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Devonian Brachiopods of The Tamesna Basin (Central Sahara Algeria and North Niger) - Part 1
Devonian Brachiopods of The Tamesna Basin (Central Sahara Algeria and North Niger) - Part 1
MICHAL MERGL
Department of Biology, University of West Bohemia, Klatovská 51, Plzeň, 30614, Czech Republic; e-mail:
[email protected]
DOMINIQUE MASSA
Universite de Nice 6, Rue J. J. Rousseau, Suresnes, 92150, France
Mergl, M., Massa, D. (2004): Devonian Brachiopods of the Tamesna Basin (Central Sahara; Algeria and North Niger). Part
1. – Acta Mus. Nat. Pragae, Ser. B, Hist. Nat. 60 (3-4): 61-112. Praha. ISSN 0036-5343.
Abstract. The Devonian brachiopod fauna from samples collected by the late Lionel Lessard in the sixties from several secti-
ons of the Lower Palaeozoic bordering the Tamesna Basin (Algeria and North Niger, south of the Ahaggar Massif) is descri-
bed. The fauna is preserved in siliciclastic rocks, often lacking fine morphological details that are necessary for the
determination of the fauna. Despite this difficulty, in total 40 taxa have been determined to generic, species or subspecies le-
vels. One new chonetoid genus Amziella is defined on newly described species A. rahirensis. The new species Arcuaminetes
racheboeufi, Montsenetes pervulgatus, Montsenetes ? drotae, Pustulatia lessardi, Pustulatia tamesnaensis, Eleutherokomma
mutabilis, Mediospirifer rerhohensis and the new subspecies Tropidoleptus carinatus titanius are described. The Pragian age
is suggested for the earliest brachiopod association, with maximum spread and diversity of the brachiopod fauna in Late Em-
sian – Early Eifelian interval. The youngest Devonian brachiopod fauna in the available samples is probably of the Givetian
age. The Upper Devonian succession is lagoonal and poorly fossiliferous and precedes the basal marine Carboniferous. The
taxonomic composition of fauna indicates an affinity to the Middle Devonian brachiopod faunas of the Eastern Americas Re-
alm. However, in the Upper Pragian and Emsian ages, the influence of the Malvinokaffric Realm is documented by the pre-
sence of several genera (Lomaella, Pustulatia, Pleurothyrella, Orbiculoidea).
■ Brachiopoda, taxonomy, Devonian, Emsian, Eifelian, Tamesna Basin, Algeria, Niger
Received November 12, 2004
Introduction of this report. The paper is divided in two parts. Part 1 pre-
sents the taxonomic work of the brachiopod fauna from the
Devonian fossils of the Tamesna Basin have been known
surface localities along the north border of the Tamesna
for over forty years but they have never been satisfactorily
Basin.
described. This extensive desert area near Algeria and North
Part 2 deals with the subsurface brachiopod data from
Niger borders remains poorly known to present time. The
the boreholes drilled in the Tamesna Basin, important non-
results of the geological mapping of the area have been pub-
brachiopod fossils from the surface and subsurface samples,
lished by Lionel Lessard (1959, 1961, 1962). He also pre-
locality data, geological setting, stratigraphy, benthic com-
sented the first more detailed stratigraphical data
munity ecology and palaeobiogeographic affinity of the fau-
concerning the Devonian in this area. He recognized the un-
na. Only a short overview of the geology and stratigraphy is
conformity between the Silurian shales and the transgres-
given in Part 1.
sive Devonian beds and he also attempted to define the
Lower/Middle and Middle/Upper Devonian boundaries in
the area, based on lithological and palaeontological data. Geological setting
Since this time, a few other authors have dealt with geology The Ténéré du Tamesna Basin, an extensive desert area
and poor brachiopod fauna (Drot 1986, Racheboeuf 1990b, near Algerian and Nigerian frontiers, is poorly known to
Herrera et Racheboeuf 2001, Mergl et Massa in print). The present time because of very difficult access. The geologi-
original Lessard (1962) stratigraphical data have been cal results in this area have been partially published, as far
called in question, mostly on the data from chonetoid bra- as the Palaeozoic is concerned, roughly between 1960 and
chiopods (Racheboeuf 1990a, 1990b, Herrera et 1985 (Lessard et Joulia 1959, Lessard 1959, 1961, 1962,
Racheboeuf 2001). Reboul et al. 1962, Claret et Tempere 1968, Greigertt et
One of the authors (D. M.) strove to bring together the Pougnet 1967a, 1967b, Perrodon 1971). Several papers and
material of the original Lessard’s samples and compiled the unpublished reports were mainly devoted to Uranium,
data of his unpublished reports. The second author (M. M.) Water or Hydrocarbons research works of Bigotte et Obel-
described and evaluated the brachiopod fauna of these sam- lina (1968), Valsardieu (1970) and Mokkadem (1990).
ples. The attempt to elucidate the geological setting and age The Ténéré du Tamesna Basin is located south of the
of the samples based on brachiopod fauna is the main aim Ahaggar Massif (Text-fig. 1). It is usually mentioned in the
61
Text-fig. 1. Devonian outcrops in the Saharan platform and NW Africa with location of referred bassins and outcrops (modified
after Boucot et al. 1983 and Racheboeuf 2004).
literature as the “Tamesna Basin”. The northern part is situ- The Cambrian and Ordovician units were defined in the
ated in South Algeria and it is called “Oua-n-Ahaggar Tas- Algerian Tassili-n-Ajjer area and refer particularly to the
sili”, a local Tuareg name. It is the north margin of the basin basin contribution (Beuf et al. 1971). According to Lessard
widely covering the North Niger (Iullemeden intracratonic et Joulia (1959), the Unit II is considered the Middle and
Basin). The eastern limit of the Tamesna Basin corresponds Upper Cambrian in age (equivalent of the Ajjers Forma-
with the Precambrian Aïr Massif. The western limit forms tion). The Unit III is of the Lower and Middle Ordovician
the Adrar-des-Iforas Massif, also of the Precambrian age, age (equivalent of the In Tahouite Formation). The Unit IV
studied by Karpoff (1960). The north-south In Guezzam is of the Upper Ordovician age (equivalent of the Tamadjert
Uplift is located in the central part of the basin separating Formation). Its diversified marine fauna has been referred
the eastern wide Tin Seririne syncline from the western by Gatinsky et al. (1966) to the Caradocian and Ashgillian
area, sometimes called Azouak in the literature. but it has not been described and only approximate faunal
Above the Precambrian of the Ahaggar Massif several list is available. The major unconformities are known north
formations are known comprising the Cambrian, Ordovi- and south of the Ahaggar Massif (Deynoux et al. 1985).
cian, Silurian, Devonian, and Carboniferous. The total The Silurian was discovered quite early (Kilian 1928).
thickness, without the Carboniferous and the Mesozoic, is This graptolite-bearing unit is present and mapped in the
some 1200–1500 m. A particularly interesting is the com- Oua-n-Ahaggar belt of outcrops (Claret et Tempere 1965).
parison between the Palaeozoic outcrops in northern side of Rapid lateral thickness variations often occur, with ranges
the Ahaggar Massif (Tassili-n-Ajjer), the eastern side of the from a few meters to almost 250 metres. The thickness vari-
Ahaggar Massif (western flank of the Murzuq-Djado Basin) ations are related with the important erosion of the Upper Sil-
and the southern margin (Tamesna Basin). They have a urian (Caledonian tectonic phase; Bellini et Massa 1981).
great significance for the regional, stratigraphic and palaeo- The age of these Silurian sequences is lower and middle
geographic purposes, as they belonged originally to the Llandoverian (Deynoux et al. 1985). A revision of the avail-
same “peri-Ahaggarian” area. The selected lithostratigraphy able graptolite collections is in preparation (Štorch et Massa,
and chronostratigraphy terminology of these units, as used in prep.). In complete uneroded sections it is possible to rec-
in present work, is given in Text-fig. 3. ognize equivalents of the “Tanezzuft” and “Acacus” facies,
62
Text-fig. 2. Geological map of of the north margin of the Tamesna Basin with the sections bearing the Devonian fauna.
TD – Toufine and In Debirene section, RB – South Tin Rerhoh Anou n’Bidek section, RA – In Rahir Tin Amzi section, OF – Oued
Felaou section, IA – In Ateï Anou Izileg section, TI – Taberia Touaret section, AZ – In Azaoua section, NWA – North West Aïr.
1 – Precambrian, 2 – Cambrian and Ordovician, 3 – Silurian, 4 – Devonian, 5 – Carboniferous, 6 – post-Carboniferous units.
described from the Western Murzuq Basin of South Libya strata were recognized south of the Ahaggar Massif in the
(Massa 1988) and the new name Tin Tarabine Formation is Tamesna Basin, preceding the Carboniferous marine cycle
proposed for this succession restricted to the Tamesna Basin. (Valsardieu 1970).
The lowest sequence of the Devonian lacks marine fos-
sils but it is rich in plant remains and carbonised woods Repository
(Idekel Formation). The age of this unit is indistinct but the
All specimens, including the types and figured ones are
Lochkovian is probable. In the western part of the Tamesna
part of the Massa and Mergl collection (abbreviation
Basin, in the Oua-n-Ahaggar belt, there the fine grained
MaMe), housed as a separate part in the palaeontological
sandstones and the associated carbonates are commonly
collection of the Department of Palaeontology of the Muse-
fossiliferous (Bazoche et Blain 1961). Trilobites, bivalves,
um of West Bohemia, Plzeň, Czech Republic.
brachiopods and tentaculites were collected from these
beds. A Pragian age of this unit (Touaret Formation) seems
adequate taking into consideration the previous publica- Acknowledgements
tions, mainly of Lessard et Joulia (1959). Above this lowest The authors have greatly appreciated the gift of the orig-
unit, the Emsian, Eifelian, Givetian and Upper Devonian inal documents and sections of the late L. Lessard, made by
63
D e s c r i p t i o n : The shell is large, with planar dorsal
valve having a weakly concave apical region. The pedicle
track is long, narrowly elliptical, with a distinct large listri-
um. Ornament consists of almost regularly spaced low con-
centric lamellose rugellae, 0.6 to 0.9 mm apart. Weak
growth fila are present in interspaces.
Remarks: This species is poorly known but its weakly
concave dorsal valve indicates an affinity to the genus Roe-
merella HALL et CLARKE, 1890. Large discinoids with
planar ventral valve are reported but otherwise poorly
known in the Lower Devonian. They are characteristic for
siliciclastic facies in several parts of the Malvinokaffric
Realm (Boucot et al. 2001). Apart of this realm, Kayser
(1892) described the large discinoid Discina siegenensis
KAYSER, 1892 from the Siegenian of Germany. This is
similar to the shell from the Tamesna Basin but until re-
cently it was neither revised nor illustrated and the original
description and drawings are of a poor quality.
D i s t r i b u t i o n : Upper Pragian.
O c c u r r e n c e : Tamesna Basin, Tin Rerhoh area (sec-
tion 3) (sample H 1079).
Orbiculoidea sp.
Madame Lessard-Moureaux. They are greatly indebted to P.
R. Racheboeuf from Université de Bretagne Occidentale, M a t e r i a l : Fourteen complete shells in sandstone.
Brest, France for sending, pro parte, the brachiopod materi- Remarks: This giant discinoid is described in a separate
al; to A. J. Boucot from the Oregon State University, Cor- paper (Mergl et Massa, in print). It represents a new species
vallis, Oregon, USA for useful comments. of the new genus, characteristic by a highly conical dorsal
valve and planar ventral valve with a very short pedicle
Abbreviations track.
DvL – length of dorsal valve; DvW – width of dorsal D i s t r i b u t i o n : Upper Pragian.
valve; VvL – length of ventral valve; VvW – width of ven- O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec-
tral valve; n – number of measured specimens. tion (sample H 1053).
64
shell is 3.5 mm wide, broadly oval with weakly convex dor- and lateral margins and obtuse cardinal extremities. The
sal and more convex ventral valve. Exterior is coarsely and cardinal process is prominent, with a pair of prominent
distantly lamellose. Interior of the ventral valve has broad lobes faced posteroventrally, extending from the short and
elevated brim along posterior and lateral margins. Ventral broad median ridge. The myophore is coarsely lamellose.
visceral field is deeply impressed in posterior half of shell, The adductor scars are large, elongate, extending in 30 % of
with large oblique crescentic anterior adductors and smaller the valve, bounded anterolaterally by a low ridge. The field
and less distinct other scars. is posterolaterally subsided below the densely pustulose
R e m a r k s : The genus has broad stratigraphic as well shell interior.
as geographic distributions. It extends from the late Ordovi- The ventral valve is subcircular, about 82 % as long as
cian to early Carboniferous, with the maximum spread in wide. The hinge line is straight, 85–100 % as wide as the
the late Silurian and early Devonian. This is the first report valve. Lateral and anterior margins are evenly curved. The
of this genus from the Devonian of North Africa. ventral interarea is distinct, low, steeply apsacline, with
D i s t r i b u t i o n : Upper Emsian. dense vertical striation that corresponds to numerous denti-
O c c u r r e n c e : Tamesna Basin, Tin Rerhoh area (sec- cles along the whole length of the hinge line. There are
tion 5) (sample H 1087). 12–14 striae and denticles, respectively, per 5 mm. The
delthyrium is small, open and narrowly triangular with
Subphylum Rhynchonelliformea WILLIAMS et al., densely and finely striated bottom of delthyrial chamber.
1996 The ventral muscle field is widely triangular, about 40 %
Class Strophomenata WILLIAMS et al., 1996 wide as the valve. The muscle scars are weakly impressed
Order Strophomenida ÖPIK, 1934 on the valve floor, being posterolaterally bounded by stout,
coarsely pustulose and widely divergent ridges, which sub-
Superfamily Strophomenoidea KING, 1846 tend about 80° angle between each other. The diductor im-
Family Leptostrophiidae CASTER, 1939 pressions are partitioned by fine but remarkably high radial
ridges of uneven length into narrow lobes. There are about
Genus Mesoleptostrophia HARPER et BOUCOT, 11–15 distinct lobes apparent in large well-preserved spec-
1978 imens, with smaller, less clearly defined lobes along the an-
terior periphery of the muscle field. The ventral process is
T y p e s p e c i e s : Mesoleptostrophia kartalensis massive, consisting of two short proximally high divergent
HARPER et BOUCOT, 1978; Lower Devonian, Emsian; ridges. The fine myophragm is low, broad and disappears
Turkey. near the midlength of the valve. The adductor scars are
minute, contiguous to proximal part of the myophragm.
Mesoleptostrophia (M.) explanata The shell interior is densely covered by fine pustules
(SOWERBY, 1842) that anterolaterally decrease in size.
