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POTENTIAL FORAMINIFERAL MARKERS FOR THE VISEAN–
SERPUKHOVIAN AND SERPUKHOVIAN–BASHKIRIAN BOUNDARIES—
A CASE-STUDY FROM CENTRAL MOROCCO
PEDRO CO
´ZAR,
1
ISMAIL SAID,
1
IAN D. SOMERVILLE,
2
DANIEL VACHARD,
3
PAULA MEDINA-VAREA,
1
SERGIO RODRI
´GUEZ,
1
AND MOSTAFA BERKHLI
4
1
UEI y Depto. de Paleontologı
´a, Instituto de Geologı
´a Econo´mica/Instituto de Geociencias CSIC-UCM, c/Jose´ Antonio Novais 2,
28040-Madrid, Spain, ,pcozar@geo.ucm.es.,,ismsaid@gmx.fr.,,pmvarea@geo.ucm.es.,,sergrodr@geo.ucm.es.;
2
UCD School of Geological
Sciences, University College Dublin, Belfield, Dublin 4, Ireland, ,ian.somerville@ucd.ie.;
3
Universite´ de Lille 1, UFR Sciences de la Terre,
UMR 8157 du CNRS, Baˆ timent SN 5, Villeneuve d’Ascq 59655, France, ,daniel.vachard@univ-lille1.fr.; and
4
Universite´ Moulay Ismaı
¨l, Faculte´ des
Sciences, De´partement de Ge´ ologie, UFR analyse et prospection des bassins se´dimentaires,
BP. 11201 Zitanne, Mekne`s, Morocco, ,mberkhli@fsmek.ac.ma.
ABSTRACT—The Carboniferous succession in Adarouch (Central Morocco, north of the Atlas Transform Fault)
contains thick carbonate beds including upper Visean, Serpukhovian and basal Bashkirian rocks. Foraminifers
enable precise recognition of the Visean/Serpukhovian (V/S), early/late Serpukhovian (eS/lS) and Serpukhovian/
Bashkirian (S/B) boundaries. Insolentitheca horrida,Loeblichia ukrainica,‘‘Millerella’’ spp. and Endostaffella? sp. 2
are regarded as regionally useful indices to the V/S boundary, whereas Eostaffellina spp., Eostaffella pseudostruvei
and some evolved species of Archaediscus exhibit greater reliability for worldwide correlation of this level. Similarly,
the eS/lS boundary is marked locally by Brenckleina rugosa,Eosigmoilina sp., and Monotaxinoides spp.
and globally by Loeblichia minima,Bradyina cribrostomata,Plectostaffella spp., Eostaffellina ‘‘protvae’’ and
‘‘Turrispiroides’’, and the S/B boundary is marked locally by Globivalulina bulloides and globally by Seminovella
elegantula, and Novella?. Occurrences of these taxa in Morocco allow correlations with the Moscow Basin, the
Urals, the Donetz Basin and North America. The Moroccan assemblages share few taxa in common with Saharan
basins south of the Atlas Transform Fault. Correlations with western European basins are difficult because of the
paucity in the latter of foraminiferal-bearing carbonate strata.
INTRODUCTION
RESEARCH ON the Visean–Serpukhovian (V/S) boundary
deals with several potential stratotypes, involves many
different fossil groups, and includes work in both deep-water
facies and shallow-water facies. The base of the Serpukhovian
was equated to the base of the Namurian, which is defined by
the first appearance of Cravenoceras leion, a datum well
established in the British Isles (Bisat, 1930).
Ammonoids, in particular C. leion, are not free of
controversy and their identification depends on well-preserved
specimens. Furthermore, their usefulness for intercontinental
correlations has been questioned (Nikolaeva and Kullmann,
2001). A third inconvenience is the strong facies control on the
ammonoid faunas but this is a similar problem observed in
other Carboniferous fossil groups. Nevertheless, ammonoid
zonal schemes similar to the British one have been used
successfully in Morocco and Algeria (e.g., Pareyn, 1961;
Lemosquet et al., 1985; Korn et al., 2007; Korn and
Ebbighausen, 2008; Wendt et al., 2009), and the British
ammonoid succession has been correlated with Germany (e.g.,
Korn, 1996, 2006; Korn and Horn, 1997), the Urals
(Nikolaeva et al., 2009a) and North America (Ramsbottom
and Saunders, 1985; Richards and Task Group, 2009).
The Visean/Serpukhovian boundary is currently under inten-
sive study worldwide in connection with efforts to select a GSSP
(see Richards and Task Group, 2009, 2010). During 2009–2010,
the task group chair intended to submit a proposal that the base
of the Serpukhovian will be defined by the first appearance
datum (FAD) of the conodont Lochriea ziegleri within the
lineage L. nodosa–L. ziegleri,butthescarceknowledgeofthe
geographic distribution of the lineage and the degree of
diachroneity of the FAD of L. ziegleri warrant a vote to be
accepted or rejected. However, in areas where conodonts are rare
or absent, such as in shallow-water platform interiors (e.g.,
Central Morocco), the V/S boundary can be recognized by
foraminifers which provide a highly useful auxillary tool.
The Serpukhovian/Bashkirian boundary is well established
as it also coincides with the mid-Carboniferous Mississippian/
Pennsylvanian boundary (e.g., Heckel and Clayton, 2006).
The GSSP is located at Arrow Canyon (Nevada, U.S.A.) and
is defined by the first appearance of the conodont Declinog-
nathodus noduliferus (Brenckle et al., 1997a, 1997b; Lane et al.,
1999). Diagnostic Bashkirian foraminifers occur some distance
above the boundary in the stratotype section in the Urals
(Groves, 1988; Groves et al., 1999; Kulagina et al., 2009).
The Carboniferous succession in Adarouch (central Mo-
rocco) ranges in age from the middle Visean up to the
Bashkirian, with excellent preservation and exposure of the
latest Visean and Serpukhovian carbonates. The succession
occurs in several faulted blocks or thrust sheets (e.g., Berkhli,
1999; Vachard et al., 2006; Fig. 1). It is characterized by cyclic
sequences, hundreds of meters thick, comprising shallow-
water platform carbonates in the lower part and deep-water
open platform to slope carbonates and shales in the upper part
(Berkhli, 1999; Berkhli and Vachard, 2001; Co´zar et al.,
2008a), foraminiferal assemblages in the carbonates are
abundant and of high diversity.
The aims for this study are to 1) document the foraminiferal
succession from the latest Visean to Bashkirian in a relatively
unknown area in central Morocco (western Paleotethys); 2)
analyze those problems related to the Visean/Serpukhovian
boundary, as well as the Serpukhovian/Bashkirian boundary;
and 3) to propose foraminiferal taxa which might be
potentially used as markers for these boundaries, which in
the case of the Visean/Serpukhovian boundary, in the absence
of the Lochriea lineage, may provide important local
alternatives in biostratigraphy.
Journal of Paleontology, 85(6), 2011, p. 1105–1127
Copyright ’2011, The Paleontological Society
0022-3360/11/0085-1105$03.00
1105
1106 JOURNAL OF PALEONTOLOGY, V. 85, NO. 6, 2011
GEOLOGIC AND BIOSTRATIGRAPHIC SETTINGS
The Carboniferous formations described in the Adarouch
region (Fig. 1D) are included in the northern part of the
Azrou-Khenifra Basin which was interpreted as a foreland
basin (Beauchamp and Izart, 1987; Fig. 1B). The oldest
sediments in the area are situated in Agouraı
¨(Berkhli et al.,
2000; Fig. 1C), dated as V2a, mid Visean. In Adarouch,
sedimentation spans only the late Asbian–Bashkirian interval.
Upper Asbian–Brigantian rocks are represented by the Tizra
and Mouarhaz formations. The best known Serpukhovian
formation is the carbonate-dominated Akerchi Formation
(Fig. 1D). This formation was originally assigned to the late
Visean by Termier and Termier (1951), Termier et al. (1975),
and Conil and Verset (1979). However, Dedok and Hollard
(1980) attributed the upper part of the formation to the early
Namurian on the basis of brachiopods. Vachard and Tahiri
(1991), Vachard and Fadli (1991) and Vachard and Berkhli
(1992) considered the Akerchi Formation as entirely lower
Serpukhovian.
Berkhli (1999), Berkhli et al. (2001) and Berkhli and Vachard
(2001) subdivided the Akerchi Formation (,100 m thick) into
two members. They considered the lower AK1 member
(predominantly limestone and shales) as uppermost Visean
(Brigantian), and the upper AK2 member (predominantly sandy
limestones and sandstones) as mostly lower Serpukhovian.
During the late Brigantian two main contrasting succession-
types are recognized. The first one is a deepening-upward
sequence into open platform to slope sediments in the upper
part (e.g., Tizra and Mouarhaz formations; Figs. 1, 2). The
second one is a shallowing-upward trend, passing into
shallow-water, nearshore siliciclastic-rich shelf facies (Akerchi
Formation). The carbonate-rich lower member of the Mouar-
haz Formation (MZ1) and the lower part of the Akerchi
Formation (AK1) contain shallow-water facies foraminifers of
late Brigantian age (Figs. 3–5; Co´zar et al., 2008a).
Said (2005) described a typical Brigantian rugose coral
assemblage from the biostrome at the base of the AK2
member (Figs. 4, 5) and assigned a Brigantian age for the
lower part of the AK2 member. Subsequently, Co´zar et al.
(2008a) documented Brigantian foraminifers from the bio-
strome but relatively poor assemblages in higher levels up to
the stromatolitic band (,35 m above the base of the section,
see Fig. 5). However, further sampling of the sandstones and
pebbly limestone beds between the stromatolite and the
biostrome revealed the presence of Serpukhovian foraminifers.
