Cecca F. 1985: On some ammonites from the “Maiolica” of Central
Apennines (Umbria, Marche e Sabina). Boll. Serv. Geol. Italia
103, 133–162 (in Italian with English summary).
Cecca F. 1999: Palaeobiogeography of Tethyan ammonites during the
Tithonian (latest Jurassic). Palaeogeogr. Palaeoclimatol.
Palaeoecol. 147, 1–37.
Cecca F., Cresta S. & Santantonio M. 1983: Ammonites from the
Malm of the Marchigiana Apennines deposited in the Museum
of the Geological Survey of Italy. Boll. Serv. Geol. Italia, 102
109–132 (in Italian with English summary).
Cecca F., Enay R. & Le Hégarat G. 1989: L’Ardescien (Tithonique
supérieur) de la région stratotypique, p. série de référence
et faunes (ammonites, calpionelles) de la bordure ardéchoise.
Doc. Lab. Géol. Fac. Sci. Lyon 107, 1–115.
Christ H.A. 1960: Beiträge zur Stratigraphie und Paläontologie
des Malm von Westsizilien. Schweiz. Paläontol. Abh. 77, 1–44.
Collignon M. 1960: Atlas des fossiles caractéristiques de Madagascar.
Fascicule VI (Tithonique). Serv. Géol. Madagascar, Tananarive,
1–175.
Dacqué E. 1934: Wirbellose des Jura. In: Gürich G. (Ed.): 7. Liefe-
rung. Leitfossilien. Borntraeger, Berlin, 1–582.
De Wever P., Geyssant J.R., Azéma J., Devos I., Duée G., Manivit H.
&Vrielynck B. 1986: La coupe de Santa Anna (Zone de Sciacca,
Sicile): une synthèse biostratigraphique des apports macro-,
micro- et nannofossiles du Jurassique supérieur et Crétacé
inférieur. Rev. Micropal. 29, 141–186.
Eliáš M. & Vašíček V. 1995: Early Berriasian ammonites from the
Štramberk Limestone of Kotouč quarry (Outer Carpathians,
Silesian Unit, Štramberk, Czech Republic).
Bull. Czech Geol.
Surv.
70, 1–32.
Enay R. & Geyssant J.R. 1975: Faunes tithoniques des chaînes
bétiques
(Espagne méridionale). Mém. Bur. Rech. Géol. Min. 86,
39
–
55.
Enay R., Boughdiri M. & Le Hégarat G. 1998: Durangites, Prota
canthodiscus (Ammonitina) et formes voisines du Tithonien
supérieur-Berriasien dans la Téthys méditerranéenne (SE France,
Espagne, Algérie et Tunisie). C.R. Acad. Sci., Paris, Série 2a,
Sciences de la Terre et des Planètes 327, 425–430.
Fallot P. & Termier H. 1923: Ammonites nouvelles des îles Baléares.
Tra. Mus. Nac. Ci. Nat., Madrid, Série Geológica 32, 1–83.
Fatmi A.N. 1973: Late Jurassic and early Cretaceous (Berriasian)
ammonites from Shaikh Budin Hills, D.I. Khan, N.W.F.P.,
Pakistan. Rec. Geol. Surv. Pak. 21, 2, ii+1–22.
Fatmi A.N. & Zeiss A. 1999: First Jurassic and Lower Cretaceous
(Berriasian) ammonites from the Sembar Formation (Belemnite
shales). Windar Nai. Lasbela — Balochistan, Pakistan. Geol.
Surv. Pak. Mem. 19, 1–114.
Ferry S. & Grosheny D. 2013: Turbidites et brèches carbonatées
(Jurassique et Crétacé du bassin subalpin). Livret-guide excur-
sion ASF-GFC (20–23 novembre 2013). http://hal.archives
ouvertes.fr/hal00908177/fr/, 1–101.
Frau C., Bulot L.G. & Wimbledon W.A.P. 2015: Upper Tithonian
Himalayitidae Spath, 1925 (Perisphinctoidea, Ammonitina)
from Le Chouet (Drôme, France): implications for the systema-
tics. Geol. Carpath. 66, 117–132.