Pl. 1, figs 1–7 The ornamentation is parvicostellate, with 8–10 costel-
lae per 5 mm anteromedianly. Concentric ornament consists
cf. 1842 Leptaena explanata; SOWERBY, J. B., p. 409, pl. 38,
of fine, slightly wavy growth fila over entire surface. There
fig. 15.
are no distinct concentric rugellae. The broad band along
cf. 1971 Leptostrophia explanata (SOWERBY, 1842); Jahnke,
p. 55, pl. 3, figs 2, 4, 5 (here older synonymy of the Euro- the shell periphery exhibits distinct internal costellation.
pean occurrences). R e m a r k s : The genus Mesoleptostrophia HARPER et
1983 cf. Leptostrophia (Leptostrophiella) explanata (SOWERBY, BOUCOT, 1978 (=Leptostrophiella HARPER et BOUCOT,
1842); Brice et al., p. 448, pl. 12, fig. 19. 1978) is known since the Silurian to the Emsian, with close-
1986 cf. Leptostrophia (Leptostrophiella) explanata (SOWERBY, ly related species in the different parts of the word. Brice et
1842); Drot, p. 513, pl. 1, figs 17, 18. al. (1983) reported less favourably preserved valves of the
M a t e r i a l : Six dorsal and seven ventral valves and nu- genus in North Morocco, and Drot (1986) referred with
merous fragments, some with original shell but mostly pre- some uncertainty to the species M. explanata a specimen
served as internal and external moulds in calcareous from the sample H 1088 collected by Lionel Lessard in the
sandstones. northern flank of the Tamesna Basin. Stratigraphically
D e s c r i p t i o n : The shell is planoconvex, very large younger leptostrophiids of the Upper Emsian and Eifelian
with the measured width of 96 mm in the largest specimen ages are currently referred to subgenus M. (Paralep-
available, thin-walled relative to shell size, with the maxi- tostrophia), which is smaller and have posteriorly faced
mum width located near or gently anterior to the hinge line. lobes of cardinal process (Cocks et Rong 2000).
The commissure is rectimarginate. The outlines of several The giant species of Mesoleptostrophia are especially
available shells are somewhat irregular, concerning espe- known from the Lower Emsian. Jahnke (1971) described,
cially the location of maximum width. figured and discussed the affinity of Mesoleptostrophia ex-
The dorsal valve is subcircular, 75–80 % as long as planata (SOWERBY, 1842) from the Siegenian and Emsian
wide, with straight posterior margin, evenly curved anterior of Harz, Germany. This author mentioned various taxa that
65
need modern revision and were formerly incorporated under Order Productida SARYTCHEVA
this species name. Jahnke (1971) noted the probability that et SOKOLSKAYA, 1959
during phylogeny of M. explanata in Siegenian to Emsian Suborder Chonetidina MUIR-WOOD, 1955
ages the ventral diductors were split by prominent radial
ridges. Having taken in account his suggestion, our speci- Superfamily Chonetoidea BRONN, 1862
mens belong to typical form of M. explanata by its large Family Anoplidae MUIR-WOOD, 1962
size and numerous radial ridges partitioning the ventral Subfamily Caenanoplinae ARCHBOLD, 1980
muscle field.
The upper range of the subgenus and the species de-
serves a discussion. Our specimens have been found in two
Genus Arcuaminetes BIZZARO, 1995
different fossil associations, probably of different age. The Ty p e s p e c i e s : Chonetes scitula HALL, 1857; Mid-
specimens in our samples are never associated with dle Devonian, Givetian; New York, USA.
Paraspirifer and Longispina and only one specimen is asso-
ciated with chonetoid Montsenetes pervulgatus sp. n. This
Arcuaminetes racheboeufi sp. n.
may be explained by the disappearance of the species before
Pl. 3, figs 17–24
the spread of the early Eifelian fauna in the area.
In Spain and Anti-Atlas of Morocco, the species ap- H o l o t y p e : Ventral valve (MaMe 0183A), figured on
peared for the first time in the Lower Emsian, extended to pl. 3, figs 17, 21, housed in the Department of Palaeontolo-
the latest Upper Emsian and disappeared before the end of gy of the Museum of West Bohemia, Plzeň, Czech Repub-
the Lower Devonian (Brice et al. 1983, Pardo et Garcia-Al- lic.
calde 1984, Garcia-Alcalde et Truyols 1994, Garcia-Al- P a r a t y p e : Dorsal valve (MaMe 0185), figured on pl.
calde 1997, 2001). The specimens from Germany are also 3, fig. 22, housed in the Department of Palaeontology of the
of the Emsian age (Jahnke 1971, Weddige et al. 1979). The Museum of West Bohemia, Plzeň, Czech Republic.
earliest but tentative report of the species is from the Ge- Ty p e h o r i z o n : Devonian, Eifelian.
dinian (Boucot 1960). The species M. explanata is never re- Ty p e l o c a l i t y : Algeria, Tamesna Basin, Tin Rerhoh
ported from beds of the Eifelian or younger age. As our area, section 5, sample H 1090.
specimens have the same morphology as the Emsian speci- E t y m o l o g y : After Dr. Patrick Racheboeuf, the fa-
mens in Europe, we also suggest their Upper Emsian age. mous chonetid brachiopod specialist.
However, because of the associated fauna, some specimens M a t e r i a l : Some twenty-five internal and external
from the Tamesna basin (sample H 884) may be also of the moulds, mostly of ventral valves, and numerous fragments
early Eifelian age. preserved in ferruginous sandstone.
The morphologically similar but stratigraphically D i a g n o s i s : The shell small, thin-shelled; dorsal valve
younger species Mesoleptostrophia (M.) casteri deeply concave, with short median septum and a pair of
(BENEDETTO, 1984) is reported from the upper Eifelian longer accessory septa; ventral valve deeply convex, with
of Venezuela. This species represents the youngest report of strong myophragm extending into low median ridge toward
the genus hitherto known. This species is also associated anterior margin; ventral muscle field weakly impressed.
with a chonetoid Montsenetes, represented by the species D e s c r i p t i o n : The shell is small, in maximum 9 mm
M. notius (BENEDETTO, 1984). The ventral valve of M. wide, thin shelled, strongly concavo-convex.
casteri differs from our specimens by less divergent ridges The dorsal valve is deeply concave, deepest at mi-
bounding the ventral muscle field, by coarser radial ridges dlength, widest in posterior third, with evenly curved mar-
partitioning the ventral diductor scars, lack of dorsal medi- gins. The dorsal interarea is very low, hypercline. The
an ridge and different shape of the cardinal process. These cardinal process is ventrally bilobed, short, supported by a
features also distinguish M. casteri from European speci- short, low median ridge. The shallow cardinal process pit is
mens referred to M. explanata. present. The median ridge is short, becoming indistinct
D i s t r i b u t i o n : The Emsian age of rocks bearing M. from shell midlength. Dental sockets are large, bordered an-
explanata is suggested, with most reports referred, to the teriorly by short and small socket ridges. Anderidia are
Upper Emsian. The species is usually associated with Tropi- short and weakly divergent. Accessory septa are well-devel-
doleptus, various chonetoids and spiriferoids. However, the oped, each with a distinct radial row of coarse endospines.
large strophomenoid shells are mostly incomplete and may, Several similar radial rows of prominent and toward com-
eventually, belong to more than one species. The differences missure larger endospines are located more laterally. Three
in associated fauna indicate that M. explanata occurs with- radial rows of endospines are intercalated between the ac-
in longer stratigraphical interval, probably from the Emsian cessory septa.
to early Eifelian. The ventral valve is broadly oval, with maximum width
O c c u r r e n c e : Tamesna Basin, Kori-Idemeg – Touaret in posterior third. The valve is strongly convex, weakly de-
area (sample H 884), In Ateï section (sample H 1003), In pressed posterolaterally, with maximum depth in posterior
Rahir Tin Amzi section (sample H 1058), Tin Rerhoh area, fourth. The VvW/VvL is 1.35 (range: 1.25 to 1.42; n = 12).
section 5 (samples H 1086, H 1087). Cardinal extremities are rectangular in small to medium
66
sized specimens becoming obtuse in large shells. The ven- anopliid indet.
tral interarea is low, apsacline to almost orthocline. Spines Pl. 4, fig. 22
are orthomorph oblique, known only in their proximal parts.
M a t e r i a l : Two dorsal valves in sandstone.
Three strongly posteriorly convergent canals (inner aper-
R e m a r k s : Poorly preserved internal moulds show dis-
tures of hollow spines) penetrate the shell on each side of
tinct, high accessory septa and absence of median septum in
the ventral umbo. The inner three pairs of apertures are ar-
weakly concave dorsal valve. Although the material is not
ranged in short intervals, the fourth pair of apertures is situ-
determined in generic level, the presence of an anopliid is
ated in much longer distance from the umbo. Teeth are
noteworthy.
massive, triangular, and widely divergent. The myophragm
D i s t r i b u t i o n : Givetian.
is thick posteriorly, anteriorly it continues to low, rounded
O c c u r r e n c e : Tamesna Basin, In Ateï section (sam-
median ridge nearly toward anterior commissure. The dor-
ples H 820 and H 1006).
sal edge of the myophragm is sharply acute. The ventral
muscle field is weakly impressed, with undivided diductor
scars and small, weakly impressed semielliptical adductor Genus Plicanoplia BOUCOT et HARPER, 1968
scars in posterior three-fourth of the valve. Almost whole
Ty p e s p e c i e s : Chonetes fornacula DUNBAR, 1920;
interior shows traces of external costellation that become
Lower Devonian, Emsian; Tennessee, USA.
coarse along the commissure. Radial rows of endospines are
short and finer than in dorsal valve, restricted into narrow
band along margins. Plicanoplia sp.
Ventral valve exterior bears some 40 costellae, half of Pl. 5, fig. 14
them extending from the umbo. Costellae are rounded, sep-
M a t e r i a l : Two internal moulds of ventral valves in
arated by shallow rounded interspaces. New costellae orig-
sandstone.
inate mostly by bifurcation in the ventral valve. The
R e m a r k s : Two poorly preserved shells have features
ornament of the dorsal valve is similar but new costellae
characteristic for the genus: small size, subtrigonal outline,
originate by implantation. There are 6 to 7 costellae per 2
extreme convexity of ventral valve and a few weak costae.
mm anteriorly.
It is most similar to P. fornacula (DUNBAR, 1920) by small
C o m p a r i s o n : This species is referred to Ar-
size and subtrigonal outline. However, other details of mor-
cuaminetes BIZZARO, 1995 although the type species is
phology remain unknown and thus generic attribution of
less convex and has different ornament of the ventral valve,
these valves remains tentative. The genus ranges from the
with the new costellae mostly originating by intercalation.
Pragian to Emsian; in Venezuela it ranges into the Upper
In the new species the costellae of the ventral valve origi-
Emsian (Benedetto 1984).
nate by bifurcation. The conformity of the new species with
D i s t r i b u t i o n : Upper Emsian.
the species A. scitulus (HALL, 1857) concerns the presence
O c c u r r e n c e : Tamesna Basin, Tin Rerhoh area, sec-
of three radial rows of endospines between accessory septa.
tion 5 (sample H 1087).
Although the shell of A. racheboeufi sp. n. is much smaller
than of other chonetoids in the area, the high convexity,
clearly impressed muscle scars and upper size limit (8.5 – Family Eodevonariidae SOKOLSKAYA, 1960
9.0 mm width) of the available shells indicate the maturity
of the preserved individuals. This difference indicates a dif- Genus Lomaella HERRERA, 1995
ferent age from specimens A. ? deynouxi RACHEBOEUF et
al., 2004 described from the Givetian of Libya and Mauri- Ty p e s p e c i e s : Lomaella primoris HERRERA, 1995;
tania (Havlíček et Röhlich 1987, Mergl et Massa 1992, Lower Devonian, Pragian; Argentina.
Racheboeuf et al. 2004). By size and convexity, the species
is also similar to other coenanoplines, especially to Lomaella amziensis HERRERA et RACHEBOEUF,
Caplinoplia HAVLÍČEK et RACHEBOEUF, 1979 but this 2001
genus has more clearly developed two pairs of accessory Pl. 2, figs 13, 14, 16–18
septa and its ornament is coarser. Small specimens of
Devonochonetes ? sp. from suggested Givetian Akara For- 1990a Loreleiella sp.; Racheboeuf, p. 320.
2001 Lomaella amziensis nov. sp.; Herrera et Racheboeuf, p. 499,
mation of the Air Massif (North Niger) (Boucot et al. 1983)
figs 7 and 8: 1–10.
may belong to the same or related species. It differs from
similar Montsenetes pervulgatus sp. n. of the Eifelian age M a t e r i a l : Six ventral valves and three dorsal valves
by smaller size, less convex ventral valve and weaker preserved as internal and external moulds in fine sandstone
myophragm in the ventral valve. and several fragments of shells showing details of morphol-
D i s t r i b u t i o n : Eifelian. ogy in calcareous sandstone.
O c c u r r e n c e : Tamesna Basin, Tin Rerhoh area, sec- R e m a r k s : This species was described in detail by Her-
tion 5 (sample H 1090). rera et Racheboeuf (2001), with description based and ma-
67
terial figured from the sample H 1052. They also reported Genus Amziella gen. n.
its range extended to some 60 m higher beds (sample H Ty p e s p e c i e s : Amziella rahirensis sp. n.; Lower
1057). These authors suggested only the late Emsian age of Devonian, Pragian; Algeria.
the beds bearing the species. However, the species is known E t y m o l o g y : After the type area of the type species in
from two distinct fossil associations. The older brachiopod In Rahir Tin Amzi.
association near the base of the Devonian succession (sam- S p e c i e s r e f e r r e d : Amziella rahirensis sp. n.; Up-
ple H 1052) yielded dominant spiriferoid Filispirifer merza- per Pragian; Algeria. ? Plicanoplia sp.; Early Emsian, Tala-
khsaiensis JANSEN, 2001 that indicates the Pragian age. casto Formation, Argentina.