Thus, the base of the Serpukhovian is situated in the bed from
which sample 3230 in Akerchi 1 section was collected and
from sample 3232 in Akerchi 2 (see Figs. 4 and 5). The base of
the late Serpukhovian is located immediately above the
stromatolite in sample 2772 (see Fig. 5). A Bashkirian age in
the upper part of the Akerchi Formation was proposed by
Dedok and Hollard (1980) based on the presence of the
brachiopod Titanaria. An unpublished re-examination of the
material (Legrand-Blain, personal commun., 2000), consid-
ered that the specimen is not the typical species found in
Algerian Sahara. This, together with its earliest occurrence in
North America (Carter, 1990), led Berkhli (1999), Berkhli and
Vachard (2001), and Berkhli et al. (2001) to assign the top
of the Akerchi Formation to a probable E2 Zone (late
Serpukhovian age). The foraminiferal assemblages from the
upper beds, although sparse, do contain significant taxa,
supporting a probable Bashkirian age (see below). Unfortu-
nately, conodonts have not been found in these youngest
siliciclastic-rich beds at Akerchi.
The Idmarrach Formation to the northeast of Akerchi
(Fig. 1D) was described by Berkhli and Vachard (2002) and
dated as Serpukhovian. Those authors only studied the
Idmarrach 1 section. According to the coral assemblages, Said
(2005) considered the formation as Brigantian. However, recent
conodont sampling has established a late Serpukhovian age for
lower levels in Idmarrach 1 section, with the discovery of
Gnathodus bilineatus bollandensis (Fig.2).Theoccurrenceofthe
conodont Declinognathodus noduliferus in the youngest limestone
beds (,40 m above the base of the Idmarrach 2 section), confirms
a Bashkirian age for the upper 10 m of the section (Fig. 2), which
shows pebbly reddish siliciclastic beds at the top (Fig. 7).
Finally, the Tirhela Formation is exposed to the north of
Idmarrach (Figs. 1D, 8). This unit was first described by
Berkhli and Vachard (2002) and dated by them as late Visean
to Serpukhovian. The Tirhela Formation (,90 m thick)
comprises marine limestones with abundant gigantoproduc-
toid brachiopods in the lower part and is attributed mostly to
the late Serpukhovian. However, foraminifers from the
uppermost limestone beds are assigned to the Bashkirian
and are comparable with those limestones from near the top of
Idmarrach 2 section (Fig. 2). The upper part of the Tirhela
Formation marks a thick transition into reddish marginal
marine and deltaic conglomerates, sandstones and shales, which
are compared to the upper red pebbly beds in Idmarrach 2.
FORAMINIFERAL MARKERS OF THE UPPERMOST VISEAN TO
BASHKIRIAN IN ADAROUCH
A summary of the stratigraphic range of the most significant
foraminiferal taxa illustrated in Figures 10 to 13 is given in
Figure 9. Details of their ranges displayed in Figures 3–8 is
based on examination of 1,050 thin-sections, prepared from
the samples highlighted in the stratigraphic logs. Thin-sections
are housed in the Palaeontological collection of the Uni-
versidad Complutense de Madrid (Faculty of Geological
Sciences).
Uppermost Visean–lower Serpukhovian.—The shallow-water
oolitic limestones (MZ 1 member) of the Mouarhaz Forma-
tion contain Globivalvulina parva,Asteroarchaediscus ex gr.
baschkiricus,Janischewskina,Endothyranopsis sphaerica and
common Euxinita efremovi (Fig. 3). All are considered
late Brigantian markers elsewhere in Europe (Co´zar and
Somerville, 2004; Co´zar et al., 2005, 2008a, 2008b, 2010).
Noteworthy, is Pseudocornuspira (Fig. 10.1–10.3) (or Recto-
cornuspira of other authors), which was commonly recorded in
the late Brigantian from the Donetz Basin by Vdovenko
(2001) and Kazakhstan (Brenckle and Milkina, 2003).
Another interesting new taxon is Globivalvulina sp. 1
(Fig. 10.6, 10.7), which can be readily included within the
r
FIGURE 1—A, location of the main Carboniferous Basins in Morocco, Algeria and Tunisia: 1, Central Massif; 2, Rehamna; 3, Djebilet; 4, Tafilalt; 5,
Be´ char; 6, Jerada; 7, Tindouf; 8, Reggane; 9, Ahnet; 10, Mouydir; 11, subsurface Carboniferous; AA5Anti-Atlas, ATF5Atlas Transform Fault,
EM5Eastern Meseta, HA5High Atlas, MA5Middle Atlas, WM5Western Meseta; B, detail of the Variscides in Morocco, pale gray areas are
Carboniferous outcrops, dark grey is the Azrou-Khenifra Basin; C, location of the Adarouch area (encased box) in the northern part of the Azrou-
Khenifra Basin, see also arrow in B;D, photogeological map showing the location of the measured stratigraphic sections in the Carboniferous
formations (modified from Weyant, 1985; Hoepffner et al., 2005; Simancas et al., 2005).
CO
´ZAR ET AL.—LATE VISEAN TO BASAL BASHKIRIAN FORAMINIFERS FROM MOROCCO 1107
Globivalvulina granulosa group but seems to be endemic to
central Morocco.
The first appearance of Neoarchaediscus postrugosus
(Fig. 11.9) is recorded in the lower part of the Akerchi
Formation of latest Brigantian age. The first appearance
datum (FAD) of this species elsewhere is in the latest
Brigantian, in England, Scotland (Co´zar and Somerville,
2004; Co´zar et al., 2008b), Ireland (Somerville and Co´ zar,
2005; Co´zar et al., 2005), Spain (Co´ zar et al. 2006), and the
Urals (Kossovaya et al., 2001; Reitlinger et al. in Einor, 1996).
Noteworthy is the first occurrence of Euxinita pendleiensis
(Fig. 11.12–11.14) in the upper part of the calcareous
succession (AK1). This species was described from Pendleian
rocks from the Midland Valley of Scotland (Co´zar et al.,
2008b). Also highlighted is the first occurrence of the Miliolata
calcivertellids (including Calcitornella,Calcivertella,Trepei-
lopsis and Ammovertella; Fig. 10.4, 10.5).
One of the most significant occurrences in the Akerchi
Formation is the genus ‘‘Millerella’’ and some species of
Endostaffella, which is included by some authors within the
synonymous genus, Zellerinella. The type species of Zellerinella
selected by Mamet and Skipp (1970) was Endothyra discoidea
Girty, 1915, and they also included most of the primitive Millerella
described in the Chesterian of the midcontinent, U.S.A. However,
Girty (1915) did not select a holotype and neither did Mamet and
Skipp, 1970 (as already noted by Brenckle, 2005), and the two
axial sections illustrated within the types are morphologically
distinct. They could be identified here as morphotype 1 (Girty,
1915, pl. 10, fig. 11), with more embracing and elongated whorls
than in morphotype 2 (Girty, 1915, pl. 10, fig. 12), although this
second morphotype was not included in the species by Mamet and
Skipp (1970). Reitlinger (1975, 1981) highlighted the similarity of
the species with Endostaffella shamordini (Rauser-Chernoussova,
1948) and both species have been commonly considered as
synonyms (e.g., Brenckle, 2005; van Ginkel, 2010). The genera
Zellerinella and Endostaffella have been also considered as
synonymous by Brenckle and Groves (1981), Baxter and
Brenkle (1982), and Rich (1986). However, Loeblich and
Tappan (1988), Rauser-Chernoussova et al. (1996) and Pinard
and Mamet (1998) kept both genera as independent taxa. As
claimed by Pinard and Mamet (1998), the lectotype selected by
Loeblich and Tappan (1988) is an equatorial section which is
not representative enough. The single axial section considered as
the lectotype by the former authors is more representative, but
as Brenckle (2005) remarked, it was not written in the original
diagnosis of the genus. The latter author published numerous
topotypes and paralectotypes, which readily correspond to the
morphotype 2, which might be considered also as Endostaffella?.
The problem may arise with specimens similar to morphotype 1,
as well as with those others species included by Mamet and
Skipp (1970) in the genus, such as Endothyra tortula Zeller, 1953
and E. designata Zeller, 1953. Those specimens can be found
referred to in the literature as Millerella,‘‘primitiveMillerella’’,
‘‘Millerella’’ and Paramillerella. Although none of the previous
genus names satisfies completely the taxonomic characters of
the specimens, for pragmatic purposes we prefer to retain the
use of ‘‘Millerella’’ here.
This genus Zellerinella and thus, the species included in
‘‘Millerella’’, is considered as mostly endemic to the North
American Realm, although rare specimens have been recorded
in eastern Europe and the Far East (see Armstrong and Mamet,
1977). More recently, ‘‘Millerella’’ tortula was recorded in the
Tarusskian and Steshevskian in the Moscow Basin (Gibshman,
2001, 2003; Gibshman and Baranova, 2007), ‘‘Millerella’’
designata and ‘‘Millerella’’ cooperi (5Eostaffella advena) from
the Serpukhovian and Bashkirian of Kazakhstan (Brenckle and
Milkina, 2003), Zellerinella spp. from Ireland and Scotland
(Somerville and Co´zar, 2005, pl. 3, figs. 19–21; Co´zar et al.,
2010, fig. 7g), and Zellerinella sp. 1, Zellerinella sp. 2, Z.
designata and Z. pressula from China (Wu et al., 2009). Most
specimens recorded in Adarouch are determined as Endostaf-
fella?discoidea (Fig. 12.14, 12.15).
Specimens identified as ‘‘Millerella’’ aff. tortula are rarely
recorded (Fig. 12.19, 12.20). Although they do not present the
same number of whorls and size as in the original material, the
evolute arrangement of the whorls and depressed umbilical
areas are rather similar. ‘‘Millerella’’ aff. tortula is exclusively
recorded in Serpukhovian rocks in Adarouch, together with
some uncommon specimens identified as ‘‘Millerella’’ sp.
(Fig. 12.16). The presence of these assemblages of ‘‘Millerella’’
highlight a local influence of the North American fauna on
Central Morocco for this period, and its paleobiogeographic
implications should be analyzed in detail.
