Geobios 45 (2012) 99–108

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Original article

New records of early Middle Jurassic belemnites in the French Subalpine Basin
and their paleobiogeographic significance§
Nino Mariotti a, Robert Weis b,*, Andrea Di Cencio c, Arnaud Clément d, Kenneth De Baets e
a
Dipartimento di Scienze della Terra, Università ‘‘La Sapienza’’, Piazzale A. Moro 5, 00185 Roma, Italy
b
Musée national d’histoire naturelle, section Paléontologie, 25, rue Münster, 2160 Luxembourg, Luxembourg
c
Via Pescara 242, 66100 Chieti Scalo, Italy
d
5b, rue de Camargue, La Cigalière, 05000 Gap, France
e
Paläontologisches Institut und Museum der Universität Zürich, Karl Schmid-Strasse 4, 8006 Zürich, Switzerland

A R T I C L E I N F O A B S T R A C T

Article history: A biostratigraphic and systematic study based on belemnites collected along with ammonites was
Received 3 December 2010 performed on four sections in the Subalpine Basin (SE France): Lac du Castillon and La Baume (Castellane
Accepted 16 November 2011 area), Galabrun and Grand Lara (Gap area). The specimens, originating from hemi-pelagic marl-
Available online 26 December 2011
limestone alternations in the lower part of the ‘‘Calcaires à Zoophycos’’ Formation, are dated from Middle
Aalenian (Murchisonae Zone) to Lower Bajocian (Humphriesianum Zone). Five belemnite taxa
Keywords: (Megateuthis elliptica, Holcobelus munieri, H. trauthi, Pachybelemnopsis roettingensis, Hibolithes sp.) have
Holcobelidae
been identified, and two more taxa are reported in an open nomenclature (Belemnitida incertae sedis sp.
Megateuthididae
Mesohibolithidae
1 and sp. 2). The biostratigraphic range of the belemnite fauna is established. The new findings
Middle Jurassic contribute to a more detailed understanding of the paleobiogeography of holcobelid belemnites that
Subalpine Basin flourished at the northern margin of the Tethys Ocean and formed a distinct sub-Mediterranean fauna.
Biostratigraphy The association herein described is similar to the fauna of the Calabro-Peloritani Arc (Calabria, Italy), a
Paleobiogeography further hint for the supposed paleogeographic position of the latter during the Middle Jurassic.
ß 2012 Elsevier Masson SAS. All rights reserved.

1. Introduction between the Boreal and Tethyan realms. Particularly, a sub-Boreal

fauna dominated by Belemnitina (Homaloteuthis, Megateuthis,
The French Subalpine Basin and its famous Mesozoic outcrops Brevibelus, Eocylindroteuthis) has been recognized in Luxembourg,
have been the subject of many paleontological studies during the Germany, NE France and northern Switzerland while a sub-
last two centuries. The Digne-Barrême-Castellane area is well Mediterranean fauna dominated by Pachybelemnopseina (Holco-
known for its rather complete Middle Jurassic hemipelagic sections belus, Pachybelemnopsis, Hibolithes, Belemnitida incertae sedis) was
rich in cephalopod faunas (Pavia, 1973, 1983; Fernández-López, evidenced for southern Italy (Calabria) and southern France
2007; De Baets et al., 2008), and also holds the Bajocian-Bathonian (Mariotti et al., 2010). With the present work, we propose a
GSSP (Fernández-López et al., 2009a, 2009b). The Upper Aalenian- biostratigraphic scheme based on new belemnite records, strictly
Lower Bajocian ammonoid faunas are well-studied (Caloo, 1970, related to local ammonite stratigraphy, and discuss their paleobio-
1971; Pavia, 1983; De Baets et al., 2008 and references within), but to geographic relevance at the Aalenian/Bajocian boundary.
our knowledge, no systematic or biostratigraphic studies have been
published on early Middle Jurassic belemnites. Cretaceous belem- 2. Geological setting
nites on the other hand are well known from this area (Combémorel,
1973; Clément, 1999, 2000; Janssen and Clément, 2002; Janssen, The study area is located in the southern part of the French
2009; Clément and Janssen, 2012). Middle Aalenian to Lower Subalpine Basin (also called ‘‘Dauphinois Basin’’ in the Jurassic).
Bajocian belemnites have been poorly studied in most European The French Subalpine Basin is characterized by a stepwise
basins. Recently, Mariotti et al. (2007, 2010) and Weis and Mariotti deepening in the Jurassic (Olivero, 2003). During the early Middle
(2007) described Middle Jurassic belemnites and suggested that a Jurassic, the study area (‘‘Digne-Gap Basin’’) corresponded to a
peculiar distribution developed at the Aalenian/Bajocian boundary subsiding basin delimited by shallow-water highs (Fig. 1). In the
southern part, a transitional area between the Provence platform
§
Corresponding editor: Pascal Neige. and the basin shows mixed and condensed sequences, as in the
* Corresponding author. vicinity of Castellane (Assenat, 1972; Olivero, 1994). A major
E-mail address: [email protected] (R. Weis). stratigraphic gap from the middle or upper Toarcian to the middle