The younger occurrences of Lomaella amziensis are from D i a g n o s i s : The shell is moderately concavo-convex,
samples H 1086 and H 1087, in the association with a large medium sized, with shortly alate ventral valve; radial orna-
chonetoid Montsenetes (?) drotae sp. n., small Plebejo- mentation of subangular costae originating from beak ex-
chonetes sp., spinocyrtiid Subcuspidella and strophomenoid cept of a few lateral costae originating above the posterior
Mesoleptostrophia, indicating a younger, probably Upper margin; costae rarely branching by bifurcation in ventral
Emsian age. Thye fragmentary state of preservation of Lo- valve and weakly widen anteriorly; ventral interarea almost
maella from younger beds does not allow satisfactory com- orthocline, dorsal interarea almost catacline, very low;
parison with the specimens from sample H 1052 and these hinge teeth large, directed anterolaterally; myophragm
younger shells may even represent another species. The short, distinct; dorsal interior with short septum supporting
whole range of L. amziensis from the Tamesna Basin is ventrally bilobate cardinal process; fused endospines form a
roughly the same as the known occurrence of the genus in pair of accessory septa; cardinal process pit shallow; an-
South America. According to Herrera (1995), Lomaella ex- deridia short, weakly divergent; interiors with prominent ra-
tends from the late Pragian in the Argentine Precordillera to dial rows of large endospines.
the Upper Emsian of Bolivia. C o m p a r i s o n : The new genus is similar to Plebejo-
D i s t r i b u t i o n : Upper Pragian to Upper Emsian. chonetes BOUCOT et HARPER, 1968 from which it is prob-
O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec- ably derived. The common features of Plebejochonetes and the
tion (samples H 1052, H 1086, H 1087). new genus comprise lateral costae crossing the posterior mar-
gin of the valves, dichotoming ventral costae, deeply bilobed
Family Chonetidae BRONN, 1862 cardinal process and short dorsal median septum (Boucot et
Subfamily Chonetinae BRONN, 1862 Harper 1968). The new genus is distinct by alate cardinal ex-
tremities in adult specimens, coarser ornament of high suban-
Genus Plebejochonetes BOUCOT et HARPER, 1968 gular costae and coarsely spinose interior with endospines
arranged into discrete radial rows. Genus Plebejochonetes is
Ty p e s p e c i e s : Chonetes plebeja SCHNUR, 1854; represented by several species in the Emsian of southwest
Lower Devonian, Emsian; Germany. Europe. Among them the Spanish species P. collensis
GARCIA-ALCALDE et RACHEBOEUF, 1978 differs by fin-
Plebejochonetes sp. er radial ornament and a weak ventral myophragm. The species
Pl. 5, figs 10, 11 P. moniellensis GARCIA-ALCALDE et RACHEBOEUF,
1978 has a less distinct internal costation and much finer inter-
M a t e r i a l : Two internal and two external moulds of
nal endospines. The morphologically nearest species is P. bu-
ventral valves in sandstone.
choti (RENAUD, 1942) from the Lower Emsian of the
D e s c r i p t i o n : Shell is small, 8 mm wide, broadly
Armorican Massif. It has rather coarse costation but its costae
semielliptical, widest at midlength, with obtuse cardinal ex-
are rounded and, apart from the flanks, the new costae are
tremities. The valve is moderate convex. Ventral interior
added mostly by intercalation. The anopliid genus Ar-
bears high and short myophragm. Ornament of ventral valve
cuaminetes BIZZARO, 1995 has similar shell morphology as
consists of about 15 simple costae, with posterolateral
the new genus but its external ornament is weaker, it has low
costae crossing the posterior margin and only some five to
cardinal process and the ventral valve is not alate. The size of
six costae extending from the umbo. Spines have not been
the new genus is also larger, unusual for anopliides.
observed in our material.
The poorly known species Plicanoplia sp. from the ear-
R e m a r k s : The origin of posterolateral costae is an im-
ly Emsian (Talacasto Formation) of Argentina may belong
portant feature of the genus. Therefore, although poorly pre-
to a new genus (Herrera 1995). It has comparatively coarse
served, the available shells are referred to Plebejochonetes
ornament but is smaller and its interior is poorly known.
BOUCOT et HARPER, 1968. By size and coarse ornament
our specimens are most similar to P. moniellensis GARCIA-
ALCALDE et RACHEBOEUF, 1978 from the Upper Em-
Amziella rahirensis sp. n.
Pl. 2, figs 1–12
sian of the Cantabrian Mountains, Spain.
D i s t r i b u t i o n : Upper Emsian. H o l o t y p e : Ventral valve (MaMe 0011A), figured on
O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi pl. 2, fig. 10, housed in the Department of Palaeontology of
(sample H 1057). the Museum of West Bohemia, Plzeň, Czech Republic.
68
P a r a t y p e : Dorsal valve (MaMe 0062), figured on pl. The new costae on ventral valve originate mostly by asym-
2, fig. 9, housed in the Department of Palaeontology of the metrical bifurcation, only rarely by an implantation. They
Museum of West Bohemia, Plzeň, Czech Republic. rapidly reach the same size as the primary costae. There are
Ty p e h o r i z o n : Lower Devonian, Upper Pragian. 28–30 costae distinct from the umbo, with 5 to 6 costae per
Ty p e l o c a l i t y : Algeria, Tamesna Basin, In Rahir Tin 5 mm anteromedianly. The large shells have 37 to 40 costae.
Amzi section, sample H 1052. D i s t r i b u t i o n : The species is a dominant element in
E t y m o l o g y : After the type area. samples from the lowest transgressive fossiliferous beds of
M a t e r i a l : Twelve dorsal valves, fifteen ventral valves the Devonian succession in four localities. These beds are a
and many fragments, often poorly preserved in fine sand- good stratigraphical marker. The Upper Pragian age is sug-
stone. gested.
D i a g n o s i s : Shell of medium size, coarsely costellate O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec-
with prominent subangular costae originating at the apex tion (sample H 1052), Tin Rerhoh area, sections 2 and 5
and above the posterior margin; hinge line shortly alate; (samples H 1074, H 1076, H 1085).
shell interior with discrete radial rows of coarse endospines.
D e s c r i p t i o n : The shell is moderately concavo-con-
Subfamily Devonochonetinae MUIR-WOOD, 1962
vex, rectimarginate, medium sized and thin-walled. The
largest preserved specimen attains 14 mm width but the av- Genus Devonochonetes MUIR-WOOD, 1962
erage shell size is smaller, ranging from 10 to 12 mm. Max-
imum width is at the hinge line, which is extended into short Ty p e s p e c i e s : Chonetes coronatus HALL, 1857;
but distinct ears. Middle Devonian, Hamilton Group; New York, USA.
The dorsal valve is moderately concave, with a flattened
posterior disc. DvW/DvL is 1.42 (range: 1.33 to 1.50; n = Devonochonetes cf. salemi MERGL et MASSA,
8). The dorsal interarea is very low, almost catacline, with
1992
notothyrium filled by the cardinal process. The cardinal
Pl. 4, figs 12–14
process is ventrally distinctly bilobed but details of its pos-
teriorly faced myophore remain unknown. The median sep- M a t e r i a l : Four dorsal valves, six ventral valves, and
tum is low and coarse, extending to midlength anteriorly, numerous fragments preserved as internal and external
laterally with a pair of weakly divergent, higher accessory moulds in sandstone.
septa of fused endospines. The anderidia are short, low and D e s c r i p t i o n : The shell is large, 20 mm wide in the
stout, diverging anteriorly in 45–50° angle. The dental sock- largest specimen, moderate concavo-convex, widest at
ets are large, deep, anteriorly bordered by stout inner sock- hinge line, rather thin-shelled.
et ridges which diverge each other in 150° angle. Outer The dorsal valve is slightly concave, with rectangular to
socket ridges are united with the hinge line. The visceral broadly angular cardinal extremities. The dorsal interarea is
area is distinct in large specimens as a large, semicircular hypercline, low. The notothyrium is filled with the cardinal
and gently elevated platform, extending anteriorly to two- process. The valve is weakly convex or almost flat in the
thirds of the valve length. Endospines are very coarse, in- posterior half. The cardinal process is moderately high, with
clined anteromedianly, arranged in simple radial rows. posteriorly faced quadrilobate myophore. The process con-
The ventral valve is about 80 % as long as wide, tinues anteriorly into the low and short median ridge, which
VvW/VvL is 1.34 (range: 1.25 to 1.47; n = 8). Anterior and disappears posterior to the shell midlength.
lateral margins are evenly curved. The valve is strongly The ventral valve is broadly oval, with evenly rounded
arched in a lateral profile and strongly convex in a trans- lateral and less rounded anterior commissure. VvW/VvL is
verse profile. Cardinal extremities are slightly depressed. 1.38 (range: 1.28 to 1.48; n = 3). The valve is moderately
The beak is short. The ventral interarea is low, apsacline, a convex in a lateral profile and evenly convex in transverse
small delthyrium is widely triangular and apically closed by profile with depressed cardinal extremities. The ventral in-
a small, highly convex pseudodeltidium. There are coarse terarea is anacline to almost orthocline, low, with a small
canal apertures on each side of the umbo, converging poste- delthyrium. The umbo is short. Spines are orthomorph
riorly, three to four on each side. Spines are cyrtomorph in- oblique but known only after their broken bases, rapidly
traversed. The ventral valve interior has stout teeth and curved dorsally. The ventral muscle field is weakly im-
lacks distinctly bordered visceral area. The median septum pressed, divided by a short and low myophragm. Visceral
is clearly defined posteriorly but weak anteriorly, in the pos- area is distinctly bordered by a low edge on valve floor. In-
terior third it is flanked by narrow and weak semielliptical ner costellation is distinct along the periphery in the ex-
adductor scars. Diductor scars are indistinct. Coarse en- travisceral area.
dospines are arranged in simple radial rows in the extravis- Ornamentation consists of fine, high and rounded costel-
ceral area. lae of uniform size, separated by deep subangular inter-
The shell exterior is covered by coarse subangular spaces. New costellae originate by bifurcation in the ventral
costae of uniform size, anteriorly regularly extending in valve and by intercalation in the dorsal valve. There are 5
size, originating at the apex and above the posterior margin. costellae per 2 mm anteromedianly. Surface of flattened
69
cardinal extremities is without costellation, with only weak anteriorly and decreasing in size laterally, becoming ob-
concentric growth lines. scure near cardinal extremities. Anteromedianly there are
R e m a r k s : The species D. cf. salemi MERGL et 3–4 costae per 2 mm.
MASSA, 1992 is similar by moderate concavo-convex pro- R e m a r k s : The genus is known from the Eifelian to
file, weak myophragm and poorly impressed ventral muscle Givetian of North and South America, Europe and north-
field and weak median ridge in dorsal interior and by cardi- western Africa (Garcia-Alcalde et Racheboeuf 1978,
nal extremities without radial ornament to the typical spec- Benedetto 1984, Racheboeuf 1990a, Racheboeuf et Feld-
imens of D. salemi MERGL et MASSA, 1992 from the man 1990, Racheboeuf et Isaacson 1993, Brice et al. 1994).
Givetian-Frasnian transition of the Ghadamis Basin. There Our specimens are most similar by coarse costellation to
are not distinct differences between Libyan and our speci- Longispina mucronata (HALL, 1843) from the Eifelian of
mens apart of less transverse outline and a more prominent New York, USA but it differs by mucronate cardinal ex-
inner costellation of D. salemi. tremities and less numerous costae. A specimen similar to
The species is also similar to the type species D. coro- our species has recently been described from a suggested
natus (HALL, 1857) from the Middle Devonian of New Givetian strata of Hodh area of Mauritania (Racheboeuf et
York, USA by a transverse outline, moderate convex ventral al. 2004). The presence of Longispina in our material indi-
and weakly concave dorsal valves but there are significant cates the post-Emsian age.
differences. In D. cf. salemi the radial ornament is absent in In the studied area, small strongly convex shells which
cardinal extremities and the maximum width is at the hinge are referred to this genus are known from two samples. In
line. Stratigraphically earlier and morphologically similar sample H 1405 several ventral valves are known, two with
chonetids in the Tamesna Basin are referred to Montsenetes spines preserved. In sample H 1404 similarly shaped but
RACHEBOEUF, 1992 but both newly described species poorly preserved one ventral valve without spines is known.
(M. pervulgatus sp. n. and M. ? drotae sp. n.) are more con- In both samples the associated devonochonetine
vex and their maximum width is more anterior, the cardinal Montsenetes pervulgatus sp. n. and a spiriferoid
extremities being always obtuse. The number of costellae Eleutherokomma support suggested the Eifelian age of the
per 2 mm anteriorly is also different in D. cf. salemi and in samples.
both species of Montsenetes. D i s t r i b u t i o n : Eifelian.
D i s t r i b u t i o n : Givetian. O c c u r r e n c e : Tamesna Basin, Oued Felaou area
O c c u r r e n c e : Tamesna Basin, In Ateï (samples H 820 (samples H 1404, H 1405).
and H 1006).
Genus Montsenetes RACHEBOEUF, 1992
Genus Longispina COOPER, 1942
Ty p e s p e c i e s : Devonochonetes notius BENEDET-
Ty p e s p e c i e s : Chonetes emmetensis WINCHELL, TO, 1984; Middle Devonian, Eifelian; Venezuela.
1866; Middle Devonian; Hamilton Group; New York, USA.
Montsenetes pervulgatus sp. n.
Longispina sp. Pl. 3, figs 1–16
Pl. 4, figs 15–21 1983 Devonochonetes ? sp.; Boucot, Massa et Perry; p. 112, pl. 3,
M a t e r i a l : Two dorsal and three ventral valves, pre- figs 28–33.
served as internal and external moulds in ferruginous sand- H o l o t y p e : Ventral valve (MaMe 0120), figured on pl.
stone. 3, figs 1, 5, housed in the Department of Palaeontology of
D e s c r i p t i o n : The shell is small, 6.5 mm wide, with the Museum of West Bohemia, Plzeň, Czech Republic.
maximum width at hinge line, strongly concavo-convex, Ty p e h o r i z o n : Devonian, Eifelian.
thin-shelled. Ty p e l o c a l i t y : Algeria, Tamesna Basin, In Ate_ sec-
The dorsal valve is deeply concave, widest at the hinge tion, sample H 1017.
line. Its interior is unknown. The ventral valve is semicircu- E t y m o l o g y : pervulgatus (Lat.) – allusion to the
lar (VvW/VvL is 1.3), with regularly curved margins. Car- abundance of the species.
dinal extremities are rectangular. The shell is strongly M a t e r i a l : Several tens of specimens in various state
convex, with depressed posterolateral edges. The ventral in- of preservation.
terarea is rather high, nearly orthocline. Teeth are small, an- D i a g n o s i s : Large, thick-shelled and strongly conca-
terolaterally supported by short and widely divergent ridges. vo-convex Montsenetes with transverse shell outline, mod-
The myophragm is high, short and not exceeding one-third erate-sized ventral myophragm, weak and short dorsal
of the valve length. The ventral muscle field is poorly im- median septum, short anderidia and evenly and coarsely
pressed. Spines are orthomorph parallel, but their number costellate shell.
and arrangements is unknown because only spine bases (pl. D e s c r i p t i o n : Shell is large, thick-shelled, in maxi-
4, figs 19, 20) have been observed. Exterior bears some mum 25 mm wide, strongly concavo-convex, thick-walled.