Fözy I. 1990: Ammonite succession from three Upper Jurassic sec-
tions in the Bakony Mts. (Hungary). In: Pallini G., Cecca F.,
Cresta S. & Santantonio M. (Eds): Atti IIº Convegno Fossili,
Evoluzione, Ambiente. Pergola, 323–339.
Fözy I. 1995: Upper Jurassic ammonites from Seno di Guidaloca
(Western Sicily). Hantk.Géczy Jub. 1, 131–143.
Geyssant J.R. 1997: Tithonien. In: Cariou E. & Hantzpergue P. (Eds.):
Biostratigraphie du Jurassique ouest européen et méditerranéen,
p. zonations parallèles et distribution des invertébrés et micro-
fossiles. Elf Aquitaine édition, 97–102
Haas J., Emoke J.E., Gidai L. Kaiser M., Kretzoi M. & Oravecz J.
1985: Geology of the Sümeg Area. Geol. Hungarica 20, 1–250.
Howarth M.K. 1998: Tithonian and Berriasian ammonites from the
Chia Gara Formation of northern Iraq. Palaeontology 35,
597–655.
Imlay R.W. 1939: Upper Jurassic ammonites from Mexico. Geol.
Soc.
Amer. Bull. 50, 1
–
78.
Ivanov M., Stoykova K. & Idakieva V. 2010: New biostratigraphic
data on the Upper Tithonian and Lower Berriasian in the
Krayshte area (Southwestern Bulgaria). Annuaire Univ. Sofia
“St. Kliment Ohridski” 102, 21–42.
Kaiser-Weidich B. & Schaier G. 1989: Stratigraphische Korrelation
von Ammoniten, Calpionellen und Nannoconiden aus Oberjura
und Unterkreider der Nördlichen Kalkalpen. Ecl. Geol. Helv. 83,
353–387.
Khimchiashvili N.G. 1976: Tithonian and Berriasian ammonites of
the Caucasus. Tbilisi, 1–180.
Kilian W. 1889: Etudes paléontologiques sur les terrains secondaires
et tertiaires de l’Andalousie. Mém. Acad. Sci. Inst. France 30,
580–733.
Krantz F. 1928: The fauna from the upper and middle Tithonian of the
Argentina Cordillera in the southern part of the Mendoza
province. Act.
Acad. Nac. Ciencas 10, 5–57 (in Spanish).
Ksiazkiewicz M. 1956: Jurassic and Cretaceous of Bachowice.
Rocz.
Pol. Tow. Geol.
23, 119–405 (in Polish).
Le Hégarat G. 1973: Le Berriasien du sud-est de la France. Doc. Lab.
Géol. Fac. Sci. Lyon 43, 1–576.
Linares A. & Vera J.A. 1966: Precisiones estratigráficas sobre la serie
mesozoica de Sierra Gorda, Cordilleras Béticas (provincia de
Granada). Estud. Geol. 22, 65–99.
Marino M. C. Andreini G., Baldanza A., D’Arpa C., Mariotti N.,
Pallini G., Parisi G. & Petti F. M. 2004: Middle Jurassic–early
Cretaceous integrated biostratigraphy (ammonites, calcareous
nannofossils and calpionellids) of the Contrada Diesi section
(south-western Sicily, Italay). Riv. Ital. Paleontol. S. 110,
357–372.
Mazenot G. 1939: Les Palaeophoplitidae tithoniques et berriasiens du
sud est de la France. Mém. Soc. Géol. France, Nouvelle Série 18
41, 1–303.
Mitta V.V. 2011: Ammonites of Tethyan origin in the Ryazanian Stage
of the Russian Platform: Genus Riasanella gen. nov. Paleontol J.
45, 13–22.
Nicolis E. & Parona C.F. 1886: Stratigraphic and palaeontologic note
on the Upper Jurassic from the Verona province. B. Soc. Geol.
559
UPPER TITHONIAN AMMONITES FROM CHARENS (DRÔME, FRANCE)
GEOLOGICA CARPATHICA
, 2016, 67, 6, 543 – 559
Ital. 4 (1885), 1–96 (in Italian).
Nikolov T.G. 1982: Les Ammonites de la famille Berriasellidae
Spath, 1922. Tithonique supérieur-Berriasien. Acad. Bul. Sci. 10,
1–251.