An unusual taxon is recorded from the basal Serpukhovian
beds in Adarouch, referred to here as Endostaffella? sp. 2
(Fig. 11.21–11.25). This taxon shows characters of that genus
(skew-coiled inner whorls and final planispiral whorls), but
differs in two aspects: 1) a differentiated wall with marked
bright clear grains, which in some cases developed a well-
defined diaphanotheca and 2) secondary basal deposits in the
form of pseudochomata, better developed than in the genus
Endostaffella. In fact, this species might be derived from an
evolved form of Euxinita, which developed more planispiral
final whorls. Some conspecific specimens might occur in the
Midland Valley of Scotland (Co´ zar et al., 2008b, 2010). A
much more common species of the genus and restricted also to
the Serpukhovian in Adarouch is identified as Endostaffella?
sp. 1 (Fig. 12.12, 12.13), which is distinctly wider and with
slightly more involute whorls than the former species, and it is
considered close to E.?discoidea, but the latter first occurs in
the latest Brigantian, as do other species of the genus. These
features, together with the plectogyroid inner whorls, give
generally, a weaker development of the depressed umbilical
areas.
Eostaffella pseudostruvei (Fig. 12.2, 12.3) first occurs from
the Serpukhovian in Adarouch, although it first occurs earlier,
in the Brigantian, in some European basins, such as the
Midland Valley of Scotland (Co´ zar et al., 2008b, 2010), and the
Urals (Kossovaya et al., 2001; Reitlinger et al. in Einor, 1996).
Two taxa which first occur in the Serpukhovian of Adarouch,
Insolentitheca horrida (Fig. 10.15) and Loeblichia ukrainica
(Fig. 11.17–11.20), both from the Idmarrach Formation could
be potentially useful as regional markers. They have been
documented from the Serpukhovian (Aisenverg et al., 1979a;
Lys, 1985; Charlot-Prat and Vachard, 1989), but also are
recorded in the Brigantian (late Visean). Loeblichia ukrainica is
well known in the late Visean in the Donetz Basin (Vdovenko,
2001), northern Bulgaria (PC, unpublished data) and in the
Tindouf Basin (Sebbar et al., 2000). Insolentitheca is also rare,
but it was documented in upper Visean rocks by Mamet (1970),
Ivanova (1973a, 1973b), Conil (1980), Brazhnikova and
Vdovenko in Aizenverg et al. (1983), Laloux (1988) and
Vdovenko (2001), although they might belong to its ancestral
form, Protoinsolentitheca Vachard and Co´ zar, 2004. The species
L. ukrainica is characterized by the skew-coiled inner whorls.
However, this character is variable within specimens and some
are almost planispiral (Fig. 11.18, 11.19), in which case it might
be confused with other species of the genus (e.g., L. ammonoides
or L. paraammonoides). On the other hand, the rounded
periphery of the lumen is characteristic, also for those slightly
1108 JOURNAL OF PALEONTOLOGY, V. 85, NO. 6, 2011
biumbilicate specimens (Fig. 11.17), which might be confused
with L. minima (with a marked acute periphery of the whorls,
Fig. 11.15–11.16). Most of the Moroccan specimens belong to
L. ukrainica.
The Archaediscus at tenuis stage (Fig. 11.2, 11.4–11.6) sensu
Conil et al. (1980) are also considered important for the
definition of the Serpukhovian, although not without some
problems. This group is characterized by the evolute
arrangement of the whorls, loose coiling and the virtual
absence of the microgranular layer. The subfamily Aster-
oarchaediscinae sensu stricto, and the evolution between the
Neoarchaediscus and Asteroarchaediscus species might be
particularly useful, because the late Brigantian–Serpukhovian
interval is the acme of those forms, with many new species.
Eostaffellina spp. (Fig. 13.1–13.4) is also recorded in the
early Serpukhovian in Adarouch. This compressed species is
also recorded in the early Serpukhovian of Kazakhstan
(Brenckle and Milkina, 2003, as Eostaffellina? sp.). In the
Moscow Basin, Aisenverg et al. (1979a) documented Eostaf-
fellina with a rounded periphery from the Tarusskian, but no
species determinations were documented, that might be the
case for those Eostaffellina spp. These specimens can be
characterized by low length/diameter ratios, around the
minimum proposed by Reitlinger (1963) in the diagnosis of
the genus (0.60), and close to the parameters described for E.
schartimiensis (Malakhova, 1956). In Tian Shan and Pamirs
areas, unsually, the rounded and subspherical species of the
genus are listed from a lower horizon of the Serpukhovian (e.g.,
E. ‘‘protvae’’, E. schartimiensis), whereas compressed forms,
attributed to E. ‘‘paraprotvae’’ are only recorded in the upper
part of the late Serpukhovian (Bensh et al. in Einor, 1996).
The last taxon that first occurs in the early Serpukhovian of
Adarouch is Endothyranopsis plana (Fig. 10.13, 10.14). This
species is also potentially a global marker, and it has been
documented in Europe and Africa, although in some cases, in
the late Serpukhovian (Poletaev et al., 1991; Krainer and
Vachard, 2002; Co´ zar et al., 2008b). In Adarouch, its FAD is
situated in the upper part of the early Serpukhovian in the
Idmarrach 3 section.
In summary, the early Serpukhovian of Adarouch allows us
to propose 1) four regional foraminiferal markers for the base
of the Serpukhovian (Fig. 9): Endostaffella? sp. 1, Eostaffella
pseudostruvei,Insolentitheca horrida, and Loeblichia ukrainica,
and 2) four more taxa or groups of taxa which potentially,
might be tested as global markers: Endostaffella? sp. 2,
Archaediscus at tenuis stage, Eostaffellina spp., and Endothyr-
anopsis plana. In addition, there are other groups, with their
acmes or abundance that also can be used for recognizing the
base of the Serpukhovian: Asteroarchaediscinae s.s., Globi-
valvulina and calcivertellids.
Upper Serpukhovian.—Monotaxinoides species (Fig. 10.16,
10.18–10.21) first occur consistently in the late Serpukhovian
in Central Morocco, and hence, the genus can be used a local
marker. Their first occurrence is established by some authors
as latest Visean (without illustrations: Conil et al., 1980, 1991;
Kossovaya et al., 2001; Poty et al., 2006), but Monotaxinoides
is illustrated in the Brigantian from the Donetz Basin
(Vdovenko, 2001, pl. 4, fig. 41) and late Brigantian in
Northern Ireland (Co´zar et al., 2005, fig. 8.16, 8.17), and
commonly documented from the base of the Serpukhovian in
FIGURE 2—Stratigraphic correlation of the sections in the Adarouch
area with first occurrence datum (FAD) of key conodont taxa: D.
noduliferus5Declinognathodus noduliferus and G. bollandensis5Gnathodus
bilineatus bollandensis.PE5Pendleian, MZ5Mouarhaz, AK5Akerchi.
r
CO
´ZAR ET AL.—LATE VISEAN TO BASAL BASHKIRIAN FORAMINIFERS FROM MOROCCO 1109
the Urals (Nikolaeva et al., 2009a, as Monotaxinoides? sp. or
Eolasiodiscus muradynus), Tunisia (Lys, 1988, as M. priscus),
England (Stephenson et al., 2010), Be´char Basin (Lys, 1975,
1985; Sebbar and Lys, 1989; as M. priscus) and Kazakhstan
(Brenckle and Milkina, 2003). Nevertheless, a primitive species
of the genus might be present from late Visean rocks
(Vachard, 1977, as M.? capriensis). In general, it is observed
that the planispiral forms, such as M. priscus, are usually
documented in older rocks than those more helicospiral forms.
It might be interpreted that forms such as M. subconica or M.
subplana are the ancestral species of the genus, and in that
case, they might be derived from Howchinia gibba. Other
related forms are assigned to ‘‘Turrispiroides’’ (Fig. 10.17,
10.22), because of the lack of a fibrous layer, but the validity
of the genus and its use is controversial, since the name was
originally used to describe a Miliolata (see discussion in
Vachard and Beckary, 1991). However, its FAD was
consistently assigned to the late Serpukhovian in Europe
and North America (e.g., Brenckle et al., 1982, 1997a; Conil
et al., 1991; Krainer and Vachard, 2002; Brenckle and
Milkina, 2003).
In Adarouch Eosigmoilina (Fig. 11.10, 11.11) and Brenck-
leina (Fig. 11.7, 11.8) are recorded in the late Serpukhovian
of Idmarrach 1 and Tirhela sections, with both genera
occurring in the same sample (3338) at the base of the late
Serpukhovian in Akerchi 1 section (Fig. 4). In general, there
is agreement that both genera show the same stratigraphic
range. However, in the Urals (see Kulagina et al., 2001, 2009;
Nikolaeva et al., 2009a) and in Tunisia (Lys, 1988) neither
taxa has been documented. In some cases only one genus is
documented (e.g., Conil et al., 1991; Gibshman, 2001). In
general, the FAD of both genera is at the base of the late
Serpukhovian (Mamet and Skipp, 1970; Mamet, 1974;
Laloux, 1988; Conil et al., 1991; Perret, 1993; Mamet et al.,
1994; Sebbar, 1998; Sebbar et al., 2000; Gibshman, 2001;
Poty et al., 2006; Pille, 2008).
Rare specimens of Eosigmoilina were illustrated from the
Tarusskian–Steshevskian horizons by Rozovskaya in Aisen-
verg et al. (1979a), but later the same specimens were revised
by Vdovenko (2001) to the base of the Prokhorovskian
Horizon (C
5
Limestone) (Fig. 14), laterally equivalent to the
Protvinskian Horizon. Eosigmoilina is considered a late
Serpukhovian marker in the Donetz Basin (Brazhnikova and
Vdovenko in Aizenverg et al., 1983; Poletaev et al., 1991;
Vdovenko, 2001), although some specimens of Brenckleina
were documented in C
4
Limestone by Brazhnikova and
Vdovenko in Aizenverg et al. (1983). The earliest occurrence
of Brenckleina might be related to the distinction made
between zones 18 and 19 of Mamet and Skipp (1970), in which
some of the characteristic markers are Rugosoarchaediscus and
Eosigmoilina, respectively. Nevertheless, as Brenckle et al.