0016-6995/$ – see front matter ß 2012 Elsevier Masson SAS. All rights reserved.
doi:10.1016/j.geobios.2011.11.012
100 N. Mariotti et al. / Geobios 45 (2012) 99–108

2.1.1. Lac du Castillon (Castellane, Alpes de Haute Provence)

The outcrop, part of a protected area, is situated along the road
on the south-eastern border of the Castillon Lake, between the
barrage of Castillon and the first tunnel direction Saint-Julien-du-
Verdon (438 530 1500 N; 68 320 1700 E). The stratigraphic sequence
(Fig. 2(1)), which is located within the Cremon unit, has first been
described by Assenat (1972) and shows:

 0.4 m Upper Pliensbachian, greyish, pyritic limestone (‘‘Calcaires

à Silex’’ Formation) (Fig. 2[1(1)]);
 0.01–0.02 m highly condensed, phosphatic and pyritic level
(Fig. 2[1(2)]) with many fossil shell fragments, gastropods,
bivalves and ammonites, ascribed to Paradumortieria distans
(Buckman, 1890) (Fig. 3(5, 6)), Catulloceras dumortieri (Thiollière
in Dumortier, 1874) (Fig. 3(8)) and Pleydellia sp. The ammonites
indicate the Upper Toarcian (base of the Aalensis Zone). This
indicates a hiatus for at least the Lower and Middle Toarcian;
 0.1–0.2 m hardground with irregular stromatolitic structures
(Fig. 2[1(3)]). At the top, the irregular surface of the hardground
is filled with a marly sediment containing numerous belemnites:
Holcobelus munieri (Eudes-Deslongchamps, 1878), Belemnitida
incertae sedis sp. 1 and sp. 2. Usually, this belemnite fauna spans
from the Middle Aalenian (Murchisonae Zone) to the Lower
Bajocian (Propinquans Zone). This belemnite-rich bed is capped
by cyclic alternations typical of the ‘‘Calcaires à Zoophycos’’
Formation, the base of which is lower Bajocian in the Cremon
unit (Kerckhove, 1976);
 about 2 m limestone-marl alternations (‘‘Calcaires à Zoophycos’’)
(Fig. 2[1(4)]), with frequent trace fossils (Zoophycos). An
ammonite from the upper part of the section, Kumatostephanus
perjucundus (Buckman, 1927) (Fig. 3(1)), indicates the Lower
Bajocian (Laeviuscula or Propinquans Zone). A complete belem-
nite rostrum found at the same level belongs to Belemnitida
incertae sedis sp. 1.