Olóriz F. 1978: Kimmeridgiense-Tithonico inferior en el Sector cen-
tral de las Cordilleras Béticas (Zona Subbética). PhD thesis,
Universidad de Granada, 1–758.
Olóriz F.
& Tavera J.M. 1979: Nuevo Simoceratinae — Simoceras
(Cordubiceras) — en la base del Tithónico Superior
de las
Cordilleras Béticas (Zona Sub-Bética). Tecniterrae
29, 1–5.
Oppel A. 1863: Über jurassische Cephalopoden. Palaeont. Mitt. Mus.
KoeniglichBayerischen Staates 3, 127–266.
Oppel A. 1865: Die tithonische Etage. Zeit. Deut. Geol. Ges. 17,
535–558.
Parent H., Scherzinger A. & Schweigert G. 2011: The Tithonian-
Berriasian ammonite fauna and stratigraphy of Arroyo
Cieneguita, Neuquén-Mendoza Basin, Argentina. Bol. Inst.
Fisio. y Geol. 79–81, 21–94.
Pervinquière L. 1907: Étude de paléontologie tunisienne. I. Cépha-
lopodes des terrains secondaires. Direction générale des Travaux
publics. Carte géologique de la Tunisie. De Rudeval, Paris,
1–438.
Pruner P., Houša V. F., Oloriz F., Košak M. M., Krs M., Man O.,
Schnabl P., Venhodová D., Tavera J.-M. & Mazuch M. 2010:
High-resolution magnetostratigraphy and biostratigraphic
zonation of the Jurassic/Cretaceous boundary strata in the
Puerto Escaño section (southern Spain). Cretaceous Res. 31,
192–206.
Remane J. 1970: Die Entstehung der resedimentären Breccien im
Obertithon der subalpinen Ketten Frankreichs. Ecl. Geol. Hel.
63, 685–740.
Roman F. 1897: Recherches stratigraphiques et paléontologiques
dans le Bas-Languedoc. Ann. Univ. Lyon 34, 1–345.
Roman F. 1936: Le Tithonique du massif du Djurdjura (province
d’Alger). Mat. Carte Géol. Algérie, Paléontologie 7, 1–43.
Roman F. 1938: Les ammonites jurassiques et crétacées. Essai de
genera. Masson, Paris, 1–554.
Salazar C.A. 2012: The Jurassic–Cretaceous boundary (Tithonian–
Hauterivian) in the Andean Basin of Central Chile:. ammonites,
bio- and sequence stratigraphy and palaeobiogeography. PhD.
thesis, RupechtKarlsUniversität Heidelberg, 1–389.
Salazar C. & Stinnesbeck W. 2015: Tithonian–Berriasian ammonites
from the Baños del Flaco Formation, central Chile. J. Syst.
Paleontol. 0, 1–34.
Salfeld H. 1921: Kiel- und Furchenbildung auf der Schalenaussen-
seite der Ammonoideen in ihrer Bedeutung für die Systematik
und Festlegung von Biozonen. Zent. Min. Geol. Pal. (1921),
343–347.
Sapunov I.G. 1977: Ammonite stratigraphy of the Upper Jurassic in
Bulgaria. IV. Tithonian, substages, zones and subzones. Geol.
Balcanica 7, 43–64.
Sapunov I.G. 1979: Fossils from Bulgaria. III. 3. Upper Jurassic
series. Ammonoidea. Izdat. B´lgarskata Akad. naukite, Sofia,
1–263 (in Bulgarian with French summary).
Shome S. & Bardhan S. 2009: A new Late Tithonian ammonite
assemblage from Kutch, Western India. J. Pal. Soc. India 54,
1–18.
Spath L.F. 1923: On ammonites from New Zealand. In: Trechmann
C.T. (Ed.): The Jurassic rocks of New Zealand. Quarterly J.
Geol. Soc. London 79, 286–308.
Spath L.F. 1925: Ammonites and aptychi. The collection of fossils
and rocks from Somaliland. Mono. Geol. Dep. Hunt. Museum 1,
111–164.
Spath L.F. 1931: Revision of the Jurassic cephalopod fauna of Kachh
(Cutch). Part IV. Pal. Indica 9, 1–945.