16–18 simple, undivided, rounded costae, evenly widening Anterior commissure is rectimarginate to very weakly
70
unisulcate in gerontic specimens. The shell shape consider- Ventral interior has a rather small, short and widely di-
ably changes with ontogeny. Small to medium sized ventral vergent teeth. Adductor muscle scar field is semielliptical,
valves are moderately convex. In adult specimens the ven- rather small, with longer, narrower and deeply impressed
tral valve becomes strongly convex, with subsequent growth posterior pair, and smaller, triangular anterior pair (pl. 3, fig.
dorsally directed. 10). The adductor muscle field is divided by a strong, acute
The dorsal valve is deeply concave, with flattened cardi- and long myophragm, which diminishes anteriorly and con-
nal extremities, broadly semielliptical, with maximum tinues as a low median ridge nearly to anterior margin.
depth at midlength. The DvW/DvL is 1.43 (range: 1.20 to Diductor muscle scars are large, flabellate, with several ra-
1.77; n = 6). The dorsal interarea is low, hypercline, with dially arranged lobes that extend over shell midlength. The
small notothyrium closed by small chilidium and the poste- ventral visceral area is large, occupying over 80 % of valve
rior face of the cardinal process. The cardinal process is length, bordered by a weak edge. Extravisceral shell interi-
large, quadrilobed, with high, thin inner pair of lobes and or is covered by regularly arranged internal radial costella-
about half-sized lateral lobes. The myophores are very high, tion intercalated by short rows of endospines. The visceral
faced posteriorly. The cardinal process is supported by a area is almost smooth.
low and proximally thick median ridge, which becomes an- Ventral valve exterior is coarsely costellate, with round-
teriorly thinner and higher. The ridge disappears at about ed costellae of uniform size, separated by narrow and deep
shell midlength. Dental sockets are small, deep, bordered interspaces. Some 10–12 costellae extend from the umbo,
posteriorly by small outer socket ridges. Inner socket ridges the new costellae originate exclusively by bifurcation; two
are obsolete, short, diverging anteriorly in about 90–100° new costellae rapidly attain the same size as the primary
from each other. Anderidia are short, thin, diverging at costella. The symmetric bifurcation is common at the same
about 40° from each other. Two rows of coarse endospines growth line, especially during earlier growth periods. Orna-
fused together to form ridges similar to accessory septa are ment of the dorsal valve consists of subangular costellae of
developed from shell midlength. Five weak radial rows of uniform size, separated by wider, rounded interspaces. The
endospines are intercalated between the anterior parts of costellae are of uniform size, with new ones originating by
these rows of endospines. Dorsal muscle field is weakly im- intercalation, commonly near the same growth line. There
pressed, with shallow triangular imprints of anterior adduc- are some 10–12 costellae extending from the umbo, and 3–4
tor between the median ridge and anderidia. costellae per 2 mm anteriorly. The large shells bear 60–65
The ventral valve is broadly semielliptical, recti- distinct costellae. The costellation is developed over all sur-
marginate, with obtuse cardinal extremities. The VvW/VvL face of the valve except of smooth depressed small corners
is 1.28 (range: 1.12 to 1.50 in juvenile shell; n = 21). Hinge near the cardinal extremities. Concentric ornament is not
line is some 90–95 % of the width. The maximum width is present. However, inside the shell wall the concentric wavy
located in posterior one-third to one-fourth. Transverse pro- lines are present. These lines are distinct in partly exfoliat-
file is strongly and evenly convex, with steeply sloping ed shell and probably are weakly expressed on internal shell
flanks and evenly convex axial part. Only small corners near surface, judging after some excellently preserved internal
cardinal extremities are depressed. Lateral profile is strong- moulds.
ly convex, with the maximum height at midlength. Margins R e m a r k s : New species is similar to the type species
are evenly rounded. The commissure is rectimarginate. The Montsenetes notius (BENEDETTO, 1984) from the Eifelian
ventral umbo is directed posteriorly to posterodorsally in of Venezuela by costellate ornamentation, shell interior and
small shells. In large shells it is directed dorsally to even convexity. These features indicate near affinity of both
weakly anterodorsally, slightly overhanging the posterior species. The main difference is a coarser ornament of the
margin. The ventral interarea is low, orthocline in small new species, with only 11–12 costellae per 5 mm anteriorly
shells but well anacline in large shells, with a weakly con- while 14–18 costellae are present in M. notius. In M. notius,
cave profile. Its surface bears fine growth lines. The the ventral median ridge is longer and more prominent. The
delthyrium is apically closed by a small pseudodeltidium. species M. boliviensis RACHEBOEUF, 1992 from the
Spines are fine and short, cyrtomorph intraversed to cyrto- Givetian of Bolivia differs from the new species by coarser
morph geniculated, with various incline toward the com- costellation, with only 7–9 costellae per 5 mm and by a
missure plane. Unfortunately, only their proximal parts are stronger median ridge on the dorsal interior.
usually preserved. Their direction may be different even in The species Devonochonetes ? sp. from the Akara For-
the same shell, with spines rapidly, just above the hinge line mation of Air Massif (Boucot et al. 1983) may belong to the
inverted posterodorsally to spines of cyrtomorph inverted new species. This species has the shell of a medium size (up
type. The spines extend in irregular, gradually increasing to 25 mm wide), highly convex ventral valve with the same
distance from the umbo in both sides of the shell, number- type of ornament and its interior is consistent with the oth-
ing three to (?) five on each side in largest shells. The first er species of Montsenetes RACHBOEUF, 1992. A difficul-
(preserved) spine (1’) appears some 1.3 mm from the umbo ty represents the age of this species. The Akara Formation is
with the next spines in various distances (2’ = 2.3 mm, 3’ = suggested of a Givetian age (Boucot et al. 1983) but the
5.5 mm, 4’ = 7.3 mm; 1 = 2.3 mm, 2 = 4.0 mm, 3 = unpre- specimens from In Guezzam area are supposed of an Eife-
served; specimen MaMe 0016). lian, even a Late Emsian age. However, the undoubted
71
Givetian fauna from the Aïr Massif is represented in the um is wide, apically closed by a chilidium. Cardinal process
younger Amesguer Sandstone by a terebratulid Rhypi- is large, high, quadrilobate posteriorly, with a posterodor-
dothyris africana BOUCOT et al., 1983. This species is sally faced myophore. Inner lobes of myophore are high and
known from Ghadamis and Murzuq Basins of Libya narrow, lateral lobes are much lower, broader. Inner cristae
(Boucot et al. 1983) from beds of a suggested early Givetian are prominent. Cardinal process is anteriorly supported by a
or even late Eifelian age. This indicates, that the Akara For- robust base of median ridge that extends into tapering ridge
mation, or at least its part, might be older, of an Eifelian age. over two-thirds of the valve. Outer socket plates are short
D i s t r i b u t i o n : The new species is very abundant; and transverse, inner socked ridges are thick, united with
sometimes it is even the commonest brachiopod in the sam- the cardinal process, widely diverging anterolaterally. Den-
ples. Its shells form low-diversity, almost monospecific ac- tal sockets are large and deep. Anderidia are long, strong,
cumulations (e.g. samples H 1017, H 358) from valves in diverging anteriorly at 40–50°. Inner surface of the valve is
various ontogenetic stages. The shells are associated with finely costellate along the commissure, with radial rows on
various fauna, usually with spiriferids Eleutherokomma, uniformly sized small endospines.
Paraspirifer and Pustulatia. The fauna by the presence of The ventral valve is strongly convex, widest at posterior
some genera indicates an Eifelian age, probably near to the third, with flattened posterolateral parts. The VvW/VvL is
Emsian/Eifelian boundary. In this respect, the new species 1.38 (range: 1.24 to 1.57; n = 13). Cardinal extremities are
is stratigraphically older than other known species of the obtuse to rectangular, depending on shell size. The valve
genus (M. notius is from the Upper Eifelian-Lower Give- margins are evenly rounded, slightly less anteromedianly.
tian, M. boliviensis from the Givetian). The ventral interarea is anacline, low and straight, distinct,
O c c u r r e n c e : Tamesna Basin, Kori-Idemeg – Touaret with fine growth lines. Hinge line is without denticles. The
section (sample H 884), In Ateï section (samples H 1004, H delthyrium is apically closed by a small pseudodeltidium.
1006, H 1016, H 1017), In Debirene section (samples H Spines are orthomorph, regularly spaced, three on each side
1106, H 1107), Oued Felaou area (samples H 1404, H in medium-sized specimens (the available large specimens
1405). have always broken spines). The spines are rapidly but un-
evenly turned ventrally. Fine thin and widely diverging
canals penetrating the shell wall are situated near the ventral
Montsenetes ? drotae sp. n.
umbo, three on each side, not extending into spines. Inner
Pl. 4, figs 1–11
apertures of spines are widely funiculate. The ventral interi-
H o l o t y p e : Ventral valve (MaMe 0086A), figured on or has short, ridge-like, obsolete and broadly diverging
pl. 4, fig. 3, housed in the Department of Palaeontology of teeth. The ventral muscle field is large, divided by posteri-
the Museum of West Bohemia, Plzeň, Czech Republic. orly distinct, low and rounded myophragm that becomes
P a r a t y p e : Dorsal valve (MaMe 0032B), figured on pl. finer in its anterior part. The adductor muscle field is
4, fig. 5, housed in the Department of Palaeontology of the semielliptical, located in the posterior third. It is differenti-
Museum of West Bohemia, Plzeň, Czech Republic. ated into parallel, more distinct posterior pair and less dis-
Ty p e h o r i z o n : Lower Devonian, Upper Emsian. tinct, triangular anterior pair that is sometimes divided into
Ty p e l o c a l i t y : Algeria, Tamesna Basin, Oued Fe- two lobes. The diductor scars are weakly impressed, of nu-
laou area, sample H 328. merous narrow, radially arranged lobes. Visceral area is
E t y m o l o g y : After Dr. Jeaninne Drot, in honour of the large, with evenly rounded and distinctly defined lateral, an-
late outstanding French palaeontologist. terior and anteromedian borders. Only a narrow slit remains
M a t e r i a l : Ten dorsal valves and eleven ventral valves between the visceral lobes along the shell axis. Internal
and numerous fragments preserved in sandstone. costellation is distinct near shell periphery on extravisceral
D i a g n o s i s : Large, thick-shelled and moderate conca- area. It consists of narrow and high costellae, in interspaces
vo-convex Montsenetes with transverse shell outline, weak with small and high endospines.
and rounded ventral myophragm, weak and short dorsal Ornament of the ventral valve consists of numerous fine,
median septum, short anderidia and evenly and coarsely rounded costellae of uniform size, numbering 4, rarely 5,
costellate shell. per 2 mm in anterior margin. Interspaces of similar size are
D e s c r i p t i o n : Shell large, thin-shelled relatively to deep. In ventral valve, the new costellae originate mostly by
shell size, moderately concavo-convex, transversely oval, in bifurcation, while intercalation is rare and restricted to pos-
maximum 31 mm wide. Spines were not found in our mate- terolateral sectors. In the dorsal valve, the intercalation pre-
rial. The inner apertures of spines are arranged symmetri- vails. The large shells bear 90–100 distinct costellae.
cally along the posterior margin, two on each side of the Concentric ornament is weak, with several concentric
large valves. growth lines.
The dorsal valve is moderately and evenly concave, R e m a r k s : New species is referred to Montsenetes, but
widest at posterior third, with rectangular to obtuse cardinal its external morphology differs significantly from other
extremities. The DvW/DvL is 1.33 (range: 1.31 to 1.69; n = species of the genus. It is much larger and less convex than
11). The dorsal interarea is low, hypercline. The notothyri- M. notius (BENEDETTO, 1984) and M. pervulgatus sp. n.
72
The anterior edge of the visceral area is much better im- RACHEBOEUF, 1992. However, this determination is ten-
pressed in the new species than in other species of the genus. tative and must be confirmed by a new better material col-
D i s t r i b u t i o n : Upper Emsian. lected in the area. The genus Pleurochonetes is known from
O c c u r r e n c e : Tamesna Basin, Oued Felaou area the Emsian (lower Icla Formation) to the Eifelian (upper
(sample H 328), In Rahir Tin Amzi section (sample H Icla Formation) of Bolivia (Isaacson 1977, Le Hérisse et al.
1057), Tin Rerhoh area, section 5 (sample H 1088), In Ateï 1992) and from the Lower Devonian of the Parecis Basin of
section (sample H 1003). Brazil (Racheboeuf in Boucot et al. 2001).
D i s t r i b u t i o n : Emsian ? or Eifelian.
Montsenetes sp. O c c u r r e n c e : Tamesna Basin, Felaou Area (sample H
Pl. 2, fig. 15 1067).
M a t e r i a l : One dorsal and three ventral and valves
preserved as internal moulds in sandstone. Order Orthotetida WAAGEN, 1884
R e m a r k s : The valves represent another species of Suborder Orthotetidina WAAGEN, 1884
Montsenetes RACHEBOEUF, 1992. It differs from M. per-
Superfamily Chilidiopsoidea BOUCOT, 1959
vulgatus sp. n. in having a deeper dorsal valve, a weak ven-
tral myophragm and a finer ornament. The ornament in M. Family Chilidiopsidae BOUCOT, 1959
pervulgatus is coarser, with 4 costellae per 2 mm anterome- Subfamily Chilidiopsinae BOUCOT, 1959
dianly, while in M. sp. there are 5 to 6 costellae.
Montsenetes sp. is even smaller, with highly convex ventral Genus Iridistrophia HAVLÍČEK, 1965
valve already in 12 mm wide specimens but in this size, the
ventral valves of M. pervulgatus are weakly to moderately Ty p e s p e c i e s : Orthis umbella BARRANDE, 1848;
convex only. Montsenetes sp. is surely a new species but due Lower Devonian, Lochkovian; Bohemia.
to poor material it is left in open nomenclature. It is note-
worthy that the shells are associated with several taxa Iridistrophia sp.
(Craniops, Salopina, Plicanoplia) unknown from other lev- Pl. 5, fig. 15
els and collecting sites in the area. M a t e r i a l : Three external moulds, several fragments.
D i s t r i b u t i o n : Upper Emsian. Description: All specimens are represented by small
O c c u r r e n c e : Tamesna Basin, Tin Rerhoh area, sec- shells, the most complete fragment is 15 mm wide but frag-
tion 5 (sample H 1087). ments indicate a larger shell size, to some 22–25 mm. The
ventral valve is weakly convex, the dorsal valve is flat with
Subfamily Notiochonetinae RACHEBOEUF, 1992 weak and broad sulcus. Cardinal extremities are almost
rectangular. Interiors are unknown. The shell exterior bears
Genus Pleurochonetes ISAACSON, 1977 fine, acute and high costellae, intercalated by slightly finer
Ty p e s p e c i e s : Chonetes (Pleurochonetes) lauriata secondary costellae. There are some 4–6 costellae per 2 mm
ISAACSON, 1977; Lower Devonian, Emsian; Bolivia. anteromedianly.
R e m a r k s : The upper limit of Iridistrophia
HAVLÍČEK, 1965 is the Emsian after Williams et Brunton
Pleurochonetes ? sp.
(2000). It is in coincidence with the suggested Upper Em-
Pl. 5, fig. 20
sian age of our samples. However, in Libya the genus is re-
M a t e r i a l : Two internal moulds of ventral valves in ported from the Bir al Quasr Formation (Eifelian) of the
sandstone. Murzuq Basin (Havlíček et Röhlich 1987), but this determi-
D e s c r i p t i o n s : The valves are large (30 mm width in nation is based on poor specimens and could not be exclud-
the largerst valve), weakly convex, with rectangular cardinal ed that this species might be referred to another chilidiopsid
extremities. The ventral interarea is very low, apsacline. The genus. Our specimens, although fragmentary preserved, are
delthyrium is small, with a small pseudodeltidium. The not resupinate and the dorsal valve is flat. Both fragments
large ventral muscle field is weakly impressed, divided by are near to I. dendritica BENEDETTO, 1984 from the Up-
low and long myophragm. The visceral area is distinctly de- per Emsian of Venezuela, from which our valves differ by
fined. Inner costellation is distinct along the shell periphery, smaller size. Out of several species of the genus in the Pra-
with 4 costellae per 2 mm anteriorly. External ornamenta- gian to Emsian strata of Argentina (Herrera et al. 1998) the
tion, spines and the dorsal valve are unknown. species I. aliciae HERRERA et al., 1998 is near to our spec-
Remarks: The valves differ by a large size, weak con- imens by size, ornament and shell outline.
vexity and a very low, apsacline interarea from other chone- D i s t r i b u t i o n : Upper Emsian.
toids in the Tamesna Basin. The valve morphology is O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec-
consistent with assignment to Notiochonetinae tion (sample H 1058), Tin Rerhoh area, section 5 (sample H
RACHEBOEUF, 1992, especially to genus Pleurochonetes 1087).