2.1.2. La Baume (Castellane, Alpes de Haute Provence)

This section (Fig. 2(2)) is part of a protected area and has been
studied by Assenat (1972), Leonide (2007) and De Baets et al.
Fig. 1. Sedimentary/structural setting of the Mesozoic ‘‘Gap-Digne Basin’’ and (2008). It is situated about 200 m above the small hamlet of La
location of the studied sections. Modified after Baudrimont and Dubois (1977). Baume, on the south-western side of the Castillon Lake (438 530 3600
N; 68 300 0500 E). De Baets et al. (2008) subdivided the section into
or upper Aalenian can be recognized in the southern sections (La 122 beds. A small Toarcian-aged condensed/lacunous unit, studied
Baume, Lac du Castillon), which largely corresponds with the by Assenat (1972) and Leonide (2007), at the base of the outcrop is
maximum regression of the hemicycle R6 of Graciansky et al. capped by the ‘‘Calcaires à Zoophycos’’ Formation that can be
(1998). The northern sections (Galabrun, Grand Lara) have a more subdivided roughly into three members (De Baets et al., 2008):
continuous sedimentation at this time. The difference in sedimen-
tary evolution between different sections lies in interaction of fault  a lower marly member 1 (beds 1–24), with ammonites ranging
activity and position of various basin highs at time of sedimenta- from the Upper Aalenian Concavum Zone to the Lower Bajocian
tion. A brief description of the studied sections is necessary as only Discites Zone. Several belemnite specimens, ascribed to Holco-
one has been published in detail before (La Baume; see De Baets belus munieri, were collected in the Concavum Zone;
et al., 2008).  an intermediate, more calcareous member 2 (beds 24–109)
corresponds roughly to the Lower Bajocian Propinquans and
2.1. Castellane Arc Humphriesianum Zones, including also ammonites from the
Laeviuscula Zone. Belemnites from this member include
The studied sections in the Castellane area (La Baume and Lac Holcobelus trauthi (Stolley, 1927) (beds 42 and 50), Belemnitida
du Castillon) are situated at the southern margin of the basin, close incertae sedis sp. 1 (bed 60), Pachybelemnopsis roettingensis
to the Moyen Verdon High (Fig. 1); thicknesses are variable in this (Schlegelmilch, 1998) (bed 68) and Hibolithes sp. (bed 77);
transitional area between the platform and the basin. In the  an upper, more marly member 3 (beds 109–122) corresponds
northern part of the Castellane Arc, the Upper Aalenian is largely to the Upper Bajocian and basal Bathonian. Belemnites
represented by an alternation of marls and limestones (La Baume were found in some beds (De Baets et al., 2008: fig. 3b), but were
section; De Baets et al., 2008). Southward, these Aalenian deposits not determinable.
are reduced to a hardground (Lac du Castillon section; Assenat,
1972). At both sections, the Lower Bajocian is well developed by 2.2. Gap Basin
further marl-limestone alternations with trace fossils (e.g.,
Zoophycos) and frequent cephalopods (‘‘Calcaires à Zoophycos’’ The two sections studied near Gap (Galabrun and Grand Lara;
Formation; Kerckhove, 1976). Fig. 1) display considerable thicknesses indicating a subsident
 N. Mariotti et al. / Geobios 45 (2012) 99–108 101

Fig. 2. Lithological succession of the investigated sections with belemnite occurrences and proposed ammonoid biostratigraphy: 1. Lac du Castillon (Castellane). 2. La Baume
(Castellane); simplified after De Baets et al. (2008). 3. Grand Lara (Gap). 4. Galabrun (Gap). 5. View of the section Galabrun near Gap (picture taken in 2010), the sampled beds
(1–6) are indicated in white.
102 N. Mariotti et al. / Geobios 45 (2012) 99–108

Fig. 3. 1. Kumatostephanus perjucundus (Buckman, 1927), Propinquans Zone, Lac du Castillon/Castellane (RNGHP 010027). 2. Skirroceras macrum (Quenstedt, 1886/87),
Propinquans Zone, Hebridica Subzone, Galabrun/Gap (MnhnL QH163). 3. Brasilia sp., Murchisonae Zone, Grand Lara/Gap (MnhnL QH164). 4. Otoites contractus (Sowerby,
1825), Propinquans Zone, Patella Subzone, Galabrun/Gap (MnhnL QH165). 5, 6. Paradumortieria distans (Buckman, 1890), Aalensis Zone (Upper Toarcian), Lac du Castillon/
Castellane (RNGHP 010025). 7. Sonninia propinquans (Bayle, 1848), Propinquans Zone, Patella Subzone, Galabrun/Gap (MnhnL QH166). 8. Catulloceras dumortieri (Thiollière in
Dumortier, 1874), Upper Toarcian, Lac du Castillon/Castellane (RNGHP 010026). Scale bars = 4 cm (1, 2), 2 cm (4–7), 1.25 cm (3), 1 cm (8).