Steuer A. 1897: Argentinische Jura-Ablagerungen: Ein Beitrag zur
Kenntniss der Geologie und Palaeontologie der argentinischen
Anden. Palaeontol. Abh., Jena new series 7, 127–222.
Steuer A. 1922: Jurassic strata from Argentina. Act. Acad. Nac. Cie.
Córdoba 7, 25–128 (in Spanish).
Szives O. & Fözy I. 2013: Systematic descriptions of Early Cretaceous
ammonites of the carbonate formations of the Gerecse
Mountains. In: Fözy I. (Ed.): Late Jurassic-Early Cretaceous
fauna, biostratigraphy, facies and deformation history of the
carbonate formations in the Gerecse and Pilis Mountains (Trans-
danubian Range, Hungary). GeoLitera Publishing House,
Szeged, 293–342.
Tavera J.- M. 1985: The ammonites from the Upper Tithonian-Berri-
asian of the Subbetic zone (Betic Cordillera). PhD. Thesis,
Universidad de Granada, 1–381 (in Spanish).
Toucas A. 1890: Étude sur la faune des couches tithoniques de
l´Ardèche. Bull. Soc. Géol. France 18, 560–629.
Uhlig V. 1910: The Fauna of the Spiti Shales. Memoirs of the
Geological Survey of India, Pal. Indica 15, 133–395.
Vigh G. 1984: Die biostratigraphische Auswertung einiger
Ammoniten- Faunen aus dem Tithon des Bakonygebirges sowie
aus dem Tithon-Berrias des Gerecsegebirges. Ann. Inst. Geol.
Hungarica 67, 1–210.
Westermann G.E.G. 2000: Biochore classification and nomenclature
in paleobiogeography: an attempt at order. Palaeogeogr.
Palaeoclimatol. Palaeoecol. 158, 1–13.
Wimbledon W.A.P., Reháková D., Pszczółkowski A., Cassellato C.E.,
Halásová E., Frau C., Bulot L.G., Grabowski J., Sobień K.,
Pruner P., Schnabl P. & Čížková K. 2013: A preliminary account
of the bio- and magnetostratigraphy of the upper Tithonian–
lower Berriasian interval at Le Chouet, Drôme (SE France).
Geol. Carpath. 64, 437–460.
Windhausen A. 1931: Geology of Argentina. Second Part, Historical
and regional geology of the territory of Argentina. 1–645 (in
Spanish).
Wippich M.G.E. 2001: Die tiefe Unter-Kreide (Berrias bis
Unter-Hauterive) im Südwestmarokkanischen Becken:
Ammonitenfauna, Bio- und Sequenzstratigraphie. PhD Thesis,
University of Ruhr, Bochum, 1–474.
Yin T.H. 1931: Etude de la faune du Tithonique coralligène du Gard
et de l’Hérault. Trav. Lab. Géol. Fac. Sci. Lyon 18, 1–193.
Zittel von K. 1868: Die Cephalopoden der Stramberger Schichten.
Pal. Mitt. Mus. KöniglichBayerischen Staates 2, 33–118.
i
FRAU, BULOT, WIMBLEDON and IFRIM
SUPPLEMENTARY DATA
Table 1: Dimensions of the type specimens and Spanish species referred to M. microcanthum. * indicates that the measurements of type
specimens were performed on plaster cast.