73
Class Rhynchonellata WILLIAMS et al., 1996 D e s c r i p t i o n : The ventral valve is subcircular, 95 %
Order Orthida SCHUCHERT et COOPER, 1932 as long as wide, thin-shelled, one complete valve is 6 mm
wide. The valve is strongly convex transversely and axially,
R e m a r k s : Orthid brachiopods, apart of the enigmatic with maximum depth in the posterior third. The maximum
genus Tropidoleptus, are rare in samples in the Tamesna width is at midlength. The ventral interarea is apsacline,
Basin. Only a few specimens, often incomplete and small small and low, about less than 70 % of valve width. Dental
sized, have been found. However, they have certain strati- plates are thin, of moderate length, diverging anteriorly at
graphical importance and therefore even this poor material about 60° angle. The ventral muscle field is weakly im-
is described and figured. pressed, as long as the dental plates. Internal ribbing is of
uneven size, with deep interspaces restricted to the shell pe-
Suborder Dalmanellidina MOORE, 1952 riphery. Exterior bears fine radial costellae of uneven size.
Superfamily Dalmanelloidea SCHUCHERT, 1913 R e m a r k s : The species is poorly known due to rare
Family Rhipidomellidae SCHUCHERT, 1913 material. It is most similar to Salopina hitchcocki
WALMSLEY et al., 1969 from the Tarratine Formation
Subfamily Rhipidomellinae SCHUCHERT, 1913
(Siegenian) of Maine (Walmsley et al. 1969, Boucot 1973).
Our specimens differ from S. hitchcocki by longer and thin-
Genus Rhipidomella OEHLERT, 1890
ner dental plates but other features are almost identical. The
Ty p e s p e c i e s : Terebratula michelini LÉVEILLÉ, Eifelian species S. hazardensis WALMSLEY et al., 1969
1835; Lower Carboniferous; Belgium. from Pennsylvania is smaller, transverse and its dental
lamellae are short and medianly curved.
Rhipidomella ? sp. D i s t r i b u t i o n : Upper Emsian.
Pl. 5, figs 3, 4 O c c u r r e n c e : Tamesna Basin, Tin Rerhoh area, sec-
tion 5 (sample H 1087).
M a t e r i a l : One internal mould of dorsal valve and incom-
plete external mould of ventral valve in calcareous sandstone.
R e m a r k s : Poor material makes the generic assign- Family Tropidoleptidae SCHUCHERT, 1896
ment of both valves speculative. The dorsal valve (pl. 5, fig.
3) is strongly convex, of circular outline indicating only a Genus Tropidoleptus HALL, 1857
small interarea. Internally, there is a prominent median Ty p e s p e c i e s : Strophomena carinatus CONRAD,
ridge extending anterior to midlength. The genus 1839; Middle Devonian, Hamilton Group; New York, USA.
Rhipidomella OEHLERT, 1890 extends from the Eifelian to R e m a r k s : As stated by Isaacson et Perry (1977), the
Upper Permian (Williams et Harper 2000) and our find genus Tropidoleptus HALL, 1857 displays remarkable mor-
would be one of the earlier reports of the genus in the area. phologic uniformity throughout its history from the Siege-
Le Maïtre (1952b) reported two species of the genus from nian to Frasnian. Apart from the type species, only a few
the Lower Eifelian of Erg Djemel in Algeria but she did not other species or subspecies have been discriminated (Isaac-
figure these specimens. The genus is common from the son et Perry 1977, Cooper et Dutro 1982, Boucot et al.
Middle Devonian to Tournaisian in Libya and Algeria 1983,). The genus has an extensive geographical distribu-
(Borghi 1939, Havlíček 1984, Mergl et Massa 1992, Brice tion, covering South America, Mexico, North America,
et Latréche 1998). Unlike our specimen, the cosmopolitan North and South Africa and Central and Southwest Europe;
genus Dalejina HAVLÍČEK, 1953, which is characteristic its distribution and evolutionary history have been com-
for Lower Devonian, lacks the prominent median septum. mented by many authors. However, variations of the shell
D i s t r i b u t i o n : Upper Emsian. morphology within particular populations (Williams 1913)
O c c u r r e n c e : Tamesna Basin, Tin Rerhoh area, sec- and only minor morphological changes through its phy-
tion 5 (sample H 1086). logeny make it useless as an index fossil (Isaacson et Perry
1977, Boucot et al. 1983).
Superfamily Enteletoidea WAAGEN, 1884 In the territory of Northwest Africa and Central Sahara
Family Draboviidae HAVLÍČEK, 1950 the genus was noted in many localities (Mauritania: Le
Subfamily Draboviinae HAVLÍČEK, 1950 Maïtre1952a, Sougy 1964, Bitam et al. 1996, Racheboeuf et
al. 2004; Algeria: Brice et Latréche 1998; Libya: Borghi
Genus Salopina BOUCOT, 1960 1939, Boucot et al. 1983, Havlíček 1984, Havlíček et Röh-
lich 1987, Mergl et Massa 1992; Niger: Boucot et al. 1983,
Ty p e s p e c i e s : Orthis lunata J. de C. SOWERBY,
Mergl et al. 2001).
1839; Silurian, Ludlow; England.
Numerous surface samples from the Tamesna Basin
yielded Tropidoleptus but the shells are often scrapped and
Salopina sp.
poorly preserved. Three morphological groups can be dif-
Pl. 5, fig. 2
ferentiated among our specimens. They probably represent
M a t e r i a l : Two internal moulds of ventral valve and different subspecies or species separated stratigraphically;
incomplete external mould. one of them is described as a new subspecies. The oldest
74
shells of this genus in the Tamesna Basin have been collect- 1006, H 1009), Anou Izileg area (H 323), and In Debirene
ed from beds with fauna referred to the Upper Pragian or section (samples H 1105, H 1106, H 1107).
Pragian/Emsian boundary interval. However, the adult
shells of this population are poorly known. Younger sand- Tropidoleptus carinatus titanius subsp. n.
stone beds, referred to the Upper Emsian, yielded remark- Pl. 6, figs 1–11
ably large specimens and this giant Tropidoleptus is
described as a separate subspecies. Younger representatives H o l o t y p e : Complete shell (MaMe 0038), figured on
of the genus have been collected near the Emsian/Eifelian pl. 6, figs 1, 3, housed in the Department of Palaeontology
boundary and these, as well as the shells in younger succes- of the Museum of West Bohemia, Plzeň, Czech Republic.
sion, are of average size, with width rarely exceeding 20 Paratype: Dorsal valve (MaMe 0037), internal mould
mm. These shells are referred to the typical subspecies T. figured on pl. 6, fig. 11, housed in the Department of
carinatus carinatus (CONRAD, 1839). No shells in our ma- Palaeontology of the Museum of West Bohemia, Plzeň,
terial display morphology consistent with the formerly de- Czech Republic.
scribed African subspecies T. carinatus nigerensis Ty p e h o r i z o n : Lower Devonian, Upper Emsian.
BOUCOT et al. 1983 and T. carinatus freuloni BOUCOT et Ty p e l o c a l i t y : Algeria, Tamesna Basin, In Ateï sec-
al. 1983 but shells from higher levels, probably of a Give- tion, sample H 1003.
tian age, are very poor. Absence of these two subspecies E t y m o l o g y : titanius (Lat.) – referring to the giant
among the available material is indicative of an earlier, pre- size of the shell.
Givetian age of most of the collected specimens from the M a t e r i a l : One complete shell, five dorsal valves,
Tamesna Basin. three ventral valves and numerous fragments preserved in
Although recently this problematic genus is not referred sandstone with carbonatic cement.
to orthids (Williams et Harper 2000), present authors retain D i a g n o s i s : Large, depressed, non-carinate Tropi-
its original position in the Superfamily Enteletoidea doleptus carinatus with low, short and thin dorsal median
WAAGEN, 1884 by Williams et Wright (1965). septum and low rounded costae.
D e s c r i p t i o n : The shell is very low, planoconvex,
weakly unisulcate, rather thick-walled, with cardinal ex-
Tropidoleptus carinatus carinatus (CONRAD,
tremity obtuse and never mucronate. The size is unusual and
1839) very large for the genus, with the width exceeding 50 mm
Pl. 5, figs 16, 17, 19, 22
(the largest specimen examined attains 56 mm width). The
1986 Tropidoleptus carinatus (Conrad, 1839); Drot, p. 513, pl. 1, shell wall is densely exopunctate.
figs 10, 11. The dorsal valve is transversely oval, with hinge line
M a t e r i a l : Some twenty-five valves in various, gener- 65–70 % of the maximum width. The lateral margins are
ally poor state of preservation in sandstones. evenly rounded, the anterior margin is only weakly curved.
R e m a r k s : Specimens from various stratigraphical lev- The valve is almost flat, with a broad and shallow sulcus ap-
els may be referred to typical subspecies T. carinatus cari- parent from valve midlength. The interarea is low, anacline,
natus (CONRAD, 1839). The size of the specimens does with the notothyrium closed by a high convex transversely
not exceed 25 mm and falls into usual size range of the type striated chilidium. The dental sockets are deeply excavated
subspecies. into shell wall, widely diverging and shoaling anteriorly.
The specimens of the Upper Pragian (sample H 1052) Their bottom is coarsely crenulated, with 5–6 transverse
age have fine dorsal median septum perfectly developed al- ridges. Outer socket ridges are thin, inner socket ridges are
ready in very small individuals (pl. 5, fig. 19). Inner periph- high, massive, united with moderately divergent short crural
ery of these small shells bears bundles of delicate radial bases. The notothyrial platform is long and narrow, anteri-
ridges. Adult specimens are some 20 mm wide and have orly supported by a low and broad proximal part of the me-
weakly convex ventral valve devoid of distinct carination. dian septum. The septum anteriorly extends into tapering,
Specimens from stratigraphically higher beds, referred low and acute myophragm, which continues towards 60–70
to the Upper Emsian, Emsian/Eifelian boundary interval % of the valve length. The cardinal process is undivided,
and Eifelian have commonly smaller shells, with width posteroventrally faced, with ventral face divided by deep
about 15 mm. The ventral valve is considerably convex and transverse slit into shorter posterior and longer anterior
in some specimens also distinctly carinate. The stratigraph- lobes. The bottom of slit has subaxially two deeper pits.
ically youngest shells in the Tamesna Basin come from Muscle scars are weakly impressed, with apparent posterior
samples H 826 and H 1009, referred to the Givetian. adductors situated in front of the notothyrial platform. An-
D i s t r i b u t i o n : Upper Pragian, Emsian, Eifelian, and terior adductors are large, triangular, situated at the centre of
Givetian. the valve.
O c c u r r e n c e : Tamesna Basin, Oued Felaou area The ventral valve is gently and evenly convex in a trans-
(samples H 333, H 1405), Kori-Idemeg – Touaret section verse profile and weakly and unevenly convex in a lateral
(sample H 884), In Rahir Tin Amzi section (samples H profile, with the maximum convexity and depth in the pos-
1052, H 1057, 1058), Tin Rerhoh area, section 5 (samples H terior third. The low fold is apparent from shell midlength.
1086, H 1087, H 1088), In Ateï section (samples H 1004, H The ventral beak is short, obtuse. The interarea is low, al-
75
most orthocline, with fine transverse striation. The broadly son et Perry 1977: pl. 1, figs 17, 23; pl. 3, figs 13, 26; pl. 4,
triangular delthyrium is covered apically by long convex fig. 20) are weaker in T. carinatus titanius. The currently
delthydial plate. The valve beak is short, curved. Teeth are described Tropidoleptus sp. from Hodh area of Mauritania
large, faced dorsolaterally, supported by low, broadly diver- (Racheboeuf et al. 2004) is smaller and belongs to another
gent dental plates, which laterally bound an extensive ven- subspecies.
tral muscle field. The muscle field is weakly impressed, D i s t r i b u t i o n : Upper Emsian.
large, wider than long, with transversely striated surface. O c c u r r e n c e : Tamesna Basin, In Ateï section (sample
The field is axially divided by a weak myophragm. H 1003), Oued Felaou area (sample H 328), Tin Rerhoh
The ornament of large shells consists of low and round- area, section 5 (sample H 1087).
ed costae of uniform size, slightly decreasing in size late-
rally, slowly widening during growth and separated by Order Rhynchonellida KUHN, 1949
shallow rounded interspaces of similar size. However, the
smaller specimens to 10 mm length have high, distinctly Superfamily Rhynchotrematoidea SCHUCHERT,
rounded costae in median and submedian sectors. The me- 1913
dian costa on the ventral valve is weakly coarser and sepa- Family Trigonirhynchiidae SCHMIDT, 1965
rated by narrower interspaces from lateral costae, which Subfamily Trigonirhynchiinae SCHMIDT, 1965
decrease in size laterally. Cardinal extremities are almost
smooth. The shells have 23–25 costae. Internal costation is Genus Trigonirhynchia COOPER, 1942
weak and obscured by secondary deposits in large shells.
Fine internal radial striation along shell periphery, which is Ty p e s p e c i e s : Uncinulina fallaciosa BAYLE, 1878;
common feature in other subspecies of the genus, is weak in Lower Devonian; France.
T. carinatus titanius. Fine pustules cover flattened peripher-
al rim of large shells, especially in the bottom of inter- Trigonirhynchia sp.
spaces. Concentric ornament is weak, restricted to several Pl. 5, fig. 12
uneven short growth lamellae or coarser growth lines near
M a t e r i a l : Three incomplete ventral valves.
periphery of large shells.
D e s c r i p t i o n : Assumed from the fragments, the shell
R e m a r k s : Tropidoleptus carinatus titanius subsp. n.
is some 12–14 mm wide, coarsely costate with short and
differs from other subspecies and species of the genus by
stout dental sockets and short and weakly impressed ventral
the large size. While other subspecies or species (T. cari-
muscle field. The costae are rounded, originating at the
natus freuloni BOUCOT et al., 1983, T. carinatus nigeren-
beak. Ventral sulcus is distinct from midlength, extending
sis BOUCOT et al., 1983, T. carinatus rhenana FRECH,
into long serrate tongue. Ventral shell has six rounded
1897, T. platys COOPER et DUTRO, 1982) are 20 to 30
costae in shallow sulcus and five lateral costae in each flank.
mm wide in average, the shells of T. carinatus titanius are
Concentric ornament consists of fine growth lines.