basin, with maxima of 450 m for the Aalenian, and 250 m for the black marls (‘‘Marnes gris sombre’’; Kerckhove et al., 1989) are
Lower Bajocian (Propinquans and Humphriesianum Zones) exposed on a hill, culminating at 940 m, to the South-West of the
according to Mouterde and Elmi (1984). The Middle and Upper hamlet Grand Lara (448 330 5900 N; 68 080 2100 E). The top of this unit
Aalenian is represented by 200 to 250 m of black marls with rare is marked by a specimen of Brasilia sp. (determined by A. Di Cencio)
calcareous beds. The marly facies continues into the Bajocian (Fig. 3(3)).
(Discites Zone), then it progressively changes to a limestone-marl
alternation, with the trace fossil Zoophycos (Propinquans and 3. Results
Humphriesianum Zones; Mouterde and Elmi, 1984).
3.1. Systematic paleontology
2.2.1. Galabrun (Gap, Hautes Alpes)
The section (Fig. 2(4, 5)) is situated along the road D6, about The belemnite classification used herein follows Riegraf et al.
100 m south from the hamlet Galabrun, in the direction of Gap (448 (1998). The belemnites have been determined by N. Mariotti and
330 2500 N; 68 070 1100 E). It displays 4.5 m of marl-limestone R. Weis, the newly collected ammonites by A. Di Cencio. Most
alternations (‘‘Calcaires gréso-argileux noirs’’; Gidon, 1971) with ammonite species reported in this study have been described
frequent Zoophycos. A bed-by-bed sampling allowed us to gather elsewhere (Pavia, 1983), including ammonoids from the La Baume
specimens of Holcophylloceras sp., Sonninia sp., Sonninia propin- section (De Baets et al., 2008). Therefore, a simple figuration of new
quans (Bayle, 1848) (Fig. 3(7)), Otoites contractus (Sowerby, 1825) specimens is deemed sufficient. The examined specimens are
(Fig. 3(4)), Skirroceras macrum (Quenstedt, 1886–1887) (Fig. 3[2]), housed in the following depositories:
Holcobelus munieri, Megatheutis elliptica (Miller, 1826), Belemnitida
incertae sedis sp. 1 and sp. 2, aptychii, and disarticulated crinoid  RNGHP: Réserve Naturelle Géologique de Haute Provence,
fragments. The paleontological record allows to assign these beds Digne-les-Bains (F), including specimens collected by F. Cecca,
to the upper part of the Lower Bajocian (Propinquans Zone), which K. De Baets, M. Guiomar and L. Leroy;
can be subdivided in two subzones (Patella and Hebridica).  MnhnL: Musée national d’histoire naturelle, Luxembourg (L),
Previous studies also indicate a Propinquans Zone (‘‘Sauzei Zone’’) including specimens collected by A. Clément.
for the lower part of this unit in this region (Gidon, 1971).
Individual descriptions also include size measurements in the
2.2.2. Grand Lara (Gap, Hautes Alpes) cases of complete or almost complete specimens (see Appendix A).
This classic section (Fig. 2(3)) has already been investigated by Dimensional adjectives are used as in Doyle and Kelly (1988); the
Haug (1891), who reported ‘‘Belemnites munieri’’. The Aalenian terms small, medium and large, related to the length of the rostrum
 N. Mariotti et al. / Geobios 45 (2012) 99–108 103

Fig. 4. 1, 2. Megateuthis elliptica (Miller, 1826), Propinquans Zone, Galabrun/Gap (MnhnL BEL038). 3, 4. Holcobelus munieri (Eudes-Deslongchamps, 1878), Murchisonae Zone,
Grand Lara/Gap (MnhnL BEL039). 5–7. Holcobelus munieri (Eudes-Deslongchamps, 1878), Propinquans Zone, Galabrun/Gap (MnhnL BEL044). 8. Pachybelemnopsis
roettingensis (Schlegelmilch, 1998), Humphriesianum Zone, La Baume/Castellane (RNGHP 010016). 9. Hibolithes sp., Humphriesianum Zone, La Baume/Castellane (RNGHP
010015). 10, 11. Holcobelus trauthi (Stolley, 1927), Laeviuscula Zone, La Baume/Castellane (RNGHP 010012). Scale bar: 2 cm.

(L), respectively refer to L < 80 mm, L between 80 and 110 mm, Type species: Belemnites giganteus v. Schlotheim, 1820 (= junior
and L > 110 mm. synonym of Belemnites suevicus Klein, 1773), by subsequent
designation (Douvillé, 1879).
Order BELEMNITIDA MacGillivray, 1840
Suborder BELEMNITINA MacGillivray, 1840 Megateuthis elliptica (Miller, 1826)
Family MEGATEUTHIDIDAE Sachs and Nalnjaeva, 1967 Fig. 4(1,2).
Genus Megateuthis Bayle, 1878 * 1826. Belemnites ellipticus Miller, p. 60, pl. 8, figs. 14–17.
104 N. Mariotti et al. / Geobios 45 (2012) 99–108