(1977) demonstrated, they both occur at equivalent strati-
graphic levels in the North American Realm. In England and
Scotland, Eosigmoilina was regarded as Arnsbergian (late
Serpukhovian) by Fewtrell et al. (1981) and Riley (1992).
In Algeria, according to Lys (1975, 1985), Eosigmoilina and
Brenckleina are first recorded in the E1 Zone in the Tindouf
Basin and Be´ char Basin whereas, other authors working in
the same basins, suggest they both first occur in the E2 Zone
(Mamet et al., 1994; Sebbar, 1997; Sebbar et al., 2000). In the
central Saharan basins only Eosigmoilina was documented in
the E2 Zone. In North America, however, Eosigmoilina/
FIGURE 3—Stratigraphic ranges of the selected microfossils in the Mouarhaz section.
1110 JOURNAL OF PALEONTOLOGY, V. 85, NO. 6, 2011
Brenckleina first occur within the Menard Limestone in the
upper part of the Chesterian (Brenckle et al., 1982; Brenckle,
1990b), a limestone correlated with the Pendleian or E1 Zone
by Brenckle et al. (1977), Baxter and Brenckle (1982) and
Brenckle (1990a), but considered as late Serpukhovian
(Protvinskian) by Kulagina et al. (2008) (Fig. 14). In Scot-
land, Cummings in Lumsden and Wilson (1961), documented
the genus Eosigmoilina as Agathammina robertsoni from the
Blae Pot Limestone in the Archerbeck Borehole5Little
Limestone of the Pennines area in northern England. Also,
in England, in the Woodland Borehole, Eosigmoilina has
been recorded recently from the Rookhope Shell-Beds
Limestone (Stephenson et al., 2010).
Other typical late Serpukhovian foraminifers recognized
are Eostaffellina ‘‘paraprotvae’’ (Fig. 13.4–13.6), Eostaffellina
‘‘protvae’’ (Fig. 13.7, 13.8), Bradyina cribrostomata (Fig. 10.10),
Loeblichia minima (Fig. 11.15, 11.16) and Plectostaffella
(Fig. 13.9–13.13, 13.19, 13.20) (e.g., Conil et al., 1991).
Gibshman (2003), Brenckle (2005) and van Ginkel (2010)
suggested E. protvae and E. paraprotvae as synonyms because in
the holotype of the former, the specimen is missing the last half
volution, and thus, the wider form ratio was considered as an
artifact of the poor orientation and preservation. Due to the only
illustration of a single holotype in each species, this proposal is
difficult to demonstrate, and futher taxonomic studies will be
necessary in the stratotypes. Certainly, more compressed forms
are published in the literature as well as wider forms, and thus,
both diagnoses seem to be found in the stratigraphical record,
although the validity of the names proposed by Rauser-
Chernoussova (1948) can be questioned. Further studies should
be necessary to clarify the validity of both species and also, the
validity of both species names. In the present study both species
will be referred to in quotation marks.
Loeblichia minima was recorded in the early Serpukhovian
in the Reggane and Ahnet-Mouydir basins (Algeria) by Lys
(1975, 1985) but the illustrated specimens probably do not
correspond to the Russian forms. The oldest record in the
Russian Platform is from the Moscow Syncline, where it first
occurs at the top of the Steshevskian Horizon, close to the
early/late Serpukhovian boundary (Fig. 14). Loeblichia mini-
ma is recorded in the late Serpukhovian at Adarouch in
Idmarrach 1 and 2 sections (Figs. 6, 7).
Bradyina cribrostomata has been documented from the early
Serpukhovian in Algeria (Lys, 1985; Sebbar et al., 2000) but
the specimens illustrated are considered to be closer to B.
rotula. The earliest record of B. cribrostomata in the Donetz
Basin is at the top of the Samarskian (Fig. 14) (Brazhnikova
and Vdovenko in Aizenverg et al., 1983), considered as the
early/late Serpukhovian boundary. Bradyina cribrostomata is
recorded in the late Serpukhovian at Adarouch in Idmarrach 2
and Tirhela sections (Figs. 7, 8).
In the upper part of the Adarouch succession, E. post-
mosquensis (Fig. 12.5, 12.6) first occurs in the late Serpukho-
vian. A similar stratigraphic range is observed in the Urals
(Kulagina et al., 2001, 2009) and in Alaska (Harris
et al., 1997), whereas in the Donetz Basin it first occurs in the
FIGURE 4—Stratigraphic ranges of the selected microfossils in the Akerchi 1 section. Lithological legend as in Figure 3.
CO
´ZAR ET AL.—LATE VISEAN TO BASAL BASHKIRIAN FORAMINIFERS FROM MOROCCO 1111
upper part of the early Serpukhovian (Brazhnikova and
Vdovenko in Aizenverg et al., 1983), and in the Be´char Basin
in the Bashkirian (Lys, 1985).
Some of the specimens identified as Plectostaffella sp. are large,
close to P. varvariensis, however, they are slightly oblique sections
and do not permit a definite identification (Fig. 13.19, 13.20).
Bashkirian.—Seminovella (Fig. 13.14–13.16) was considered
by Conil et al. (1980, 1991) as a marker of the Bashkirian in
Western Europe and in North Africa (Lys, 1985). However, it
is present from the latest Serpukhovian in England (Grind-
stone Limestone, Stephenson et al., 2010). This could be a
laterally equivalent level to the Zapaltyubinskian Horizon in
the Donetz Basin (Fig. 14), where it is the oldest record of the
genus (Brazhnikova and Vdovenko in Aisenverg et al., 1983).
In general, in the Russian Platform it is Bashkirian in age
(Vachard and Maslo, 1996). In Adarouch, Seminovella
FIGURE 5—Stratigraphic ranges of the selected foraminifers in the Akerchi 2 section. Lithological legend as in Figure 3.
1112 JOURNAL OF PALEONTOLOGY, V. 85, NO. 6, 2011
elegantula occurs in the youngest limestone beds in Akerchi 2,
Idmarrach 2, and Tirhela sections (Figs. 5, 7, 8). Also recorded
but rare in these beds are Novella? sp. (Fig. 13.17, 13.18),
Globoomphalotis? (Fig. 10.11) and Globivalvulina bulloides (with
a differentiation of the wall in the inner whorls; Fig. 10.9).
The genus Novella is only known in the Bashkirian
(Vachard and Maslo, 1996; Maslo and Vachard, 1997),
although a rather similar specimen was illustrated from the
Lisbourne Group in Alaska, also identified as Novella? sp., by
Harris et al. (1997, fig. 7.23), about 16 m below the
Mississippian/Pennsylvanian boundary, a level that can be
interpreted as latest Serpukhovian. However, Baesemann et al.
(1998), studying the same outcrops, only recorded specimens
identified as Novella sp. in the Bashkirian.
Typical species of Globoomphalotis are described from the
middle and late Visean, and thus, its stratigraphic range does
not correspond to the Baskirian recorded in Adarouch.
However, the granular aspect of the wall, as well as the
tectoria differentiation, seem to be conspecific with a specimen
illustrated by Harris et al. (1997, fig. 8.8), as Endothyra sp., a
meter below the Morrowan/Atokan boundary (Bashkirian) of
the Lisburne Group of Alaska.
Globivalvulina bulloides usually first occurs in the late
Serpukhovian in Western Europe, where there was migration
to the midcontinent U.S.A., where it first occurs in the
Bashkirian (e.g., Brenckle et al., 1997a, 1997b). In between, in
the Arctic area, its first occurrence is situated a few meters
below the base of the Bashkirian, and thus, in the uppermost
latest Serpukhovian (e.g., Harris et al., 1997; Baesemann et al.,
1998). Its late occurrence in central Morocco is possibly related
to facies control or an artifact due to the scarcity of specimens.
In Idmarrach 2, the conodont Declinognathodus noduliferus
occurs at sample 2355 (5 m below the top of the limestones)
(Fig. 7). This conodont was used for defining the Mississippian/
Pennsylvanian GSSP in Arrow Canyon (Brenckle et al., 1997a,
1997b; Lane et al., 1999).
Comparison with other basins Western Europe.—Western
Europe is generally poor in Serpukhovian carbonates, except
FIGURE 6—Stratigraphic ranges of the selected foraminifers in the composite Idmarrach 3 and 1 sections (left column). Lithological legend as in Figure 3.
CO
´ZAR ET AL.—LATE VISEAN TO BASAL BASHKIRIAN FORAMINIFERS FROM MOROCCO 1113
for those of deep-water setting in the Cantabrian Mountains,
where apart from goniatites, only conodonts are biostrati-
graphically useful (e.g., Nemyrovska, 2005). In consequence,
the number of publications based on new data on the
Serpukhovian foraminifers is limited to a restricted suite of
papers (e.g., Laloux, 1988; Conil et al., 1991; Perret, 1993;
Krainer and Vachard, 2002; Co´zar, 2003; Poty et al., 2006; Pille,
2008). In addition, the correlation of the E1 (5Pendleian)
and E2 (5Arnsbergian) ammonoid biozones with the early/late
Serpukhovian of the Moscow Basin is not well established yet,
and different levels of correlation can be found in the literature
(Fig. 14). Thus, correlation can be found with the E1/E2 zonal
boundary equating with 1) the Steshevskian/Protvinskian
boundary (e.g., Aizenverg et al., 1979a, 1979b; Vachard and
Tahiri, 1991; Hecker and Osipova, 2007; Co´ zar et al., 2008b;
Nikolaeva et al., 2009b); 2) at a level within the middle part of
FIGURE 7—Stratigraphic ranges of the selected foraminifers in the Idmarrach 2 section. Lithological legend as in Figure 3.
1114 JOURNAL OF PALEONTOLOGY, V. 85, NO. 6, 2011
FIGURE 8—Stratigraphic ranges of the selected foraminifers in the Tirhela section. Lithological legend as in Figure 3.