50 mm wide or even larger as can be assumed from the as-
R e m a r k s : Poor material does not allow accurate
sociated shell fragments. There are limited exact data about
generic determination but nature of the costae and the out-
the size of other subspecies. The maximum width noted by
line of the tongue indicate affinity to Trigonirhynchia
Isaacson et Perry (1977) are less than 35 mm, in paper
COOPER, 1942. This genus occurs in the Pragian and Em-
Isaacson et Chlupáč (1984) even less than 27 mm. All de-
sian (Brice et al. 2000). The species seems to be rather sim-
scribed specimens from Bolivia, where the species is typi-
ilar to the type species T. fallaciosa (BAYLE, 1878), but its
cal element in the Emsian brachiopod fauna, are less than
size and ornament are similar also to T. occidens
27 mm wide (Isaacson 1977a). Also specimens of T. cari-
(WALCOTT, 1884) from the Emsian (Eurekaspirifer piny-
natus from the Hamilton Group (Givetian) of New York,
onensis Zone) of Nevada (Johnson 1970).
USA are smaller (Williams 1913). In addition, a compara-
D i s t r i b u t i o n : Upper Pragian.
tively small and low median ridge of T. carinatus titanius
O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec-
distinguishes it from other subspecies of the genus. Only T.
tion (sample H 1052).
carinatus freuloni almost lacks the median septum but
there are other differences between these subspecies, main-
ly in stronger myophragm, deeper impression of adductors Genus Stenorhynchia BRICE, 1981
and prominent internal radial striation of T. carinatus Ty p e s p e c i e s : Terebratula nympha BARRANDE,
freuloni. Specimens from the Hamilton Group of New York 1847; Lower Devonian, Pragian; Barrandian, Bohemia.
(Williams 1913) and the Silica Shale of Ohio have higher
and more prominent dorsal median septum, T. carinatus
rhenana from the Lower Emsian of Germany differs by
Stenorhynchia ? sp.
Pl. 5, figs 5–7, 9
coarser costation and distinctly bilobed cardinal process.
The coarser median costae on shell exterior that is devel- M a t e r i a l : Three incomplete external moulds and one
oped in specimens from various worldwide sites (cf. Isaac- internal mould of dorsal valves, one ventral valve.
76
D e s c r i p t i o n : The available fragments of a small Order Athyridida BOUCOT, JOHNSON et
rhynchonellid have low dorsal sulcus distinct from the STATON, 1964
umbo. Costae are prominent from the beak, simple, an- Suborder Athyrididina BOUCOT, JOHNSON et
gular, separated by deep angular interspaces. Flanks bear STATON, 1964
slightly curved asymmetrical costae of comparable size.
The dorsal valve interior has distinct strong median sep- Superfamily Athyridoidea DAVIDSON, 1881
tum. The ventral valve is subpentagonal, with the erected Family Athyrididae DAVIDSON, 1881
beak and shallow sulcus. The shell has some 16 costae, Subfamily Athyridinae DAVIDSON, 1881
out of which four are in the dorsal fold.
R e m a r k s : It is impossible to make an accurate de- Genus Athyris M’COY, 1888
termination of these fragments. However, they indicate
the presence of a small rhynchotrematoid in benthic com- T y p e s p e c i e s : Terebratula concentrica VON
munities. The shape of our shells is most similar to small BUCH, 1834; Middle Devonian, Eifelian; Germany.
specimens of Stenorhynchia nympha (BARRANDE,
1847) reported from many sites in Europe and North Athyris sp.
Africa (Havlíček 1961, Brice 1981). Small specimen of Pl. 5, figs 18, 21
S. nympha from the Upper Emsian of Morocco figured by
M a t e r i a l : Two internal moulds and one external
Drot (1964; pl. 19, fig. 6) has less curved lateral costae.
mould of ventral valves.
The fragments also show similarity to Cupularostrum sp.
D e s c r i p t i o n : The ventral valve has rounded subpen-
reported by Boucot et Johnson (1968) from the Emsian
tagonal outline and is slightly wider than long (VvL/VvW =
(Bois Blanc Formation) of New York, USA.
0.90; n = 2) with maximum width at midlength. The larger
D i s t r i b u t i o n : Upper Emsian.
shell is 26 mm wide. Posterolateral margins are almost
O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi
straight, lateral margins are strongly curved. Anterolateral
section (samples H 1057, H 1058), In Ateï section (sam-
margin is weakly curved. The anterior margin is truncated.
ple H 1003), In Debirene section (sample H 1108).
The valve is evenly convex in lateral and transverse profiles.
The sulcus is narrow, shallow, distinct from about mi-
Superfamily Camarotoechioidea SCHUCHERT, dlength. It extends into long, rounded tongue, occupying
1929 some 50 % of the shell width.
Family Leiorhynchidae STAINBROOK, 1945 Dental plates are distinct, short, subparallel, with the
Subfamily Leiorhynchinae STAINBROOK, 1945 bases slightly converging anteriorly and ventrally. The ven-
tral muscle field is clearly defined, with long, narrowly tri-
angular, subparallel diductor scars, flabellate along the
Genus Leiorhynchus HALL, 1860
anterior margins. Adductor scars are narrow, parallel, encir-
Ty p e s p e c i e s : Orthis quadracostata VANUXEM, cled by diductors. The muscle field is long, extending over
1842; Middle Devonian, Givetian; New York, USA. the shell midlength.
Shell exterior bears thin concentric lines of uniform size,
of which a few extend into short lamellae. There are some
Leiorhynchus ? sp.
10–12 lines per 5 mm anteromedianly.
Pl. 5, fig. 13
C o m p a r i s o n : Athyridids are rare in our material and
M a t e r i a l : One poorly preserved external mould in apart from the figured specimens, only a small fragment of
siltstone. ventral valve has been found in another sample (H 1087).
R e m a r k s : Although poorly preserved, elongate Outline, convexity and external ornament of the shells from
rounded outline, weak radial plications in a midsector of the Tamesna Basin are consistent with the attribution to
the ventral valve and smooth flanks indicate leiorhynchid Athyris M’COY, 1888. Shells are similar to the type species
affinity of the shell. It is the only report of the camaro- A. concentrica VON BUCH, 1834 as newly defined by Al-
toechioids brachiopod in the material from the Devonian varez et al. (1996) but our species differs by transversely
outcrops of the Tamesna Basin, unlike other North strongly convex ventral valve and bigger shell. The Spanish
African basins with the Middle and Upper Devonian suc- Emsian species Athyris campomanesi (DE VERNEUIL et
cession. However, a subsurface data indicate the pres- D’ARCHIAC, 1845) is less transverse, with shorter ventral
ence of other leiorhynchids in the Tamesna Basin tongue and has distinct radial plications that are absent in A.
(Part 2). sp. The species Athyris cf. cora HALL, 1857 from the Eife-
D i s t r i b u t i o n : The Givetian or even younger age of lian of Venezuela (Benedetto 1984) differs by much smaller
the sample is suggested. size, weaker concentric ornament and weak ventral tongue.
O c c u r r e n c e : Tamesna Basin; Kori-Idemeg – Touaret Le Maïtre (1952b) reported some athyridids from western
area (sample H 888). Algeria, among others also A. concentrica. The figured shell
77
(Le Maïtre 1952b; pl. 19, figs 41, 42) differs from our spec- M a t e r i a l : Six dorsal valves, eight ventral valves, nu-
imens by smaller size and more distinct sulcus. In Central merous valves of juvenile specimens and several fragments
Sahara, poorly preserved athyridids were also reported by with microornament preserved in fine sandstone.
Havlíček et Röhlich (1987) and Mergl et Massa (1992) from D i a g n o s i s : Pustulatia with broad dorsal sulcus and
the Eifelian of Libya (Ghadamis and Murzuq basins). These weak ventral median plications, with cardinal process with
Libyan athyridids are much transverse with a deeper ventral small ctenophoridium of about seven plates; lateral plica-
sulcus and their morphology is more consistent with a sub- tions weak and rapidly decreasing in size, distinct from mi-
genus Alvarezites STRUVE, 1992. dlength; ventral myophragm absent.
D i s t r i b u t i o n : Upper Emsian to Lower Eifelian. D e s c r i p t i o n : The shell is small, planoconvex, weak-
O c c u r r e n c e : Tamesna Basin, Tin Rerhoh area, section ly sulcate, thin-shelled, transversely oval, with width maxi-
5 (sample H 1087), In Debirene section (sample H 1106). mum at or posterior to midlength. The shell width attains
some 9.0 mm in large specimens. Lateral margins are even-
ly curved, frontal margin has a weak emargination.
Order Spiriferida WAAGEN, 1883
The dorsal valve is flat, with rounded cardinal extremi-
Suborder Spiriferidina WAAGEN, 1883 ties. DvW/DvL ratio is about 135 %. The dorsal interarea is
Superfamily Ambocoelioidea GEORGE, 1931 anacline, low but clearly defined. The shallow and broad
Family Ambocoelidae GEORGE, 1931 sulcus is distinct from valve midlength. The dental sockets
are deep, triangular, supported by triangular, moderately di-
vergent (at about 65°) inner socket plates. The crural bases
Genus Pustulatia COOPER, 1956 are poorly defined, attached to surface of weakly defined
Ty p e s p e c i e s : Vitulina pustulosa Hall, 1860; Middle triangular notothyrial platform. The platform is low, broad
Devonian, Hamilton Group; New York, USA. and long, unsupported by a median ridge. The ctenophorid-
R e m a r k s : The genus has typical pustulose microorna- ium is small, faced posteroventrally, of seven vertical radi-
ment, which is lacking in the externally similar Lower ally arranged plates.
Devonian genus Plicoplasia BOUCOT, 1959. Other differ- The ventral valve is high, with broad and shallow sulcus.
ences in these homoeomorphous genera concern of the The plications bounding the sulcus are much more promi-
myophragm and the median septum, which are poor in Pus- nent than the lateral plications. The delthyrium is large,
tulatia COOPER, 1956. The presence of Pustulatia in the open. The bottom of delthyrial chamber is highly raised
Upper Pragian of the Tamesna Basin indicates that the ori- above adjacent valve floor. Teeth are large and robust in re-
gin of the genus could be sought outside of North America. lation to shell size, unsupported by dental plates. The ven-
The genus has more likely a Gondwanan origin. It is known tral muscle field is poorly impressed, with large paired
from the Emsian and early Eifelian, possibly also the Give- diductors and much smaller, posteromedianly situated scars
tian from Niger, South Africa, Bolivia, and Brazil. Pustula- of adductors. The myophragm is absent.
tia is also reported from Argentina Precordillera from beds The ornament consists of low rounded plications, four in
already of the Emsian age (Herrera 1995). The famous oc- each flank, rapidly diminishing in size laterally. The lateral
currence on Pustulatia pustulosa (HALL, 1860) in the plications become distinct from shell midlength, being ob-
Hamilton Group (New York, USA) represents the latest scure near shell beaks. The bottom of ventral sulcus bears
member in the evolutionary history of the genus. The oc- weaker but distinct median plication. The dorsal sulcus has
currence of P. pustulosa in the Hamilton Group of New York correspondingly sized median interspace between the pair
associates with the elements having also other than North of plications. Microornament consists of almost regularly
American origin (e.g. Tropidoleptus). spaced radial fila with regular rows of fine elongate pus-
tules. There are seven to ten fila on plications bounding the
sulcus. Concentric ornament consists of a few, low, distinct
Pustulatia lessardi sp. n. growth lamellae. Growth fila are unevenly sized, distinct
Pl. 7, figs 1–12, 15
between lamellae along the shell margin of large shells.
H o l o t y p e : Dorsal valve (MaMe 0052B), figured on R e m a r k s : This new species is the stratigraphically
pl. 7, fig. 5, housed in the Department of Palaeontology of earliest species of the genus in our samples from the Tames-
the Museum of West Bohemia, Plzeň, Czech Republic. na Basin, and probably also the earliest known representa-
P a r a t y p e : Ventral valve (MaMe 0050H), figured on tive of the genus yet recorded. The species differs from the
pl. 7, fig. 11, housed in the Department of Palaeontology of stratigraphically subsequent species Pustulatia tamesnaen-
the Museum of West Bohemia, Plzeň, Czech Republic. sis sp. n. in the same area by a broader sulcus having a low
Ty p e h o r i z o n : Lower Devonian, Upper Pragian. median plication in the ventral valve, by the lateral plica-
Ty p e l o c a l i t y : Algeria, Tamesna Basin, In Rahir tions distinct from the shell midlength, and by rapidly re-
section, sample H 1052. duced size of lateral plications. The dorsal valve of P.
E t y m o l o g y : After Dr. Lionel Lessard, the famous lessardi sp. n. is flat, while that of P. tamesnaensis sp. n. is
French geologist. moderately convex. The new species differs from the speci-
78
men found in the Taoudeni Basin by Villemur et Drot from the umbones.
(1957) and referred to P. pustulosa (HALL, 1860) by a dis- D e s c r i p t i o n : The shell is ventribiconvex, weakly
tinctly broader dorsal sulcus and uneven size of plications. unisulcate, thin-shelled, transversely oval, 9.0 mm wide in
The presence of P. pustulosa in the Tamesna Basin was the largest known specimen.
commented by Le Maïtre (1961). She noted the occurrence The dorsal valve is moderately convex, widest in the
of P. pustulosa in three different levels, from Siegenian to posterior third, with a shallow, rather narrow sulcus. The
upper Emsian, but without any description or illustrations of dorsal interarea is very low, anacline. Cardinal extremities
these specimens. It is probable that her determination was are rounded. Dental sockets are long, narrow, deep and
approximate and comprised two currently described broadly divergent. The dental sockets are bound by broadly
species. The typical Pustulatia pustulosa (HALL, 1860) divergent (at about 130°) inner socket ridges. The cardinal
from the Middle Devonian (Hamilton Group; upper Eifelian process is raised, without distinct ctenophoridial blades.
and Givetian) of New York, USA differs by even narrower The notothyrial platform is distinctly raised, short, support-
dorsal sulcus, distinct dorsal myophragm and median ridge, ed by a short median ridge.
and coarser and more numerous lateral plications. The ventral valve is highly convex in lateral view, sub-
The genus Pustulatia is reported, apart from the classi- carinate in transverse profile. The ventral sulcus is shallow
cal Middle Devonian occurrence in New York of USA, from and narrow. The delthyrium is open. Teeth are large, robust,
Brazil (Rathbun 1874), Bolivia and Argentina (Thomas unsupported by dental plates. The ventral interarea is small,
1905, Isaacson 1977b, 1993), and South Africa (Reed catacline. Muscle scars are not impressed in the available
1903). Two species, the stratigraphically earlier (Emsian shells.
and early to mid-Eifelian) species P. curupira (RATHBUN, External ornament consists of distinct plications origi-
1874) and the latter (late mid-Eifelian) species P. branisi nating at umbones. Sulcus-bounding plications are stronger
ISAACSON, 1977, are known from Bolivia. However, their than the lateral plications. There are five plications on each
stratigraphical range may be younger, as suggested by flank of the ventral valve and four plications on each flank
Racheboeuf et al. (1993a, 1993b). The earlier P. curupira of the dorsal valve. The median plication in dorsal sulcus is
differs from the new species by more transverse outline, al- well developed, originating at beak and is slightly smaller
most rectangular cardinal extremities, narrower sulcus and than sulcus-bounding plications. Concentric ornament is not
coarser plications. An important difference concerns the preserved. Microornament of radial rows of elongate pus-
cardinal process. While in P. curupira and P. branisi the car- tules is poorly preserved.
dinal processes are large, bilobate proximally and quadrilo- R e m a r k s : The species differs from P. lessardi sp. n.
bate distally, in the new species the cardinal process is low, by a narrower dorsal sulcus and more prominent lateral pli-
with a chevron-like arrangements of high ctenophoridial cations distinct already from umbones, by shorter and more
plates. divergent inner socket ridges and by convex dorsal valve,
D i s t r i b u t i o n : Upper Pragian. which is flat in P. lessardi. The species P. pustulosa (HALL,
O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec- 1860) has the sulcus-bounding plications only weakly
tion (sample H 1052), Tin Rerhoh area, section 5 (sample H coarser than other lateral plications and the median plica-
1085). tion in sulcus evenly widens anteriorly. In the new species,
the sulcus bounding plications are much stronger than oth-
er lateral plications, leaving the median plication nearly
Pustulatia tamesnaensis sp. n.
evenly wide from the shell midlength. The new species has
Pl. 7, figs 13, 14, 16–22
also more rounded outline and less distinct dorsal
H o l o t y p e : Dorsal valve (MaMe 0128), figured on pl. myophragm than P. pustulosa. The species is similar to P.