v. 2006. Megateuthis elliptica (Miller) – Weis, p. 157, figs. 9, 10 guished Holcobelus deshayesi (Mayer) from H. munieri. In our view,
(cum syn.). these variations conserve always the typical morphological
v. 2007. Megateuthis elliptica (Miller) – Weis and Mariotti, p. characters (e.g. length of the ventral groove, cylindriconical outline
166, pl. 7, fig. 1. and profile, shape and length of the epirostrum, lateral compres-
Material and occurrence: Eight complete juvenile-adult rostra sion), so that H. deshayesi is included in the range of variation of
from the section Galabrun (Gap, Hautes Alpes), Propinquans Zone, H. munieri and becomes a junior synonym.
Patella Subzone (coll. MnhnL BEL038).
Holcobelus trauthi Stolley, 1927
Description: Large-sized, elongate, cylindriconical rostrum.
Fig. 4(10, 11)
Both profile and outline are almost symmetrical and cylindrico-
*v. 1927 Holcobelus trauthi Stolley, p. 118, pl. XXIV, figs. 7, 8.
nical. The apical region is elongated and characterised by an
v. 2010 Holcobelus trauthi Stolley – Mariotti et al., fig. 4k, l.
epirostrum, visible in longitudinal sections. Two dorsolateral
Material and occurrence: One adult and two juvenile rostra
grooves are present, although weakly developed. The transverse
from the section La Baume (Castellane area, Alpes de Haute
sections are elliptical and compressed.
Provence), beds 42 and 50, Lower Bajocian, Laeviuscula Zone (coll.
Remarks: The studied specimens are typical of M. elliptica.
RGHP).
Their relatively small size (180 mm for the longest specimen) is
Description: Small-sized, robust and cylindriconical rostrum.
typical for those stratigraphically older individuals from the
The outline is symmetrical and slightly subhastate. The profile is
Laeviuscula and Propinquans Zones.
symmetrical and cylindriconical. The apical region is elongated,
tapering regularly towards a moderately acute apex. A well incised
Suborder PACHYBELEMNOPSEINA Riegraf in Riegraf et al., 1998
ventral groove extends from the apical region towards the alveolar
Family HOLCOBELIDAE Gustomesov, 1977
border. Transverse sections of the rostrum are highly compressed.
Genus Holcobelus Stolley, 1927
Remarks: The studied specimens match well the character-
Type species: Belemnites munieri Eudes-Deslongchamps, 1878.
istics of H. trauthi as described by Stolley (1927): elongated apical
Holcobelus munieri (Eudes-Deslongchamps, 1878) region and high lateral compression. The only difference with
Fig. 4(3–7), Fig. 5(5, 6) Stolley’s specimens from Normandy is the robustness of our
* 1878. Belemnites munieri Eudes-Deslongchamps, p. 63, pl. V, material, compared to the elongate, gracile types, which represent
figs. 3–6, 12–14; pl. VI, figs. 5–11.0 probably subadult individuals.
v. 1994. Holcobelus munieri (Eudes-Deslongchamps) – Combé-
Family MESOHIBOLITHIDAE Nerodenko, 1983
morel et al., p. 15, pl. 2, figs. 7, 8.
Genus Hibolithes Denys de Montfort, 1808
v. 2010. Holcobelus munieri (Eudes-Deslongchamps) – Mariotti
Type species: Hibolithes hastatus Denys de Montfort, 1808.
et al., fig. 4i, j.
Material and occurrence: Seventeen complete juvenile-adult Hibolithes sp.
rostra and more than one hundred fragments from the section Fig. 4(9)
Grand Lara (Gap, Hautes Alpes), Middle Aalenian (coll. MnhnL 2007. Hibolithes sp. – Mariotti et al., p. 11, pl. 2, fig. 5.
BEL039-041). Two complete subadult-adult rostra from the section Material and occurrence: One fragment (62 mm) from the La