species
no. Specimen
D
U
Wh
Wb
U/D
Wh/D
Wb/D
Wb/Wh
D. mutari
F.G1.12.1
170
85
47.2
46.6
0.5
0.28
0.27
0.99
129.2
61.7
39.2
42.1
0.48
0.3
0.33
1.07
T.G1.12.67
118.2
54.7
36.2
39
0.46
0.31
0.33
1.08
87.6
39.7
28
29
0.45
0.32
0.33
1.04
T.G1.12.70
127.5
64
36.2
36.5
0.5
0.28
0.29
1.01
97.9
47.4
28.6
29.1
0.48
0.29
0.3
1.02
D. mediterraneum
T.GA1.1.1
148
77
39.8
37.2
0.52
0.27
0.25
0.93
126
69
32
29
0.55
0.25
0.23
0.91
F.G1.12.64
131.3
64.6
37
37
0.49
0.28
0.28
1
D. sinuosum
W.GA10.3.15
156
77
46.5
45
0.49
0.3
0.29
0.97
114.4
57.3
33.5
29
0.5
0.29
0.25
0.87
H. gen. et sp. Indet
T.GA6b.1.45
98.1
51.4
28.8
27
0.52
0.29
0.28
0.94
T.GA8.-3.5
112
56.9
31.3
31.6
0.51
0.28
0.28
1.01
M. (M.) microcanthum
AI-29
88.4
48
23.5
26
0.54
0.27
0.29
1.11
70.3
39.2
17.8
20.4
0.56
0.25
0.29
1.15
T.G23a.R.5.82
80.5
39.7
24
23
0.49
0.3
0.29
0.96
61.9
30.6
17.9
18
0.49
0.29
0.29
1.01
T.Gax
109
61.2
26.9
29.3
0.56
0.25
0.27
1.09
85.8
48
22.5
22.4
0.56
0.26
0.26
1
T.G23a.4.5
85.8
48
22.5
22.4
0.56
0.26
0.26
1
52.5
29.3
13.7
14.2
0.56
0.26
0.27
1.04
T.GA10.2.1.
44.7
23.8
11.3
12.3
0.53
0.25
0.28
1.09
82.3
47.7
21.1
25
0.58
0.26
0.3
1.18
T.G23a.4.120
64.3
38
14.1
18
0.59
0.22
0.28
1.28
29.9
14.8
8.9
10.2
0.49
0.3
0.34
1.15
25
12.4
7.5
9
0.5
0.3
0.36
1.2
T.G23a.R.5.113
32.8
17.3
9.5
12
0.53
0.29
0.37
1.26
26.6
13.4
7
10
0.5
0.26
0.38
1.43
T.GA6.4.28
30
16.2
9.3
10.6
0.54
0.31
0.35
1.14
T.GA6b.4.28
37.3
20.4
9.6
12.4
0.55
0.26
0.33
1.29
T.A2c.10.1
45.2
25
13.2
12
0.55
0.29
0.27
0.91
35.3
18.6
9.4
0.53
0.27
0
0
T.G23a.R.5.97
65.8
38.7
15
17
0.59
0.23
0.26
1.13
T.G23a.5.19
43.6
26
10.4
10.6
0.6
0.24
0.24
1.02
35.7
21.4
8.3
9.2
0.6
0.23
0.26
1.11
M. (M.) cf. brightoni
W.G19b.5.1
69.3
37
17
22
0.53
0.25
0.32
1.29
50.4
27.4
12.7
15.9
0.54
0.25
0.32
1.25
M. (C.) lotenoense
HOLOTYPE
27.7
10.6
10.4
11.5
0.38
0.38
0.42
1.11
F.G1.12.3
31.9
14.3
10.1
10.2
0.45
0.32
0.32
1.01
F.G1.12.64
26.9
12.3
8.7
9.2
0.46
0.32
0.34
1.06
22
9.6
7.9
8.2
0.44
0.36
0.37
1.04
F.G1.12.21
40.5
20
12.7
13.4
0.49
0.31
0.33
1.06
29.6
13.8
9
11
0.47
0.3
0.37
1.22
M. (C.) radians
W.GA10.3.32
48.7
22.9
15.1
15.2
0.47
0.31
0.31
1.01
37.7
17.4
12
11.6
0.46
0.32
0.31
0.97
T.Co.R.3
40.4
18.5
12.8
12.6
0.46
0.32
0.31
0.98
33.4
14.5
11
10
0.43
0.33
0.3
0.91
W.PR.8.1
49.1
22.5
15.3
16.2
0.46
0.31
0.33
1.06
38.7
18
12.9
12.7
0.47
0.33
0.33
0.98
W.PR.8.4
36.3
16.5
11.6
11.2
0.45
0.32
0.31
0.97
28.3
12.2
9.3
9.4
0.43
0.33
0.33
1.01
M. (C.) rhodanicum
NS-22
63.4
35
16
18
0.55
0.25
0.28
1.13
50.9
28
13
13.7
0.55
0.26
0.27
1.05
T.G23a.R.5.100
62.3
33.4
15.8
16.9
0.54
0.25
0.27
1.07
50.3
25.7
14.2
14.4
0.51
0.28
0.29
1.01
T.G1.11.49
49
25.6
13.2
13.9
0.52
0.27
0.28
1.05
ii
UPPER TITHONIAN AMMONITES FROM CHARENS (DRÔME, FRANCE)
Table 1 (continuation): Dimensions of the type specimens and Spanish species referred to M. microcanthum. * indicates that the measurements
of type specimens were performed on plaster cast.