7, figs 13, 16, 17 housed in the Department of Palaeontology curupira (RATHBUN, 1874) but the South American
of the Museum of West Bohemia, Plzeň, Czech Republic. species is more transverse, having narrower and deeper sul-
P a r a t y p e : Ventral valve (MaMe 0127C), figured on pl. cus. The specimen referred to P. pustulosa (HALL, 1860)
21, figs 21, 22 housed in the Department of Palaeontology of by Villemur et Drot (1957) from the Taoudeni Basin cannot
the Museum of West Bohemia, Plzeň, Czech Republic. be referred to P. tamesnaensis. The specimens from the
Ty p e h o r i z o n : Middle Devonian, Eifelian. Taoudeni Basin differ by a broader dorsal sulcus with a pair
Ty p e l o c a l i t y : Algeria, Tamesna Basin, In Debirene of weak median plications, while the latter has only a single
section, sample H 1107. undivided median plication. In addition, the new species has
E t y m o l o g y: After the studied area. coarser plications than the specimen figured by Villemur et
M a t e r i a l : Five dorsal valves, five ventral valves and Drot (1957).
several fragments preserved as internal and external moulds D i s t r i b u t i o n : This species is a common element in
in sandstone. samples referred to the Eifelian, associated among others
D i a g n o s i s : Pustulatia with convex dorsal valve, nar- with the spiriferids Paraspirifer, Eleutherokomma and a
row dorsal sulcus without ventral median plication; inner chonetoid Montsenetes pervulgatus sp. n. but it occurs al-
socket ridges broadly divergent; strong plications distinct ready in beds of the suggested Upper Emsian age.
79
O c c u r r e n c e : Tamesna Basin, In Debirene section halved by a short fine myophragm, which anteriorly sepa-
(sample H 1107), Oued Felaou area (sample H 333), Tin rates the parallel elongate adductor scars. Posteriorly, the
Rerhoh area, section 5 (samples H 1087, H 1090). myophragm becomes thickened and passes into massive
vertical septum inside the umbonal chamber. The muscle
scars impressions are distinctly striated (pl. 8, figs 10, 13).
Suborder Delthyridina IVANOVA, 1972
Internal plication is distinct, commonly with 6 or 7 pli-
Superfamily Delthyridoidea PHILLIPS, 1841 cations but only 4 to 5 plications remains distinct in the
Family Hysterolitidae TERMIER et TERMIER, gerontic specimens.
External ornament consists of seven to ten distinct
1949
rounded and undivided lateral plications. Their size rapidly
Subfamily Hysterolitinae TERMIER et TERMIER, decreases laterally. The fold and sulcus are smooth. Fine
1949 concentric ornament is obscure, but several coarse, often
imbricating growth lamellae are anteriorly developed. The
Genus Filispirifer JANSEN, 2001 lamellae are well defined internally by thickening of the
valve. The microornament is unknown.
Ty p e s p e c i e s : Filispirifer merzakhsaiensis Jansen,
R e m a r k s : The morphology and ontogeny of this
2001; Lower Devonian, Pragian; Morocco.
species was widely discussed by Jansen (2001). Although
the microornament of our specimens is unknown, the shell
Filispirifer merzakhsaiensis JANSEN, 2001 morphology is identical with A. merzakhsaiensis JANSEN,
Pl. 8, figs 3–17 2001. In coincidence with specimens from Morocco, our
specimens lack the notothyrial platform, have deeply im-
S y n o n y m y : See Jansen (2001).
pressed ventral muscle field and the exterior with fewer pli-
M a t e r i a l : Seventeen ventral and five dorsal valves
cations but they are slightly smaller than in the specimens
preserved as internal and external moulds in fine sandstone.
described by Jansen (2001). The stratigraphically younger
D e s c r i p t i o n . The shell is biconvex, strongly unipli-
species Filispirifer fallax (GIEBEL, 1858) is larger and has
cate, some 40 mm wide in the largest specimens, with the
distinct notothyrial platform. The species Filispirifer fallax
hinge line extended into short mucronations. The shell wall
is reported from the early Lower Emsian in several sites in
is thickened, especially in the posterior of the ventral valve.
Germany and Spain. The species F. merzakhsaiensis is
The dorsal valve is almost semicircular, 50 % as long as
stratigraphically earlier, restricted to the Middle and Upper
wide, subcarinate, with prominent fold. The flanks are
Siegenian of Morocco. Nearly identical specimens are re-
weakly convex, passing laterally into flattened mucrona-
ported also in various parts of West Europe (Jansen 2001).
tions. Lateral profile of the valve is considerably convex,
D i s t r i b u t i o n : Upper Pragian.
with the maximum depth in midlength. The dorsal interarea
O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec-
is very low, anacline, with widely triangular notothyrium.
tion (sample H 1052), Tin Rerhoh area, section (sample H
The dorsal valve interior lacks the notothyrial platform. The
1080).
ctenophoridium rests at the posterior concavity of the um-
bonal chamber. The ctenophoridium is deeply concave in
transverse profile and consists of 22 to 30 high vertical Genus Hysterolites SCHLOTHEIM, 1820
lamellae. Dental sockets are short and rather small. The cru- Ty p e s p e c i e s : Hysterolites hystericus SCHOLT-
ra are short and massive. Posterior adductor scars are small HEIM, 1820; Lower Devonian, Siegenian, Germany.
and weakly impressed in the posterior fourth of the valve.
Anterior adductors are located at the centre of the valve and
Hysterolites hystericus SCHLOTHEIM, 1820
are deeply impressed, having transversely striated surface.
Pl. 8, figs 1, 2
The scars are separated from each other by a prominent me-
dian septum. 1963 Hysterolites (Hysterolites) hystericus SCHLOTHEIM, 1820;
The ventral valve is transverse, strongly convex, with Solle, p. 174 (here complete older synonymy).
convex flanks and depressed mucronations. The sulcus is 1989 Hysterolites hystericus SCHLOTHEIM, 1820; Gourvennec,
p. 111, pl. 9, figs 11–29 (here synonymy after 1963).
deep, rounded, extending into prominent long tongue. The
ventral interarea is high, apsacline, slightly concave in late- M a t e r i a l : One dorsal valve, one ventral valve and
ral profile. The delthyrium is triangular, bordered by stout several shell fragments in fine sandstone.
bases of large teeth. The ventral valve interior undergoes D e s c r i p t i o n : The shell is strongly transverse, thin-
distinct changes during growth. The distinct, short and shelled, ventri-biconvex, with acute extremities, about 30
broadly divergent dental plates of juvenile specimens are al- mm wide.
most completely sealed in gerontic shells. The ventral mus- The dorsal valve is weakly convex, with anacline inter-
cle field is posteriorly deeply impressed into the valve floor area and wide notothyrium filled by cardinal process. The
but anteriorly remains shallower and is bordered by a low triangular ctenophoridium form some 10 to15 lamellae. The
raised ridge. The surface of the muscle field is posteriorly dental sockets are deep and widely divergent. The crural
80
plates are distinctly developed, the extending crura are ternal moulds of ventral valves, six external moulds and
widely divergent. The muscle scars are indistinct. The me- several fragments.
dian septum is weak. D e s c r i p t i o n : The shell is 35–40 mm wide, slightly
The ventral valve has a deep sulcus. The ventral inter- dorsi-biconvex, with posteriorly strongly thickened shell in
area is steeply apsacline and slightly curved. The ventral adult specimens.
muscle field is weakly impressed, with distinct myophragm, Dorsal valve is moderately convex, with narrow fold
posterolaterally bounded by extrasinal, basally long and slightly broader than bordering plications. Dorsal interior is
evenly thick dental plates. The shell ornament consists of unknown.
high subangular costae, 11–12 on each flank. The ventral valve is broadly semielliptical, 40–50 % as
R e m a r k s : Our specimens are nearly identical by the long as wide but may be significantly longer with age (pl. 9,
size and internal morphology with the specimens from West fig. 5), with the maximum width at the hinge line. The beak
and Central Europe (Solle 1963, Vandercammen 1963, is prominent. Anterior and lateral margins are evenly
Gourvennec 1989) as inferred from our limited material. curved, anterior margin is without prominent emargination.
The genus Hysterolites SCHLOTHEIM, 1820 is widespread Cardinal extremities are acute, sometimes with weak and
in the Pragian and probably extends to the Lower Emsian short mucronations. The valve is moderately convex in late-
(Gourvennec 1989). ral profile and moderately and weakly arched in transverse
D i s t r i b u t i o n : The species is associated with Fil- profile with arched flanks. The sulcus is shallow, only
ispirifer merzakhsaiensis JANSEN, 2001 but it is rare in slightly wider than interspaces between plications, with
both samples. The Upper Pragian age is probable. rounded bottom. The sulcus extends into short rounded
O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec- tongue. The ventral interarea is apsacline, remarkably low,
tion (sample H 1052) and Tin Rerhoh area, section 3 (sam- weakly concave in lateral profile with a wide open delthyri-
ple H 1080). um. Surface of interarea is smooth. The ventral valve interi-
or has large teeth, in small specimens supported by very
Genus Arduspirifer MITTMEYER, 1972 short dental plates, which subtend 80–85° angle between
each other. The adult specimens have the ventral muscle
T y p e s p e c i e s : Spirifer arduennensis SCHNUR, field completely subdued into thick posterior shell wall and
1853; Lower Devonian, Emsian; Germany. the dental plates are obsolete. The ventral muscle field is
R e m a r k s : A revised diagnosis of the genus has been large, broadly oval, halved by a weak and broad median
recently given by Jansen (2001). This author also critically ridge.
revised the taxa referred by various authors to this genus Ornament consists of coarse, low rounded plications.
and presented precise data about the distribution of the Their height rapidly decreases laterally. There are 6 to 7
genus in Germany, Belgium, France, Spain, and Morocco. lateral plications in large ventral valves, but the most lateral
The genus appeared in the Middle Siegenian and extended ones are already poorly defined. Internally, there are 4 to 6
into the Lower Eifelian. In Morocco, the genus is known distinct plications. Interspaces are narrower than plications,
from the Upper Siegenian to early Upper Emsian (Jansen with rounded bottom. Concentric ornament is weak, with
2001). Our limited material from the Tamesna Basin falls several short lamellae along the periphery of large shells.
into variability range of only one species of the suggested Microornament consists of concentric rows of microspines.
Upper Emsian age. There are two concentric rows of microspines per 1 mm,
with 6 to 7 microspines per 1 mm in a row.
Arduspirifer cf. mosellanus (SOLLE, 1953) R e m a r k s : The presence of Arduspirifer MITT-
Pl. 9, figs 1–9 MEYER, 1972 in the area has been reported by Drot (1986).
1952a Acrospirifer speciosus speciosus AUCT.; Le Maïtre, p. 334, She figured four specimens, detail of its microornamenta-
pl. 2, figs 4–9. tion, and suggested the Upper Emsian age. Our specimens
1952b Acrospirifer speciosus speciosus AUCTORUM; Le Maïtre, from the same sample H 1088 and also from the probably
p. 126, pl. 15, figs 16–18. corresponding level (sample H 1058) are similar to the
cf. 1953 Hysterolites (Acrospirifer) mosellanus n. sp.; Solle, p. 83, specimens referred by Solle (1953) to A. mosellanus mosel-
pl. 8, figs 119–123, pls. 9–11, pl. 12, figs 176–178. lanus SOLLE, 1953. Our specimens have less extended
1964 Acrospirifer mosellanus mosellanus SOLLE, 1953; Drot, p. hinge line than the specimens from Germany, especially the
54. subspecies A. mosellanus gracilis (SOLLE, 1953) and A.
1964 Acrospirifer mosellanus cf. mosellanus SOLLE, 1953; Drot,
mosellanus steiningeri (SOLLE, 1953) (= probably subjec-
p. 54.
1964 Acrospirifer mosellanus cf. steiningeri SOLLE, 1953; Drot, tive synonyms of A. mosellanus mosellanus). This species is
p. 54. a characteristic younger member of the evolutionary lineage
1964 Acrospirifer cf. mosellanus steiningeri SOLLE, 1953; Drot, of the genus Arduspirifer. The species is known from the
p. 54. Upper Emsian in Germany (Solle 1953), France (Heddebaut
1986 Arduspirifer sp. gr. mosellanus (SOLLE, 1953); Drot, p. 513, 1981) and Celtiberia (Carls et al. 1972, Carls 1988). It is
pl. 1, figs 1–4, 7. known, apart from the already mentioned paper of Drot
M a t e r i a l : External mould of a complete shell, six in- (1986), probably also in Algeria and Mauritania (Le
81
Maïtre1952a, 1952b; p. 126, Acrospirifer speciosus specio- tinct from the beak. The plications dichotomously divide in-
sus SCHNUR, 1853). Our specimens are externally similar to two slender ribs at about midlength, separated by shallow
also to the species Arduspirifer supraspeciosus (LOTZE, interspaces. Only four to five plications adjacent to fold and
1928) from the Eifelian of Germany and Belgium (Solle sulcus, respectively, bifurcate, the remaining lateral plica-
1953, Vandercammen 1963) but our specimens are less tions remain undivided. Each flat side of the fold and sul-
transverse and lack distinct dental plates present in the Ger- cus, respectively, has weakly defined radial bands separated
man specimens. Our specimens can be compared with oth- by obscure interspaces from the axial part of the fold and
er members of the A. intermedius evolutionary lineage sulcus. There are 15 plications distinct from the beaks, in to-
(Solle 1953). Their shared morphology, with few low tal some 20 along the anterior margin of large shells.
rounded plications and narrow sulcus, indicates a late Em- R e m a r k s : The poor preservation and lack of complete
sian or early Eifelian age of the specimens from Tamesna shells make determination of particular specimens difficult.