CO
´ZAR ET AL.—LATE VISEAN TO BASAL BASHKIRIAN FORAMINIFERS FROM MOROCCO 1115
the Protvinskian (e.g., Vachard and Berkhli, 1992; Vdovenko,
2001; Nikolaeva and Kullmann, 2001; Krainer and Vachard,
2002; Alekseev et al., 2004); or 3) without any precise location
to the above-mentioned levels (e.g., Einor et al., 1979;
Vdovenko et al., 1990).
In Austria, Krainer and Vachard (2002) only recognized
Mississippian (late Serpukhovian) carbonates assigned to the
E2 Zone. This attribution was supported by some species of
Eostaffella,Eolasiodiscus dilatatus, and Bradyina concinna.A
similar stratigraphy was described in southern France (in
Ardengost) and Belgium (Tramaka Encrinite), although there,
exceptionally, some H to G and Westphalian A beds
(Chokierian to Langsettian) were recorded (see Laloux, 1988).
In the Montagne Noire and Mouthoumet Massif (southern
France), early Serpukhovian outcrops (Roc de Murviel, La
Serre-vignoble, and col de Couise) have been described based
on the foraminiferal and algal assemblages (e.g., Vachard
and Aretz, 2004; Pille, 2008), although they are not free of
controversy, because none of the listed taxa seem to be
exclusively Serpukhovian, and they might be upper Brigantian
rocks. The most reliable data come from the Ardengost
Limestone in the Pyrenees, already established as E2 Zone
(Perret and Vachard, 1977; Massa and Vachard, 1979; Perret,
1993; Pille, 2008), as well as the Tramaka Encrinite in Belgium
(Austin et al., 1974; Laloux, 1988). Late Serpukhovian markers
in those outcrops are mostly: ‘‘Turrispiroides’’, Loeblichia
minima (the single illustrated specimen by Laloux, 1988, is
questioned here), Brenckleina,andBrownediscus (endemic), and
hence the comparison with the rich assemblages of Morocco is
unsustained. Conil et al. (1991) (and Poty et al., 2006, who
mostly followed the same data) considered also Bradyina
cribrostomata as an E2 Zone marker, whereas apart from the
Ardengost section, the only record of the species in Western
Europe was documented by Laloux (1988) in the Bashkirian.
Globivalvulina s.s. was considered as a Bashkirian marker, while
Monotaxinoides and Archaediscus at tenuis stage were first
recorded in the latest Visean. We recognize the first occurrence
of Monotaxinoides in the latest Visean (see Co´ zar et al., 2005,
figs 8.16, 8.17), but this is in an exceptional case, and no
convincing illustrated specimen is known by us in the literature
from Western Europe. On the other hand, true A.attenuis stage
illustrated from Europe, are all included in E2 Zone outcrops,
except those recently published from Britain (e.g., Co´ zar and
Somerville, 2004; Co´zar et al., 2008b; Stephenson et al., 2010),
but always in E1 Zone outcrops in the Great Limestone, and
thus, they are exclusively considered as Serpukhovian.
A continuous Visean–Serpukhovian succession is recorded
in Britain, even though, up to the Baskhirian there are only
rare thin shallow-water carbonates, in predominantly silici-
clastic-rich sequences. These include the Midland Valley of
Scotland (Co´ zar et al., 2008b, 2010), and the Pennines in
northern England, with numerous natural sections and
boreholes (e.g., Rookhope Borehole, Co´ zar and Somerville,
2004; Woodland Borehole, Stephenson et al., 2010), as well as
in the transitional areas (Archerbeck Borehole, Lumsden
and Wilson, 1961). There, Monotaxinoides and Archaediscus
at tenuis stage occur from basal E1 Zone limestones,
while Eostaffellina and Plectostaffella are recorded from
rocks attributed to the Arnsbergian (E2). The FAD of
Eosigmoilina is in lower levels in E1 Zone, but it becomes
abundant in E2 Zone levels. It is also noteworthy, that rare
specimens of Seminovella,whichiscommonintheBashkir-
FIGURE 9—Summary of the stratigraphic ranges of the selected foraminifers in the Adarouch area. Dashed line5rare occurrences, solid line5common
occurrences. Stratigraphical stages and substages are situated in the left column. Bash.5Bashkirian.
1116 JOURNAL OF PALEONTOLOGY, V. 85, NO. 6, 2011
ian, occur at the top of a limestone of E2 Zone (Grindstone
Limestone in the Woodland Borehole), as well as Plectos-
taffella bogdanovkensis, a taxon which generally occurs in
the Bashkirian.
North American basins.—The establishment of the Missis-
sippian/Pennsylvanian boundary at Arrow Canyon, Nevada
led to detailed studies in the region and surrounding areas.
Although mostly based on conodonts, foraminifers were also
analyzed (Brenckle, 1973, 1990b; Brenckle et al., 1977, 1982,
1997a, 1997b; Skipp et al., 1985; Groves et al., 1999). To
establish the Visean/Serpukhovian boundary, American au-
thors used the recognition of Zone 17 of Mamet and Skipp
(1970), in which the main marker is the rapid development of
Asteroarchaediscus baschkiricus, although as the authors
recognized, some specimens can occur in older levels. Mamet
and Skipp (1970) assigned the Glen Dean Limestone of the
type Chesterian section (Kentucky, Mississippi Valley) to
Zone 17, and it was equated with the Ship Formation, situated
above the Battleship Wash Formation in Nevada. However,
Brenckle (1973) assigned the latter formation to Zone 17.
Later, Baxter and Brenckle (1982) recognized the acme of
A. baschkiricus within the Beech Creek Limestone-Haney
FIGURE 10—1,Pseudocornuspira sp. 1, sample Pc3267, Mouarhaz, late Brigantian; 2,Pseudocornuspira sp. 2, sample Pc3278, Mouarhaz, late
Brigantian; 3,Pseudocornuspira sp. 3, sample Pc3366, Mouarhaz, late Brigantian; 4,Ammovertella sp., sample Pc3390, Idmarrach 3, early Serpukhovian;
5,Calcivertella sp., sample Pc3195, Akerchi 2, late Serpukhovian; 6–7,Globivalvulina sp. 1: 6, sample Pc3391, Idmarrach 3, early Serpukhovian;
7, sample Pc3299, Tirhela, late Serpukhovian; 8, Globivalvulina parva Chernysheva, 1948, sample Pc2317, Idmarrach 1, late Serpukhovian; 9,
Globivalvulina bulloides (Brady, 1876), sample Pc3202, Akerchi 2, Bashkirian; 10,Bradyina cribrostomata Rauser-Chernoussova and Reitlinger in
Rauser-Chernoussova and Fursenko, 1937, sample Pc2325, Idmarrach 2, late Serpukhovian; 11,Globoomphalotis? sp., sample Pc3333, Tirhela, late
Serpukhovian; 12,Janischewskina delicata (Malakhova, 1956), sample Pc2325, Idmarrach 2, late Serpukhovian; 13,14,Endothyranopsis plana
Brazhnikova in Brazhnikova et al., 1967: 13, sample Pc3365, Tirhela, Bashkirian; 14, sample Pc3362, Tirhela, Bashkirian; 15,Insolentitheca horrida
(Brazhnikova in Brazhnikova et al., 1967), sample Pc3393, Idmarrach 3, early Serpukhovian; 16,Monotaxinoides subplana (Brazhnikova and Yartseva,
1956), sample Pc2338, Idmarrach 2, late Serpukhovian; 17, 22,‘‘Turrispiroides’’ spp.: 17, sample Pc2303, Idmarrach 1, late Serpukhovian; 22, sample
Pc3341, Akerchi 1, late Serpukhovian; 18,Monotaxinoides priscus (Brazhnikova and Yartseva, 1956), juvenile, sample Pc2771, Akerchi 2, late
Serpukhovian; 19, 21,Monotaxinoides ex gr. subplana (Brazhnikova and Yartseva, 1956): 19, sample Pc3309, Tirhela, late Serpukhovian); 21, sample
Pc2356, Idmarrach 2, Bashkirian; 20, Monotaxinoides cf. gracilis (Dain in Reitlinger, 1956), sample Pc2306, Idmarrach 1, late Serpukhovian. Scale
bar5100 microns.
CO
´ZAR ET AL.—LATE VISEAN TO BASAL BASHKIRIAN FORAMINIFERS FROM MOROCCO 1117
Limestone interval, underlying the Glen Dean Limestone
(Fig. 14). For other authors, the top of the Glen Dean
Limestone (the Tar Springs Sandstones) was correlated with
the Visean/Namurian boundary (Baxter et al. [1975] based on
foraminifers and Collinson et al. [1971] on conodonts).