species
no. Specimen
D
U
Wh
Wb
U/D
Wh/D
Wb/D
Wb/Wh
M. (C.) minor
T.GA1.1.4
23
11.4
7.1
8
0.5
0.31
0.35
1.13
20
9.8
6
7
0.49
0.3
0.35
1.17
T.GA8.-3.7
30
13.5
10.4
10.6
0.45
0.35
0.35
1.02
23.1
9.9
8.1
9.4
0.43
0.35
0.41
1.16
M. (C.) mendozanum
T.GA6b.1.52
35
17
10.9
9.5
0.49
0.31
0.27
0.87
31.6
15.2
9.9
8.1
0.48
0.31
0.26
0.82
M. (C.) flexuosum
T.GA1.1.2
60
32.3
15.2
16.2
0.54
0.25
0.27
1.07
50.5
27
13.3
12.6
0.53
0.26
0.25
0.95
T.GA1.1.5
75.7
37.5
21.5
19
0.5
0.28
0.25
0.88
64.4
33.9
16.3
16.6
0.53
0.25
0.26
1.02
T.GA6b.1.40
72.2
36.5
20.9
19.6
0.51
0.29
0.27
0.94
57.3
28.6
16.5
15.4
0.5
0.29
0.27
0.93
W.GA10.3.21
76.4
38.8
21.8
21.5
0.51
0.29
0.28
0.99
52.5
27
14.5
13.6
0.51
0.28
0.26
0.94
W.GA10.3.22
58.5
30.3
16.8
15.2
0.52
0.29
0.26
0.9
53
27.1
14.5
13.8
0.51
0.27
0.26
0.95
W.GA10.3.25
59.5
30
17.5
17
0.5
0.29
0.29
0.97
46.1
22.6
13.2
15
0.49
0.29
0.33
1.14
W.GA10.3.36
71.6
35.5
20.7
18.8
0.5
0.29
0.26
0.91
54.3
26.8
15.2
14.7
0.49
0.28
0.27
0.97
T.G1.12.66
67.9
35.5
20.1
17.8
0.52
0.3
0.26
0.89
52.8
27.3
14.1
12.8
0.52
0.27
0.24
0.91
M. (C.) koellikeri
T.GA7.26.30
63
29.8
20
18.4
0.47
0.32
0.29
0.92
47
22.3
14.8
13.2
0.47
0.31
0.28
0.89
M. (C.) ornatum
T.GA8.-3.4
77
38.3
22.1
24.4
0.5
0.29
0.32
1.1
W.GA10.3.18
75.6
37
23
25
0.49
0.3
0.33
1.09
52.2
25.6
15.2
17.2
0.49
0.29
0.33
1.13
M. (C.) symbolum
T.G1.11.47
36.2
19
9.7
13
0.52
0.27
0.36
1.34
T.G23a.R.5.77
58
34.1
14
18.6
0.59
0.24
0.32
1.33
47.6
27
11.1
16.6
0.57
0.23
0.35
1.5
T.Ce.R.6
81.3
49.2
17.8
22.4
0.61
0.22
0.28
1.26
67.7
38.6
17
21.8
0.57
0.25
0.32
1.28
65.8
40.4
13.9
19.6
0.61
0.21
0.3
1.41
W.GA10.3.17
85
46.8
21.2
25.4
0.55
0.25
0.3
1.2
M. (C.) leanzai
T.GA6b.1.46
70.7
36
19.7
0.51
0.28
0
0
T.GA6b.1.47
77.8
38.2
22.6
24.2
0.49
0.29
0.31
1.07
57.7
28.6
17.1
18.2
0.5
0.3
0.32
1.06
T.GA8.-3.2
108
55.5
30.2
33.4
0.51
0.28
0.31
1.11
89.3
44.7
24.7
0.5
0.28
0
0
T.GA8.-3.3
80
42
21.7
22.8
0.53
0.27
0.29
1.05
A. parvulus
T.G23a.R.5.90
33.6
17.6
9.5
9
0.52
0.28
0.27
0.95
27.2
13.8
8
8.8
0.51
0.29
0.32
1.1
A. sulcatus
T.GA1.1.8
78.5
35.5
25
21
0.45