Galabrun (Gap, Hautes Alpes), Propinquans Zone, Patella Subzone Baume section (Castellane area, Alpes de Haute Provence), bed 77,
(coll. MnhnL BEL032, 036). One complete adult rostra and Lower Bajocian, basal Humphriesianum Zone (coll. RGHP).
numerous fragments from the section La Baume (Castellane, Alpes Description: This single fragment corresponds to the middle
de Haute Provence), Upper Aalenian, Concavum Zone (coll. RGHP). part of a large-sized individual with the following characteristics:
Five subadult-adult rostra and several fragments from the section outline hastate, broad ventral alveolar canal, transverse sections
Lac du Castillon (Castellane area, Alpes de Haute Provence), depressed.
Aalenian/Bajocian boundary (coll. MnhnL BEL010). Remarks: Although only a part of the rostrum is conserved, the
Description: Medium to large-sized, elongate, cylindriconical specimen is similar to large-sized Hibolithes as figured in Mariotti
rostrum. The outline is symmetrical and cylindriconical, the profile et al. (2007: pl. 2, fig. 5).
is cylindriconical or slightly asymmetrical in case of a deformed
Genus Pachybelemnopsis Riegraf, 1980 (pro Belemnopsis auct.,
epirostrum. The apex is elongated and moderately acute. The
not Bayle, 1878)
epirostrum is well developed in most adult specimens. A long
Type species: Belemnites canaliculatus v. Schlotheim, 1820.
intermediate ventral groove extends from close the tip of the apex
until the alveolar end, without reaching the alveolar border. The Pachybelemnopsis roettingensis (Schlegelmilch, 1998)
transverse sections of the rostrum are elliptical and compressed. Fig. 4(8)
The phragmocone penetrates approximately one quarter of the v. 1980. Belemnopsis (Longibelemnopsis) cf. beyrichi (Oppel) –
rostrum in adults; this proportion may vary in case of a particularly Riegraf, p. 188, text – fig. 172, pl. 2, figs. 20, 21.
long epirostrum. * 1998. Belemnopsis roettingensis Schlegelmilch, p. 78, pl. 17,
Remarks: Some individuals from the Concavum Zone of Gap Figs. 2–4.
reach a remarkable size with up to 150 mm in rostrum length, v. 2007. Pachybelemnopsis roettingensis Schlegelmilch – Weis
compared to the maximal length of 100 mm reached by rostra from and Mariotti, p. 170, pl. 7, fig. 9.
the type area in Normandy. Moreover, the material herein studied Material and occurrence: One complete rostrum (67 mm)
shows a relatively wide morphological variation; this can be from the La Baume section (Castellane area, Alpes de Haute
explained by morphological differences between material of Provence), bed 68, Lower Bajocian, basal Humphriesianum Zone
different ontogenetic stages (e.g., rostra of H. munieri had (coll. RGHP).
previously been described as a separate species, H. subblainvillei Description: Small-sized, slender and gracile rostrum. The
Eudes-Deslongchamps, 1878, now considered a subjective syno- specimen is partly embedded in the rock and only the outline can
nym of the former), different preservations and/or different be observed. The outline is symmetrical and subhastate. The apical
stratigraphic ages. Especially the individuals from the Lower region is elongated and acute. The well marked ventral alveolar
Bajocian evolved towards a more gracile rostrum. Based on canal extends from the alveolar border towards the apical region; it
material from Bulgaria, Stoyanova-Vergilova (1993) has distin- fades out at 20 mm from the tip of the apex. The transverse section
 N. Mariotti et al. / Geobios 45 (2012) 99–108 105