Foraminiferal markers in the Tar Springs Sandstones for the
E1 Zone included the Archaediscus at tenuis stage by Baxter et
al. (1975), which were equated with the Archaediscus of the
Pennington Formation of Tennessee (also with Eosigmoilina;
see Hewitt and Conil, 1969). This archaediscid assemblage, in
turn, was compared with the Tramaka Encrinite of Belgium,
which is currently assigned to the E2 Zone. The first
appearance of ‘‘Millerella’’ tortula/designata was located
within the Glen Dean Formation. Nevertheless, Brenckle
(1990a) correlated the first occurrence of those species with the
first occurrence of Janischewskina in the Franco-Belgian
basins, in the V3c or Brigantian. The Menard Limestone
was correlated with the base of the Pendleian, due to the FAD
of Eosigmoilina and Brenckleina, whereas no latest Chesterian
markers were distinguished. Only in the upper part of the
uppermost Chesterian, are the earliest Globivalvulina (possibly
G. moderata) and ‘‘Turrispiroides’’ recognized. Monotaxinoides
and evolved Millerella (M. pressa and M. marblensis) were
already recognized in the Pennsylvanian, and not necessarily
from the basal beds (Skipp et al., 1985). Recently, Kulagina et
al. (2008) has correlated the occurrence of ‘‘Millerella’’ with
those from the basal Serpukhovian in the Zabory Quarry, and
considered the Beech Creek Limestone as the base of the
Serpukhovian, whereas the Menard Limestone was situtated
in levels equivalent to the late Serpukhovian. If the occurrence
FIGURE 11—1,Kasachstanodiscus sp., sample Pc3367, Tirhela, Bashkirian; 2, 4–6,Archaediscus spp. at tenuis stage: 2, P sample c3366, Tirhela,
Bashkirian; 4, sample Pc2775, Akerchi 2, late Serpukhovian; 5, sample Pc2774, Akerchi 2, late Serpukhovian; 6, sample Pc2376, Akerchi 1, early
Serpukhovian; 3,Archaediscus aff. velguriensis Grozdilova and Lebedeva, 1954, sample Pc3343, Tirhela, late Serpukhovian; 7,8,Brenckleina rugosa
(Brazhnikhova, 1964): 7, sample Pc3338, Akerchi 1, late Serpukhovian; 8, sample Pc2774, Akerchi 2, late Serpukhovian; 9,Neoarchaediscus postrugosus
(Reitlinger, 1949), sample 2369, Akerchi 1, late Brigantian; 10,11,Eosigmoilina sp.: 10, sample Pc3338, Akerchi 1, late Serpukhovian; 11, sample Pc2304,
Idmarrach 1, late Serpukhovian; 12–14,Euxinita pendleiensis Co´ zar et al., 2008b: 12, sample Pc3367, Tirhela, Bashkirian; 13, sample Pc3340, Akerchi 1,
late Serpukhovian; 14, sample Pc2358, Idmarrach 2, late Serpukhovian; 15,16,Loeblichia minima Brazhnikhova, 1962: 15, sample Pc2313, Idmarrach 1,
late Serpukhovian; 16, sample Pc2347, Idmarrach 2, late Serpukhovian; 17–20,Loeblichia ukrainica Brazhnikhova, 1956: 17,19, sample Pc2337,
Idmarrach 2, late Serpukhovian; 18, sample Pc2335, Idmarrach 2, late Serpukhovian; 20, sample Pc3392, Idmarrach 3, early Serpukhovian; 21–25,
Endostaffella? sp. 2: 21, sample Pc3337, Akerchi 1, late Serpukhovian; 22, sample Pc3375, Tirhela, Bashkirian; 23, sample Pc2774, Akerchi 2, late
Serpukhovian; 24, sample Pc3347, Akerchi 1, late Serpukhovian; 25, sample Pc3338, Akerchi 1, late Serpukhovian. Scale bar5100 microns.
1118 JOURNAL OF PALEONTOLOGY, V. 85, NO. 6, 2011
of Eosigmoilina is the key marker for the recognition of Zone
19 of Mamet, this implies that in the midcontinent, it would be
represented during the late Serpukhovian, contrary to the
early suggestion by Brenckle et al. (1977), in which, zones 17
to 19 would correspond to the early Serpukhovian. In
contrast, despite numerous published correlations of Mamet’s
zonal scheme, assigning Zones 17 and 18 to the early
Serpukhovian and Zone 19 to the late Serpukhovian, or Zone
17 to the early Serpukhovian and Zones 18–19 to the late
Serpukhovian (e.g., in Perret, 1993, fig. 129), the three zones
(17–19) correspond with the early Serpukhovian in the Tethys,
and no new zone exists for the late Serpukhovian.At the base
of the Bashkirian no diagnostic foraminifers exist in the so-
called ‘‘No Man’s Land’’ in Alaska as described by Mamet
and Batz (1989). Reitlinger (1980) observed a similar interval
lacking age diagnostic foraminifers in Europe. The Zone 20 is
the first Bashkirian zone, approximately equivalent to the
Bogdanovskian/Siuranskian horizons, which is characterized
by the incoming of the first Globivalvulina bulloides and large
Millerella in the Midcontinent (Skipp et al., 1985).
The fauna is distinct in the Arctic American basins/Taimyr-
Alaska Realm (e.g., Armstrong and Mamet, 1977). It occurs in
FIGURE 12—1,Eostaffella angusta Kireeva, 1949, sample Pc3304, Tirhela, late Serpukhovian; 2, 3,Eostaffella pseudostruvei Rauser-Chernoussova and
Reitlinger in Rauser-Chernoussova et al., 1936: 2, sample Pc2314, Idmarrach 1, late Serpukhovian; 3, sample Pc2347, Idmarrach 2, late Serpukhovian; 4,
Eostaffella acutiformis Kireeva in Rauser-Chernoussova et al., 1951, sample Pc3331, Tirhela, late Serpukhovian; 5,6,Eostaffella postmosquensis Kireeva
in Rauser-Chernoussova et al., 1951; 5, sample Pc3202, Akerchi 2, Bashkirian; 6, sample Pc2311, Idmarrach 1, late Serpukhovian; 7,Eostaffella aff.
postmosquensis Kireeva in Rauser-Chernoussova et al., 1951, sample Pc3390, Idmarrach 3, early Serpukhovian; 8,Eostaffella sp., sample Pc3311,
Tirhela, late Serpukhovian; 9, 10,Eostaffella cf. exilis Grozdilova and Lebedeva, 1950, Tirhela, late Serpukhovian: 9, sample Pc 3313; 10, sample Pc
3318; 11,‘‘Millerella’’? sp., sample Pc2304, Idmarrach 1, late Serpukhovian; 12,13, Endostaffella? sp. 1: 12, sample Pc3300, Tirhela, late Serpukhovian;
13, sample Pc3302, Tirhela, late Serpukhovian; 14, 15,Endostaffella?discoidea (Girty, 1915): 14, sample Pc3346, Akerchi 1, late Serpukhovian; 15,
sample Pc3198, Akerchi 2, late Serpukhovian; 16,‘‘Millerella’’ sp., sample Pc3329, Tirhela, late Serpukhovian; 17, 18,‘‘Millerella’’? sp. or Eostaffella?
sp., Tirhela, late Serpukhovian: 17, sample Pc3300; 18, sample Pc3326; 19, 20, ‘‘Millerella’’ aff. tortula (Zeller, 1953), sample Pc2375, Akerchi 1, early
Serpukhovian. Scale bar5100 microns.
CO
´ZAR ET AL.—LATE VISEAN TO BASAL BASHKIRIAN FORAMINIFERS FROM MOROCCO 1119
a transitional zone between the prolific Tethyan and the
impoverished sub-Arctic North American basins. This area
contains numerous taxa never or rarely encountered further
south (Harris et al., 1997), and conversely, explains the route
of migration and dispersion of typical North American taxa in
the Tethys. Despite this cosmopolitan distribution of bio-
stratigraphically useful taxa, they usually first occur in younger
rocks than in their respective realms, with a variable degree of
diachroneity (Harris et al., 1997; Baesemann et al., 1998).
Moscow Basin.—This is one of the best known basins due to
the fact that the Serpukhovian Stage was defined by Nikitin
(1890) in the southern outcrops of the Moscow Basin.
Unfortunately, the base of the Serpukhovian exposed in
Zabory Quarry contains a non-sequence and the uppermost
Serpukhovian is poorly developed in the basin due to the
erosion and non-deposition of the Pestovo ‘Series’ (Fig. 14),
in which foraminifers have been rarely studied (Fomina,
1977). Gibshman (2001) has proposed the subdivision of the
Tarusskian-Steshevskian-Protvinskian interval in four fora-
miniferal zones for the basin: Neoarchaediscus postrugosus,
Pseudoendothyra globosa,Eostaffellina decurta,andEostaf-
fellina ‘‘protvae’’ Zones. The basal zonal taxon, N. post-
rugosus shows different stratigraphic ranges elsewhere: e.g.,
in the Algerian basins and Kazakhstan, Bashkirian or late
Serpukhovian (Lys, 1985; Brenckle and Milkina, 2003);
latest Brigantian in Morocco (Co´ zar et al., 2008a), Britain
(Co´ zar et al., 2008b), Ireland (Co´ zar et al., 2005), and the
Urals (Kossovaya et al., 2001). P. globosa is restricted by
strong facies control (Vdovenko, 2001), and it is relatively
rare or absent at equivalent stratigraphic levels outside of
the Moscow Basin. Eostaffellina is recognized as a more
cosmopolitan genus, being recorded in the entire Paleo-
tethys, and at more or less equivalent levels. In Morocco,
more compressed Eostaffellina occur in older beds than E.
FIGURE 13—1–4,Eostaffellina sp.: 1,2sample Pc2769, Akerchi 2, early Serpukhovian; 3, sample Pc3364, Tirhela, Bashkirian; 4, sample Pc3230,
Akerchi 1, early Serpukhovian; 5–6,Eostaffellina ‘‘paraprotvae’’ Rauser-Chernoussova, 1948: 5, sample Pc3201, Akerchi 2, late Serpukhovian; 6, sample
Pc3338, Akerchi 1, late Serpukhovian; 7,8,Eostaffellina ‘‘protvae’’ Rauser-Chernoussova, 1948: 7, sample Pc3341, Akerchi 1, late Serpukhovian; 8,
sample Pc3364, Tirhela, Bashkirian; 9,10, Plectostaffella varvariensiformis tenuissima Brazhnikhova and Vdovenko in Aizenverg et al., 1983: 9, sample
Pc3311, Tirhela, late Serpukhovian; 10, sample Pc2345, Idmarrach 2, late Serpukhovian; 11–13, Plectostaffella varvariensiformis Brazhnikhova and
Vdovenko in Aizenverg et al., 1983: 11, 12, sample Pc3300, Tirhela, late Serpukhovian; 13, sample Pc3364, Tirhela, Bashkirian; 14,Seminovella aff.
elegantula Rauser-Chernoussova in Rauser-Chernoussova et al., 1951, sample Pc2356, Idmarrach 2, Bashkirian; 15,16,Seminovella elegantula Rauser-
Chernoussova in Rauser-Chernoussova et al., 1951: 15, sample Pc2776, Akerchi 2, Bashkirian; 16, sample Pc3377, Tirhela, Bashkirian; 17,18,Novella?
sp., sample Pc 3365, Tirhela, Bashkirian; 19,20, Plectostaffella sp.: 19, sample Pc2331, Idmarrach 2, late Serpukhovian; 20, sample Pc3311, Tirhela, late
Serpukhovian. Scale bar5100 microns.