0.32
0.27
0.84
T.GA6.1.8
78.1
37.4
24.3
20
0.48
0.31
0.26
0.82
T.GA10.3.3
81.1
38.5
24.7
21.4
0.47
0.3
0.26
0.87
T.G2.2.3
64.3
31.1
19.1
16
0.48
0.3
0.25
0.84
T.G2.2.3
73
35.5
22.4
20
0.49
0.31
0.27
0.89
T.G23a.R.13
64.5
30.5
19.2
18.5
0.47
0.3
0.29
0.96
51.2
23.5
15.8
15.7
0.46
0.31
0.31
0.99
A. venustus
W.PR8.5
43.8
18.5
14.6
15.4
0.42
0.33
0.35
1.05
36.3
15.8
12.2
13.4
0.44
0.34
0.37
1.1
85
42.5
25.3
23.5
0.5
0.3
0.28
0.93
A. microcanthus
Lectototype*
60.7
27.85
20.9
20.6
0.46
0.34
0.34
0.99
A. microcanthus
Paratype*
140.1
65.3
45.15
35.5
0.47
0.32
0.25
0.79
A. symbolus
Lectotype*
79.25
44.5
21.45
26.4
0.56
0.27
0.33
1.23
A. koellikeri
Lectotype*
160
83
43
42
0.52
0.27
0.26
0.98
iii
FRAU, BULOT, WIMBLEDON and IFRIM
Table 2: Dimensions of Micracanthoceras microcanthum (Oppel in
Zittel, 1868) from Charens. Yellow squares indicate approximated
dimensions.
no. Specimens
D
U
Wh
U/D
Wh/D
MPP-CHT.29/1
41
18.9
14.25
0.46
0.35
MPP-CHR.34/1
28.6
MPP-CHR.34/2
17.7
MPP-CHR.37/1
MPP-CHR.37/2
MPP-CHR.37/3
190
88.2
55
0.46
0.29
MPP-CHR.45/1
26.3
MPP-CHR.50/1
MPP-CHR.50/2
12.9
MPP-CHR.50/3
28.42
11.94
9.8
0.42
0.34
MPP-CHR.50/3x
MPP-CHR.51/2
27
11.5
9.7
0.43
0.36
MPP-CHR.53/2
MPP-CHR.53/4a-c
10.17
7.96
MPP-CHR.54/1a
51
17.5
0.34
MPP-CHR.54/1b
14.2
MPP-CHR.60/1
18.8
MPP-CHR.60/2
25
16
MPP-CHR.60/3
15.3
MPP-CHR.60/4
50.6
15.5
0.31
MPP-CHR.60/6
39.3
18.3
12
0.47
0.31
MPP-CHR.60/7
MPP-CHR.62/1
11.2
MPP-CHR.63/1
9
MPP-CHR.67/4
33.4
15.4
12.26
0.46
0.37
MPP-CHR.67/7
MPP-CHR.67/9
38
17.2
11.5
0.45
0.3
MPP-CHR.68/1
16.25
MPP-CHR.68/2
20.95
9.22
7.26
0.44
0.35
MPP-CHR.82/1
56.5
26.8
16.2
MPP-CHR.87/1
41.3
20.7
10.7
MPP-CHR.92/2
26.5
MPP-CHR.92/4
51.4
24.8
14.5
0.48
0.28
MPP-CHR.92/5
28.7
MPP-CHR.92/10
27.5
10.9
8.4
0.4
0.31
MPP-CHR.92/11
51.4
25.2
15.75
0.49
0.31
MPP-CHR.92/13
32.4
15.56
10.8
0.48
0.33
MPP-CHR.92/28
18.07
c14.5
MPP-CHR.98/1
58
30.7
18.3
0.53
0.32
MPP-CHR.R100/6
57.4
26.7
18.3
0.47
0.32
Table 3: Dimensions of Protacanthodiscus andreaei (Kilian, 1889)
from Charens. Yellow squares indicate approximated dimensions.