Fig. 5. 1, 2. Belemnitida incertae sedis sp. 2, Propinquans Zone, Hebridica Subzone, Galabrun/Gap (MnhnL BEL043). 3, 4. Belemnitida incertae sedis sp. 2, Propinquans Zone,
Galabrun/Gap (MnhnL BEL034). 5, 6. Holcobelus munieri (Eudes-Deslongchamps, 1878), Murchisonae Zone, Grand Lara/Gap (MnhnL BEL040). 7, 8. Belemnitida incertae sedis
sp. 1, Propinquans Zone, Patella Subzone, Galabrun/Gap (MnhnL BEL037). Scale bar: 2 cm.

of the rostrum is depressed in the apical region, but compressed at present. The transverse sections of the rostrum are compressed on
the alveolar end. the whole length.
Remarks: The present rostrum is only partially preserved; Remarks: The present form, along with Belemnitida incertae
nonetheless it is a typical specimen of P. roettingensis, as figured by sedis sp. 2, shows morphological characters which are found
Weis and Mariotti (2007: pl. 7, fig. 8). separately in belemnites belonging to different genera and
families: a long ventral groove, post-alveolar compression of the
Belemnitida incertae sedis sp. 1
rostrum, extremely short termination of the apical part, great
Fig. 5(7, 8)
width of the groove, subhastate shape. The specimens from SE
pars 1994. Belemnitida incertae sedis – Combémorel et al., p.
France are similar to the belemnites described as Belemnitida
48, pl. 1, figs. 1, 6–11.
incertae sedis from peri-Tethyan sediments in Calabria, Italy
v. 2007. Belemnitida incertae sedis – Mariotti et al., p. 12, figs.
(Combémorel et al., 1994; Mariotti et al., 2007). A more
1–3 [non figs. 6, 10].
comprehensive paleontological study on material from several
Material and occurrence: One subadult rostrum from the
European localities by the authors (R. Weis, N. Mariotti) is in
section Lac du Castillon (Castellane, Alpes de Haute Provence),
progress, which will define the systematic position of the present
Lower Bajocian (coll. RGHP, former coll. Leroy). One subadult
form, including the description of new taxa.
rostrum from the section La Baume (Castellane, Alpes de Haute
Provence), Lower Bajocian, Propinquans Zone (coll. RGHP). Four Belemnitida incertae sedis sp. 2
subadult rostra from the section Galabrun (Gap, Hautes Alpes), Fig. 5(1–4)
Lower Bajocian, Propinquans Zone (coll. MnhnL BEL035, 037). pars 1994. Belemnitida incertae sedis – Combémorel et al., p.
Description: Small or medium-sized, elongate rostrum. The 48.
outline is symmetrical and cylindrical or slightly subhastate. The v. 2007. Belemnitida incertae sedis – Mariotti et al., p. 12, fig. 10
profile is symmetrical and cylindrical. The short apical region [non figs. 1–3, 6].
closes with a rounded and mucronate apex. A long ventral groove Material and occurrence: Three juvenile-adult rostra from the
extends towards the apical region (fading out 5 mm before the tip section Galabrun (Gap, Hautes Alpes), Lower Bajocian, Propin-
of the apex) and towards the alveolar border, which it reaches quans Zone, Patella and Hebridica Subzones (coll. MnhnL BEL034,
under form of a shallow depression. Lateral lines are distinctly 043). One adult rostrum from the section Lac du Castillon
106 N. Mariotti et al. / Geobios 45 (2012) 99–108

(Castellane, Alpes de Haute Provence), Lower Bajocian (coll. MnhnL

Megateuthis elliptica

Belemnitida inc. sedis

Substage

Holcobelus munieri
Holcobelus trauthi
Mesohibolithidae
Megateuthididae
BEL013).

? Hibolithes sp.
P. roettingensis
STAGE
Period

Holcobelidae
Description: Medium-sized, reasonably robust and cylindrical Ammonite
rostrum. The outline is symmetrical and cylindrical. The profile is
asymmetrical, due to the dorsally orientated apex, and cylindrical. Zones

sp.1
sp.2
The apical region is short, mucronate and ‘‘beak’’-like, due to the
position of the apex, which is orientated towards the dorsal side. A
broad, ventral groove extends from the apical region (starting Humphrie-
5 mm form the tip of the apex) towards the alveolar border. Lateral sianum

BAJOCIAN
lines are weakly distinguishable on our specimens. The transverse

Middle JURASSIC

Lower
sections of the rostrum are compressed along the whole rostrum. Propinquans
The phragmocone penetrates approximately one half of the
rostrum. Laeviuscula
Remarks: See also ‘‘Remarks’’ under Belemnitida incertae sedis
sp. 1. The present form (sp. 2), is close to the previously described
(sp. 1), but differs slightly by possessing a more robust and Discites
cylindrical rostrum, a broader ventral groove, and an asymmetri-