1120 JOURNAL OF PALEONTOLOGY, V. 85, NO. 6, 2011
‘‘paraprotvae’’ or E. ‘‘protvae’’. Aisenverg et al. (1979a)
mentioned Eostaffellina in the Tarusskian Horizon, but no
species were determined. Reitlinger et al. in Einor (1996)
also acknowledged this record but they attributed it to
‘‘rounded eostaffellids.’’
Other taxa that first occur in the Tarusskian Horizon are
Pseudocornuspira (Cepekia auct.) and Planoendothyra aljuto-
vica (Gibshman, 2001). The former genus is well-known in
the Brigantian in the Ukraine (Vdovenko, 2001), whereas the
latter taxon is commonly attributed to the latest Serpukho-
vian or Bashkirian (Brazhnikova and Vdovenko in Aizenverg
et al., 1983; Kulagina et al., 2001). In Morocco, Pseudocor-
nuspira is commonly observed in thin-section from the late
Brigantian (Co´ zar et al., 2008a) and Rectocornuspira (cf.
Vdovenko, 2001) are rarely recognized in the late Serpukho-
vian of Idmarrach (only recorded as 3D cast samples
processed for conodonts). Gibshman (2001) identified Cepe-
kia from the Steshevskian Horizon. Although unrecorded by
the previous author, Aisenverg et al. (1979a) considered
Loeblichia ukrainica to be from the Tarusskian Horizon in the
Moscow Basin. Other taxa recorded from the Steshevskian
Horizon in the Moscow Basin by Gibshman (2001) are
Loeblichia minima,Eostaffellina paraprotvae,‘‘Millerella’’
tortula,andMillerella pressa. The latter species, is generally
used as a marker of the Bashkirian in the sub-Arctic North
American basins, in concert with other large species of
Millerella (e.g., Brenckle, 1990b), even its rarity in Arctic
North American basins prevents its use as a potential marker
(Harris et al., 1997).
For the Protvinskian Horizon, the key taxa include the well-
rounded Eostaffellina protvae and Eostaffellina shartimensis,
as well as Brenckleina.Eosigmoilina does not occur in the
Moscow Basin.
In Aisenverg et al. (1979a), Bradyina cribrostomata is also
recognized from the Protvinskian Horizon. Fomina (1977) is
the only study of the Pestovo foraminifers, an interval which
was characterized by the occurrence of new species of the genera
Eostaffella,Eostaffellina, and Pseudoendothyra. It is surprising
that while Howchinia occurs up to the Protvinskian, the
common Monotaxinoides species of the Donetz Basin and the
Urals have not been documented in the Moscow Basin.
Urals.—Eostaffella pseudostruvei is rarely recorded at the
top of the Visean, but is common in the Serpukhovian
(Reitlinger et al. in Einor, 1996). As mentioned previously,
Neoarchaediscus postrugosus first occurs in some outcrops
from the late Brigantian (Kossovaya et al., 2001), although in
other sections of the Urals, it first occurs in the lower
Serpukhovian Kosogorskian Horizon (Nikolaeva et al., 2001,
2009a; Fig. 14). Other taxa that have been documented in this
horizon are Globivalvulina minima and G. parva (Kossovaya
FIGURE 14—Correlation of the Serpukhovian stage and substages between North America, Western Europe, Moscow Basin, Urals, and Donetz
Basin. Correlation with Western Europe and North America is still controversial; source for the correlation is exposed and discussed in the text.
Correlation between Moscow, Urals, and Donetz basins are mostly based on Aizenverg et al. (1979a, 1979b), Nikolaeva et al. (2009a), Vdovenko (2001),
and Einor (1996). Mamet’s foraminiferal zones 17–20 shown in the midcontinent U.S.A. column, where the type Chesterian limestones in southern
Illinois are based on Brenckle et al. (2005) and its correlation based in Kulagina et al. (2008).
CO
´ZAR ET AL.—LATE VISEAN TO BASAL BASHKIRIAN FORAMINIFERS FROM MOROCCO 1121
et al., 2001; Reitlinger et al. in Einor, 1996), Monotaxinoides?
sp. (Nikolaeva et al., 2009a) and Eostaffellina subsphaerica
(Reitlinger et al. in Einor, 1996).
The Khudolozovian Horizon is correlated with the upper
part of the Steshevskian and the entire Protvinskian horizons
of the Moscow Basin (Fig. 14), and thus, the early/late
Serpukhovian boundary interval is included within the middle
part of this horizon (see Nikolaeva et al., 2009a). Taxa that
first occur in the Khudolozovian Horizon are numerous:
Monotaxinoides priscus,M. subplana,M. transitorius,Eostaf-
fellina decurta,E. paraprotvae,E. protvae,Eostaffella mirifica,
Plectostaffella varvariensis, as well as other primitive Plectos-
taffella, and Eosigmoilina. No major differences are observed
between the lower and upper part of this horizon, and thus,
the boundary between the early and late Serpukhovian cannot
be precisely correlated with the Moscow Basin.
The youngest horizon in the Urals is the Chernyshevian,
which is laterally equivalent to the Pestovo beds (Fig. 14). This
is characterized by numerous species of Monotaxinoides
unrecorded in lower levels, including M. gracilis and M.
planus (Nikolaeva et al., 2009a), Globivalvulina bulloides
(Groves, 1988; Groves et. al., 1999), Eostaffella postmosquen-
sis,Planoendothyra aljutovica,Globivalvulina kamaensis (Ku-
lagina et al., 2001), and Bradyina cribrostomata (Groves, 1988;
Kulagina et al., 2001).
Several new species of Eostaffella,Plectostaffella,Millerella,
and Plectomillerella are first recorded in the basal Bashkirian
Bogdanovskian Horizon in the type section in Bashkiria, south
Urals (Reitlinger, 1980; Groves, 1988; Kulagina et al., 2001).
Donetz Basin.—Four Serpukhovian substages are consid-
ered in the Donetz Basin, the Samarskian (equivalent to the
Tarusskian and Steshevskian horizons combined), the Pro-
khorovskian (lower part of the Protvinskian), Novolyubovs-
kian (upper part of the Protvinskian) and Zapaltyubinskian
(5Pestovo Beds) (Fig. 14).
In the Samarskian and older rocks of the Visean,
Brazhnikova and Vdovenko in Aizenverg et al. (1983),
Poletaev et al. (1991) and Vdovenko (2001) recorded the first
Rectocornuspira,Loeblichia ukrainica,Eostaffella ex gr.
pseudostruvei, and Monotaxinoides (M. subplana,M. subco-
nica,M. priscus,M. declivis; considered as Howchinia by
Vdovenko, 2001, except for M. priscus). In the top limestone
bed of this horizon (C
4
), Bradyina cribrostomata,Eostaffella
postmosquensis,E. pseudostruvei s.s., and Endothyranopsis
plana are recorded (Brazhnikova and Vdovenko in Aizenverg
et al., 1983). Brenckleina is also listed from the C
4
–C
5
lime-
stones. However, as well as Eosigmoilina, only specimens from
the D
3
Limestone or younger horizon (Zapatulyubinskian)
were illustrated. Poletaev et al. (1991) and Vdovenko (2001)
revised and reillustrated a specimen of Eosigmoilina, consid-
ered here as valid, from the C
5
Limestone. In contrast to older
publications, the C
5
Limestone was considered as the basal
part of the Prokhorovskian Horizon, and as the base of the
late Serpukhovian by Vdovenko (2001). Other markers of the
Prokhorovskian Horizon are Eostaffellina protvae,E. para-
protvae, and Eostaffella mirifica.
In the Novolyubovskian, only a few new taxa occur, such as
M. transitorius,Globivalvulina ex gr. pulchra, and G. ex gr.
eogranulosa (Brazhnikova and Vdovenko in Aizenverg et al.,
1983).
The Zapaltyubinskian Horizon can be characterized by
the first appearances of Loeblichia minima,Planoendothyra
spirillinoides,Globivalvulina moderata,Plectostaffella varvar-
iensis,P. bogdanovkensis (with other species and subspecies of
the genus), rare Millerella,Seminovella ex gr. elegantula, and
some new species of Eostaffellina (Brazhnikova and Vdovenko
in Aizenverg et al., 1983). Most Millerella and Seminovella
species do not occur up to the Bashkirian (Voznesenskian
Horizon) (Fig. 14).
Saharan basins and Tunisia.—Three main areas of Carbon-
iferous sedimentation can be distinguished in this region of
northern Africa (Fig. 1A): central Saharan basins (e.g.,
Reggane and Ahnet-Mouydir, southern Algeria), western
Saharan basins (e.g., Be´char and Tindouf, northwestern
Algeria/southern Morocco), and Tunisia. Taxa described and
used as markers in the Sahara have little in common with
Adarouch. These faunal differences could be attributed to
several factors: 1) inconsistent stratigraphic ranges for the taxa
in the same basin; 2) an incomplete foraminiferal database,
because in many publications, no location of the samples were
provided, only stratigraphic summaries of the species lists,
split by stages or substages (e.g., Lys, 1975, 1985; Sebbar,
1997; Mamet et al., 1994; Sebbar et al., 2000); 3) the number of
illustrated specimens is scarce, or poorly illustrated specimens
of key taxa such as Bradyina cribrostomata,Loeblichia minima,
and Eostaffella mirifica, or even non-existent illustrations of
Loeblichia ukrainica; 4) lack of a close calibration between
ammonoid and foraminiferal zonal schemes that might be
helpful in validating the stratigraphic ranges of the foramin-
ifers in the Sahara; and 5) strongly marked paleobiogeo-
graphic contrast north and south of the Atlas Transform
Fault.