no. Specimens
D
U
Wh
U/D
Wh/D
MPP-CHR.79/4
52.46
19.03
19.2
0.4
0.4
MPP-CHR.19/5
12.8
4.6
4.5
0.4
0.4
MPP-CHR.21/3
41
18.2
13.6
0.4
0.3
MPP-CHR.21/6
27.48
11.7
11.11
0.4
0.4
MPP-CHR.21/19
43
20.5
15
0.5
0.3
MPP-CHR.21/20
17.4
MPP-CHR.21/25
28.7
13.35
9.2
0.5
0.3
MPP-CHR.21/29
24.26
8.4
10
0.3
0.4
MPP-CHR.21/32
13.84
5.14
4.57
0.4
0.3
MPP-CHR.21/34
6
MPP-CHR.21/42
25.5
12.9
6.4
0.5
0.3
Table 4: Dimensions of Boughdiriella chouetense Frau, Bulot &
Wimbledon, 2015 from Charens. Yellow squares indicate approxi-
mated dimensions.
no. Specimens
D
U
Wh
U/D
Wh/D
MPP-CHR.19/1
23.3
7.2
11.7
0.3
0.5
MPP-CHR.21/4
35.2
14.5
11.5
0.4
0.3
MPP-CHR.21/8
31.64
12.8
11.03
0.4
0.3
MPP-CHR.21/11
26.07
9.7
9.9
0.4
0.4
MPP-CHR.21/24
27.96
11.36
9.2
0.4
0.3
MPP-CHR.21/27
29
14.9
9.5
0.5
0.3
MPP-CHR.21/33
31.7
15.57
13.61
0.5
0.4
MPP-CHR.21/35
12.5
12.1
8
1
0.6
MPP-CHR.21/40
8.7
Table 5: Dimensions of Pratumidiscus elsae Bulot, Frau & Wimble-
don, 2014 from Charens.
no. Specimens
D
U
Wh
U/D
Wh/D
MPP-CHR.21/43
9.8
Table 6: Dimensions of Busnardoiceras busnardoi (Le Hégarat,
1973) from Charens. Yellow squares indicate approximated
dimensions.
no. Specimens
D
U
Wh
U/D
Wh/D
MPP-CHR.19/1x
36.98
15.6
14.6
0.4
0.4
MPP-CHR.19/3
32.24
10.63
12.57
0.3
0.4
MPP-CHR.21/7
66.3
23.94
24.25
0.4
0.4
MPP-CHR.21/10
48
16.8
15.2
0.4
0.3
MPP-CHR.21/12
53.35
20.5
18.3
0.4
0.3
MPP-CHR.21/13
27.77
9.84
10.05
0.4
0.4
MPP-CHR.21/14
32.2
11.85
11.8
0.4
0.4
MPP-CHR.21/15
39.43
14.35
14.3
0.4
0.4
MPP-CHR.21/16
23.65
9.42
8.83
0.4
0.4
MPP-CHR.21/17
22.59
7.8
7.95
0.3
0.4
MPP-CHR.21/18
MPP-CHR.21/26
60.37
20.97
0.3
MPP-CHR.21/28
25.61
11.1
9.6
0.4
0.4
MPP-CHR.21/36
94.7
37.87
34
0.4
0.4
MPP-CHR.21/37
28.4
11.7
11
0.4
0.4
MPP-CHR.21/38
53.6
20.7
19
0.4
0.4
MPP-CHR.21/42
40.5
15
14.8
0.4
0.4
iv
UPPER TITHONIAN AMMONITES FROM CHARENS (DRÔME, FRANCE)
Fig. S1. U=f(D) for Spanish specimens referred to M. microcanthum and comparison with type specimens.
Fig. S2. Wh=f(D) for Spanish specimens referred to M. microcanthum and comparison with type specimens.
v
FRAU, BULOT, WIMBLEDON and IFRIM
Fig. S3. Wb=f(D) for Spanish specimens referred to M. microcanthum and comparison with type specimens.
Fig. S4. Wb=f(Wh) for Spanish specimens referred to M. microcanthum and comparison with type specimens.