L. Middle U.
cal, ‘‘beak’’-like apex. Concavum

AALENIAN
3.2. Biostratigraphy
Murchisonae
The biostratigraphic control of the belemnite finds is provided
by associated ammonoids (Fig. 3). We herein use the ammonite Opalinum
zonal schemes for the Aalenian by Contini et al. (1997) and
Bajocian by Rioult et al. (1997). Attempts to use Middle Jurassic
Fig. 6. Simplified stratigraphical distribution of the studied belemnite species in the
belemnites for a stratigraphic zonation have been achieved in the
Gap-Digne Basin, plotted against ammonite zones.
past (Stoyanova-Vergilova, 1990; Doyle and Bennett, 1995;
Combémorel, 1997). A reliable biostratigraphic scheme based on
belemnites obviously needs detailed systematic and stratigraphic 4. Paleobiogeography and discussion
studies of numerous sections located in different sedimentary
environments. The French Subalpine Basin is of interest for such a In Middle Jurassic times, the French Subalpine Basin repre-
study, due to its paleogeographic position at the boundary of the sented the north-western margin of the Alpine Tethys and a
European and Peri-mediterranean Tethyan domains. The present transitional area between the epicontinental sea of the Paris Basin
study is limited to four stratigraphic sections located in the basinal and the ocean of the Piedmont domain (Olivero, 2003; Olivero and
and the transitional facies. Our sections often contain Zoophycos, Mattioli, 2008). Due to its transitional geographic position, the
which in the Aalenian and Bajocian seems to be typical for the studied area is of special importance to study belemnite
outer shelf and slope environments in the French Subalpine Basin distribution patterns.
(Olivero, 2003). This is the first attempt to use Middle Jurassic According to Thierry (2000) and Callomon (2003), during the
belemnites for a biostratigraphic scheme in the Gap-Digne Basin. Middle Jurassic, the north-western European area was character-
The stratigraphic analysis allows the following remarks: ized by Boreal and sub-Boreal ammonite bioprovinces, whilst the
Tethyan area of southern Europe was characterized by a
 the Middle and Upper Aalenian is marked by an abundant, Mediterranean ammonite bioprovince; additionally, the southern
monospecific fauna characterized by Holcobelus munieri. This epi-Variscan basins (including the French Subalpine Basin)
species, which continues in the Lower Bajocian with less represent a sub-Mediterranean bioprovince with elements of
frequency, can be considered a valid marker for field paleontol- the sub-Boreal bioprovince. A similar pattern, with few differences,
ogy; can be inferred from belemnites, according to recent data by
 The range of Holcobelus trauthi marks the Lower Bajocian Mariotti et al. (2010) and the present study (Fig. 7). The fauna of the
(Laeviuscula Zone); Gap-Digne Basin and the similar fauna of the Calabro-Peloritani
 the Propinquans Zone is characterized by the presence of Arc (Combémorel et al., 1994; Mariotti et al., 2007) show
Belemnitida incertae sedis sp. 1 and sp. 2. Only in the Galabrun significant differences from the fauna of parts of the north-
section (Fig. 2(4)), Belemnitida incertae sedis sp. 1 was collected western European area (Luxembourg, NE France, SW Germany, N
in the Patella Subzone and Belemnitida incertae sedis sp. 2 in the Switzerland). This is also expressed in the Jaccard coefficient of
Hebridica Subzone; similarity (=C/N  100, where C = shared species and N = total
 the first occurrence of mesohibolithids (Pachybelemnopsis number of species; compare Doyle, 1987: 238) between the sub-
roettingensis and Hibolithes sp.) marks the base of the Hum- Boreal and the sub-Mediterranean fauna (Mariotti et al., 2010),
phriesianum Zone; which is equivalent to 20 for the Discites-Propinquans Zones
 Megateuthis elliptica has a stratigraphic distribution that covers interval (Lower Bajocian). The association of holcobelids
the Bajocian (Laeviuscula to Parkinsoni Zone); for this reason it (H. munieri, H. trauthi), early mesohibolithids (Pachybelemnopsis,
was chosen by Combémorel (1997) as a zonal index. This species Hibolithes) and Belemnitida incertae sedis sp. 1 and sp. 2, combined
is present with many individuals at the Galabrun section (Gap) in with the near absence of Belemnitina, is characteristic of this sub-
sediments of the Propinquans Zone, but was so far not found in Mediterranean belemnite fauna. Sub-Boreal taxa Megateuthis
the Digne-Castellane area. elliptica and Brevibelus sp. (Combémorel et al., 1994) are rare
elements that testify faunal exchanges with the north-western
All these data allow us to hypothesize a biostratigraphic scheme European area. In this context, it is noteworthy that Megateuthis
(Fig. 6) for the Middle Aalenian to Lower Bajocian of the Gap-Digne elliptica is abundant in the northern part of the Gap-Digne Basin
Basin. (Galabrun section) and almost absent in the southern part
 N. Mariotti et al. / Geobios 45 (2012) 99–108 107

English language. KDB would also like to thank Fabrizio Cecca

(Paris) and Jacques Verniers (Gent). Martin Košt’ák (Prague) and
Guillaume Dera (Dijon) reviewed the manuscript and greatly
contributed with helpful suggestions. The present research was
supported by the project ‘‘Jurassic Coleoids’’ at the National
Museum for Natural History, Luxembourg.

Appendix A. Supplementary data

Supplementary data associated with this article can be found, in

the online version, at doi:10.1016/j.geobios.2011.11.012.

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