Analysis of the foraminiferal taxa in the Saharian basins
and Tunisia suggest that some species may have arisen in
Algeria and hence are older than in the western Basins of
Russia, including: 1) Bradyina cribrostomata from the basal E1
Zone (Tindouf and Be´char, Lys, 1985; Sebbar, 1998; Mamet
et al., 1994; Sebbar et al., 2000), including the B. ex gr.
cribrostomata from the Visean (Lys, 1985); 2) Eosigmoilina
from the base of E1 Zone in Tindouf, Tunisia and Be´char
(Lys, 1975, 1985, 1988; Sebbar, 1998); 3) Eostaffella mirifica
from the basal E1 Zone in Tindouf and Reggane/Ahnet-
Mouydir (Lys, 1975, 1985; Sebbar, 1998; Sebbar et al., 2000);
4) Globivalvulina moderata from the basal E1 Zone in Tindouf,
Reggane/Ahnet-Mouydir, Tunisia (Lys, 1975, 1985, 1988;
Sebbar and Lys, 1989; Sebbar, 1998); 5) Loeblichia minima
also from the base of the E1 Zone in Be´char and Reggane/
Ahnet-Mouydir (Lys, 1975, 1985; Sebbar and Lys, 1989); and
6) Eostaffellina paraprotvae in the basal E1 Zone (Lys, 1985).
In addition, the use of the two zonal schemes (Mamet’s
foraminiferal zones and the classical Eumorphoceras ammonoid
Zone) also contributed more confusion to the data, because, as
was described in the North America sections, all the markers
defined for Mamet’s Zones 17, 18 and 19 are recorded in the E1
Zone, but in some publications the Zone 18 and/or 19 are
indistinctly used as equivalent to the E2 Zone in the Sahara
(e.g., Mamet et al., 1994; Sebbar, 1998; Sebbar et al., 2000). This
problem is actually even more acute, because the E1 and E2
zones and Mamet’s zones are not precisely calibrated with the
Serpukhovian substages of the Russian Platform (Fig. 14).
PERSPECTIVES FOR BIOMARKERS OF THE
VISEAN/SERPUKHOVIAN AND SERPUKHOVIAN/
BASHKIRIAN BOUNDARIES
Globivalvulina parva and other species such as G. minima or
G. moderata that in some works are considered as synonyms
(e.g., Sebbar et al., 2000) can have only limited use as potential
markers for recognizing the Visean/Serpukhovian (V/S)
boundary because primitive species of the genus first occur
from the late Brigantian, and even in older rocks. Other more
1122 JOURNAL OF PALEONTOLOGY, V. 85, NO. 6, 2011
evolved species with a clear light diaphanotheca occur much
later, close to the Serpukhovian/Bashkirian (S/B) boundary
(e.g., Globivalvulina bulloides). The new species G. sp. 1 with a
granular wall, seems to be endemic to Morocco, and has a
range from late Brigantian to Serpukhovian.
Neoarchaediscus postrugosus and Monotaxinoides species
can be potentially useful for recognizing the V/S and early/late
Serpukhovian boundaries respectively, but have different
stratigraphic ranges elsewhere.
In the case of ‘‘Millerella’’, although mostly a North
American genus, it seems to have a worldwide distribution,
but the Moroccan, European and Chinese species are difficult
to compare with the species recognized in the U.S.A. or
Canada.
In the case of the genus Loeblichia, some species first appear
in the Brigantian (e.g., L. paraammonoides,L. ukrainica). In
Adarouch L. ukrainica first appears later, in the early
Serpukhovian while L. minima occurs much later, around
the early/late Serpukhovian boundary (Fig. 9).
The archaediscids are still of great significance, not only for
the subdivision of the Visean, but possibly also for the
Serpukhovian. For recognizing the V/S boundary species of
Neoarchaediscus, particularly N. postrugosus, may be impor-
tant in North African basins, despite the fact that it is recorded
in uppermost Brigantian rocks in Europe. In Adarouch it first
occurs only a few meters below the V/S boundary. The
significance of Asteroarchaediscus baschkiricus in Zone 17, as
defined in Mamet’s zones, is minimized, because its FAD is
recognized earlier in Zone 16, a fact frequently demonstrated
in many Paleotethyan and North American basins. Neverthe-
less, the A. baschkiricus group is still important, because of its
worldwide distribution, and hence has several advantages.
Firstly, within the group, can be recognized 1) several
morphotypes/species; 2) different stages of evolution; 3)
different number of whorls; and 4) different development of
the microgranular layer, from A. pustulus to A. baschkiricus
s.s., and even other larger specimens with a reduced
microgranular layer (unnamed and usually included in A.
baschkiricus). These features also coincide with the reduction
of the number of pores in the fibrous layer, characteristic of
the Archaediscus at tenuis stage and also in the species of
Asteroarchaediscus and Neoarchaediscus (e.g., Hewitt and
Conil, 1969, pl. 2, figs. 31–33).
In addition, within the archaediscids there is another group,
the Archaediscus at tenuis stage. The use of this group in the
Brigantian by Conil et al. (1980) is confusing, because they
described ‘‘transitional forms to the Archaediscus at tenuis
stage’’, and later (Conil et al., 1991) they used those taxa as key
markers for the Cf7 Zone (Namurian). The only publication in
which various Archaediscus at tenuis stage were illustrated in
Western Europe is in Austin et al. (1974), from the Tramaka
Encrinite, and subsequently in Perret (1993) from the Arden-
gost Limestone, and always from the E2 Zone. The small
primitive specimens generally occurring at the base of the
Serpukhovian, and with uniform wall thickness from the inner
whorls seems to be derived from Archaediscus stilus, with some
tenuis final whorls that are already present in the Brigantian.
Furthermore, among the archaediscids, Eosigmoilina is also
an excellent marker as it has a worldwide distribution and is
relatively common, but its FAD is situated in the early
Serpukhovian in some basins, whereas in the majority it can be
used as marker for the early/late Serpukhovian boundary.
A similar case to Eosigmoilina seems to be Endothyranopsis
plana, occurring in younger beds than the current position of
the V/S boundary. However, it must be admitted that the
stratigraphic range and geographic distribution of this species
is not well established yet.
The FAD and species determination of the primitive
Eostaffellina is not well established in the literature. It would
be necessary to establish precisely if this FAD can be located
at the base of the Serpukhovian or in slightly younger beds, as
in the case of Eosigmoilina, which is not clear enough in the
published data.
In Western European basins, Endostaffella? sp. 2 is also
potentially an excellent marker for the base of the Serpukho-
vian, but this needs to be confirmed in other basins, especially
in Eastern Europe.
CONCLUSIONS
A continuous uppermost Visean to Bashkirian sequence is
recorded in the Adarouch area of central Morocco. It is a
predominantly limestone and shale sequence passing up at the
top into red bed siliciclastic rocks.
The Adarouch sequence is one of the few successions in
Morocco where limestones have yielded a rich suite of late
Brigantian and Serpukhovian foraminifers, algae, corals and
conodonts.
The Visean/Serpukhovian boundary at Adarouch is marked
by the first appearance of Eostaffellina sp., Eostaffella
pseudostruvei, and Archaediscus at tenuis stage. It is also
characterized by unusual and rare taxa such as ‘‘Millerella’’,
Endostaffella? sp. 1, Endostaffella? sp. 2, Insolentitheca horrida,
and Loeblichia ukrainica, which might have more local
regional significance. In slightly higher beds in the early
Serpukhovian, Endothyranopsis plana first occurs.
An early/late Serpukhovian boundary is recognized at
Adarouch, based on the first appearance of Brenckleina,
Eosigmoilina,Loeblichia minima,Bradyina cribrostomata,
Plectostaffella spp., Eostaffellina ‘‘paraprotvae’’, E. ‘‘protvae’’,
‘‘Turrispiroides’’, and locally by Monotaxinoides spp. Higher
in the late Serpukhovian is recorded Eostaffella postmosquensis
and Globoomphalotis?. Also recorded in this E2 zonal interval is
the zonal conodont Gnathodus bilineatus bollandensis.
The Serpukhovian/Bashkirian (Mississippian/Pennsylvanian)
boundary at Adarouch is marked by the first occurrence of rare
Seminovella elegantula and Novella? The base of the Bashkirian is
marked by the zonal conodont Declinognathodus noduliferus.
Comparisons are made between the Adarouch area and other
Paleotethyan regions containing Serpukhovian-Bashkirian sec-
tions, including the Sahara, Donets Basin, Moscow Basin and
the Urals, as well as North America.
A significant aspect of the Adarouch succession is that it is
composed of thick limestone intervals, unlike many other
Mississippian sequences in the western Paleotethys, which are
usually characterized by cyclothems dominated by siliciclastic
rocks, in which limestones form only a minor component.
Hence, it is much more likely to be able to trace evolutionary
lineages in foraminifers in Adarouch.
The Adarouch succession represents a late Mississippian–
Pennsylvanian sequence from the northern margin of Gond-
wanaland, but yet shows all the characteristic taxa of western
Paleotethyan basins, and even shows some affinities with North
American successions, particularly in the Serpukhovian.
ACKNOWLEDGMENTS
The authors would like to thank the collaboration of the
Department of Mine Development of the Ministe`re de l’Energie
et des Mines of Morocco, who kindly gave us permission to
carry out field work. Also, thanks to M. Howe (Chief Curator,
Palaeontological Unit, Keyworth) and M. Dean (Palaeonto-
CO
´ZAR ET AL.—LATE VISEAN TO BASAL BASHKIRIAN FORAMINIFERS FROM MOROCCO 1123
logical Unit, Edinburgh) of the British Geological Survey who
kindly allowed the examination of the material from the
Woodland and Archerbeck boreholes. This research was
supported by the projects CGL2006-03085 and CGL2009-
10340 of the Spanish Ministry of Research and Innovation. We
thank the two reviewers D. Altiner and J. Groves for their
helpful comments which significantly improved this paper.
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