Basin. By shell shape, the species shows distinct affinity to P. cul-
A g e : Upper Emsian to Lower Eifelian. trijugatus (ROEMER, 1844). Both our specimens and the
O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi typical specimens of P. cultrijugatus cultrijugatus
(sample H 1058), Tin Rerhoh area, section 5 (sample H (ROEMER, 1844) from Belgium and Germany (Vander-
1088). cammen 1963, Solle 1971) have an acute fold and angular
to subangular bottom on the ventral sulcus. Our species dif-
fers only by less acute bottom of the ventral sulcus but it
Subfamily Paraspiriferinae PITRAT, 1965
still remains less rounded than in the European species P.
cultrijugatus frechi SOLLE, 1971, P. praecursor SOLLE,
Genus Paraspirifer WEDEKIND, 1926
1971, and P. curvatissimus SOLLE, 1971. By morphology,
Ty p e s p e c i e s : Spirifer cultrijugatus F. ROEMER, our specimens fall into the cultrijugatus group of the Euro-
1844; Lower Devonian, Emsian; Germany. pean paraspiriferinids (Solle 1971), which are characteristic
for the late Upper Emsian and early Eifelian time.
A close affinity of the Saharan species to the American
Paraspirifer aff. cultrijugatus (ROEMER, 1844)
representatives of the genus was suggested by earlier au-
Pl. 9, figs 10–19
thors (Racheboeuf 1990a, 1990b). However, the American
1952a Spirifer (Paraspirifer) cultrijugatus (F. ROEMER): Le paraspiriferinids of which revision has been published by
Maïtre, p. 335. Godefroid et Fagestrom (1983) can be differentiated from
M a t e r i a l : One incomplete shell, two ventral valves, our specimens. The species P. conradi GODEFROID et
one external mould of ventral valve and an incomplete in- FAGESTROM, 1983 and P. clarkei GODEFROID et
ternal mould of a dorsal valve. FAGESTROM, 1983 differ by rounded folds and sulcus. P.
D e s c r i p t i o n . The shell is large, slightly transverse, of bownockeri (STERWARD, 1927) differs by earlier dichoto-
bilobate outline, widest in anterior third, dorsi-biconvex, my of plications. P. halli GODEFROID et FAGESTROM,
highly uniplicate, with moderately thick shell. The maximum 1983 and P. acuminatus (CONRAD, 1839) are most similar
shell width estimated from incomplete shells is 58 mm. to our specimens. The latter has less numerous lateral plica-
The dorsal valve is highly carinate, with feebly convex tions, its fold is less acute, flanks of the dorsal valve are
and steeply sloping flanks. In transverse profile the flanks more convex and its fold is narrower than in our specimens.
subtend 100° angle. The fold is prominent, subangular, with P. halli is the most similar of all North American species but
rounded crest. There is only a weak angle between the side it is more globose and its shell is more transverse. Poorly
of fold and the adjacent flank. The dorsal interarea is very preserved paraspiriferinids (P. aff. cultrijugatus, P. aff. P.
low, anacline. The notothyrium is open. The shell interior is acuminatus, and P. ? sp.) are known from early to late Eife-
poorly known, with internal ribbing distinct only near the lian of Venezuela (Benedetto 1984) and these taxa need a re-
anterior commissure. vision.
The ventral valve is strongly convex, with significantly In northwest Africa the genus is known north as well as
arched flanks in the transverse profile. The valve is deepest south of the Reguibat Shield (Le Maïtre, 1952a, 1952b,
at the posterior third. The hinge line occupies about 80 % of Drot 1964) and its stratigraphical significance has repeated-
the shell width. The interarea is rather small, apsacline, ly been discussed by Racheboeuf (1990a, 1990b). Although
strongly curved. The delthyrium is open. The sulcus is deep, a mere presence of the genus can indicate at least the late
with straight slopes and rounded bottom, evenly and signif- Upper Emsian to Givetian age, the species of the P. cultri-
icantly expanding anteriorly at about 25–28° angle. It occu- jugatus group are restricted to the Upper Emsian and Low-
pies 40–45 % of shell width at anterior margin. The tongue er Eifelian in Europe and the morphologically similar North
is short, triangular with rounded apex. Interior is known on- American species P. acuminatus and P. halli are of the Eife-
ly in small specimens. The dental plates are short, thin, lian age (Onondaga Limestone, Jeffersonville Limestone).
widely divergent. Muscle scars are deeply impressed in Our specimens differ from specimens of P. cultrijugatus fig-
large specimens but very weak in small individuals. ured by Le Maïtre (1952a; pl. 14, figs 1–6) and Drot (1964;
Ornament consists of asymmetrical radial plications dis- pl. 4, fig. 5) by a subangular fold and they cannot be re-
82
ferred to the same species. These previously described bust, short to very long but invariably present. However,
African species are similar to the subspecies P. cultrijugatus their length significantly fluctuates and sometimes there are
frechi SOLLE, 1971 or to younger American species with differences in their length also between the left and right
long and rounded fold (e.g. P. clarkei). Consequently, the side of the same specimen. The mucronations extend late-
stratigraphical extension of the genus in Central Sahara can rally into evenly wide extensions, with low striated cata-
take longer interval, with few discrete species. However, the cline posterior face and nearly smooth and flat anterior face.
morphology of our specimens from the Tamesna Basin is The anteromedian shell margins are gently curved, with a
uniform and falls into the morphology of the earliest mem- shallow emargination at the edge of the fold. Small dorsal
bers of the genus. Therefore, the late Upper Emsian or ear- valves are nearly flat, but the valves of adults become weak-
ly Lower Eifelian age is suggested. ly arched in lateral profile, with a tendency to geniculation
Distribution: Late Upper Emsian to Lower Eifelian. along the anterior periphery of gerontic specimens. This
Occurrence: Tamesna Basin, Tin Rerhoh area, section 5 geniculation is commonly formed by overlapping short
(sample H 1090), In Debirene section (samples H 1105, growth lamellae and internally is marked by a step-like shell
1106), In Ateï section (sample H 1017). floor. Transverse profile at midlength of the valve is weak-
ly convex, with somewhat flattened cardinal extremities.
The fold is prominent, bald, high, rounded, regularly but
Family Mucrospiriferidae BOUCOT, 1959
weakly expanding anteriorly, bordered by deep interspaces.
Subfamily Mucrospiriferinae BOUCOT, 1959 However, in single specimens (sample H 884), the median
groove, slightly asymmetrical to the shell axis has been
Genus Eleutherokomma CRICKMAY, 1950 found. The dorsal interarea is low, anacline, with broadly
Ty p e s p e c i e s : Eleutherokomma hamiltoni CRICK- triangular notothyrium, which is almost entirely filled by
MAY, 1950; Upper Devonian, Givetian; USA. small, widely triangular, slightly raised ctenophoridium of a
crescentic outline. The ctenophoridium consists of 20–22
high, weakly diverging plates of uniform size. The no-
Eleutherokomma mutabilis sp. n. tothyrial platform is short and thick, highly raised above
Pl. 10, figs 1–19
valve floor, supported by very short and low median ridge.
? 1964 Acrospirifer arduennensis arduennensis (SCHNUR, 1853); Short and thick crural bases extend anteriorly with 90–100°
Drot, p. 50. angle from each other, but they are distinct only in small in-
? 1964 Acrospirifer cf. mosellanus mosellanus SOLLE, 1953; dividuals. Dental sockets are narrowly triangular, rapidly
Drot, p. 54. deepening laterally and bordered by thin and high inner
H o l o t y p e : Complete shell valve (MaMe 0125), fig- socket ridges. A thin myophragm extending towards mi-
ured on pl. 10, figs 15, 18, housed in the Department of dlength of the valve is distinct in small individuals, general-
Palaeontology of the Museum of West Bohemia, Plzeň, ly becoming obscure in large specimens but its strength is
Czech Republic. variable. Muscle scars are deeply impressed, with subtrian-
Ty p e h o r i z o n : Middle Devonian, Eifelian. gular posterior adductor scars onto the second and third pli-
Ty p e l o c a l i t y : Algeria, Tamesna Basin, In Ateï sec- cations. The anterior pair of adductors is situated on the top
tion, sample H 1017. and slopes of the fold at about valve midlength. The anteri-
E t y m o l o g y : mutábilis (Lat.) – allusion to the varia- or adductors scars are divided into a narrower and longer el-
tion of the mucronation length. liptical posterior pair and a shorter, subcircular anterior pair.
M a t e r i a l : Six complete specimens, eight dorsal and The internal plication is distinct over the whole internal sur-
eighteen ventral valves and numerous fragments in sand- face except for the gerontic specimens having posteromedi-
stone. anly thick and deeply pitted pad.
D i a g n o s i s : Unequally biconvex, large Eleutherokom- The ventral valve is strongly convex in a lateral profile,
ma with smooth sulcus, with six to eleven lateral ribs and some 25 to 30 % as deep as long, with the maximum depth
mucronations of variable length, posteriorly thickened inte- at midlength. The flanks are gently convex, extending into
rior of ventral valve with prominent pitted pad; short, pos- lateral extension bordering anterior edge of mucronations.
teriorly thick dental plates; prominent apical pad in ventral This extension is of variable length, making an angle sub-
umbonal chamber and distinctly impressed ventral and dor- dued by mucronation and the lateral margins strongly vary-
sal muscle fields; crural plates absent in adult specimens; ing. The point of inflexion lies immediately before the hinge
concentric ornament of densely spaced short lamellae. line, the lateral margins are weakly to moderately curved.
D e s c r i p t i o n : The shell is unequally biconvex to al- The sulcus is prominent, deep, with rounded bottom, ex-
most plano-convex, in maximum 50 mm wide, rather thick- tended dorsally into a short, semielliptical tongue. The ven-
shelled in adults, with uniplicate commissure. The shell tral interarea is high, apsacline, strongly concave in the
outline is broadly semicircular to subtrapezoidal. Maximum lateral profile, with prominent transverse striation along the
width is at the hinge line. whole length. The delthyrium is open. Teeth are strong, di-
The dorsal valve is strongly transverse, some 30 % as rected almost laterally. The dental plates are distinctly ex-
long as wide (without mucronations). Mucronations are ro- trasinal, diverging at 60–70° degrees from each other.
83
Dental plates are always clearly developed, short, located in posterior of the ventral valve is unknown in E. monodi.
between the second and third lateral plications. Young spec- Plications and interspaces of the new species are higher and
imens have thin dental plates but their bases become strong- subangular, compared with the rounded plication of E.
ly thickened in adults, leaving only acute vertical edges monodi. However, some specimens of A. mutabilis (sample
anteriorly. The gerontic specimens have stout bases of den- H 333) also have low rounded plications separated by shal-
tal plates, especially on inner sides, leaving only narrow and low interspaces. This is better considered to be the result of
deep delthyrial chamber. The shell floor between proximal shell abrasion because these specimens also lack concentric
parts of dental plates is remarkably thick. This pad posteri- lamellose ornament. Rounded plications of E. monodi are
orly borders the muscle field and extends axially as a short probably primary because based on the data of Racheboeuf
ridge between adductor scars. The ventral muscle field is et al. (2004), the shells show minimum transport and fine
poorly impressed in young specimens. In adults the field be- concentric ornament is preserved on these rounded plica-
comes posteriorly deeply impressed and subdued below tions. The remarkable morphological variation of E. muta-
valve floor but its anterior border remains weakly defined. bilis concerns the number of plications, corresponding
The field is broadly triangular, anteriorly bisected by thin outline and length of mucronations. Length of mucronations
low myophragm. The adductor scars are small, long, nar- cannot be used for specific determination because our spec-
rowly elliptical. The accessory diductors are deeply im- imens have both very long, ridge-like mucronations (as de-
pressed. The main diductors are less clearly impressed, with scribed in E. monodi) to short mucronations and some
radially striated surface in anteromedian parts. Posterome- specimens lack any mucronations at all. Similar variation of
dian floor of the large valve is strongly thickened, with mucronation length is known in other spiriferids (e. g.
deeply pitted surface. Mediospirifer spinosum BENEDETTO, 1984 or Mucro-
Ornament on the flanks of the dorsal valve generally spirifer hastatus MERGL et MASSA, 1992) and therefore
consists of 9 to 11 straight subangular plications, regularly their length is not a reliable diagnostic feature. Even the
expanding during growth. Some specimens have only six length of mucronations on the left and right side of the same
plications and their number may significantly vary among specimen may be different (pl. 10, fig. 7).
individuals. Lateral plications rapidly decrease in size, with Although other species of Eleutherokomma are of the
thin and short plications near the cardinal extremity. The Givetian or Frasnian age, the beds bearing the new species
subangular interspaces are narrower than the plications. All are referred to the Eifelian. The species E. mutabilis is as-
plications extend from the umbones. Concentric ornament sociated with the species Paraspirifer aff. cultrijugatus
consists of short low lamellae, evenly distinct along the (ROEMER, 1844) and also other fauna (chonetoids,
whole length and distinct also on shell surface of cardinal Mesoleptostrophia) indicates an earlier age than suggested
extremities behind the most lateral plication. Lamellae are in North American and other species. Concluded from the
arranged in 0.6 to 1.0 mm long intervals, with shorter dis- morphological similarity of both species, the Upper Give-
tance along periphery of large shells, where they common- tian E. monodi is probably a descendant of E. mutabilis.
ly overlap. Radial microornament was not observed. However, it is worth noting that Eleutherokomma is un-
R e m a r k s : The genus Eleutherokomma is characteristic known in the presumed Givetian succession of the Illizi
spiriferid in the Givetian and Frasnian succession of the North Basin of east Algeria (Brice et Latréche 1998), and in the
American fauna (Crickmay 1950). It is also known from Givetian of the Ghadamis and Murzuq Basins (Mergl et
France (Brice 1988), Belgium (Mailleaux 1940), Poland Massa 1992). The mucrospiriferinids are represented there
(Baliñski 1979), Afghanistan (Brice 1971) and China (Su by several species of Mucrospirifer GRABAU, 1931 invari-
1976). Recently, the new species of the genus has been ably with the median rib in the sulcus.
described from the beds of the Upper Givetian to Lower Fras- D i s t r i b u t i o n : Eifelian.
nian age in Hodh area of Mauritania (Racheboeuf et al. 2004). O c c u r r e n c e : Tamesna Basin, Oued Felaou area
A detailed comparison of formerly described species with the (samples H 332, H 333, H 1404, H 1405), In Ateï section
species from Mauritania has been given by Racheboeuf et al. (samples H 1016, H 1017), and In Debirene section (sam-
(2004) and their conclusions are in a great part valid also for ples H 1105, H 1106, H 1107).
the new species. Unlike the North American species, the new
species has bald fold and sulcus (except for a single, probably Superfamily Cyrtospiriferoidea TERMIER et
malformed shell from sample H 884; pl. 10, fig. 19), and the TERMIER, 1949
number of lateral plications is smaller. Family Spinocyrtiidae IVANOVA, 1959
The new species is morphologically closest to E. mono-
Subfamily Spinocyrtiinae IVANOVA, 1959
di RACHEBOEUF et al., 2004 from Mauritania. Unlike the
specimens of E. monodi, the new species differs by few
Genus Mediospirifer BUBLICHENKO, 1956
characters that look stable within the otherwise broad phe-
notypic variability. The muscle scars of the new species are Ty p e s p e c i e s : Delthyris medialis HALL 1843; Mid-
more clearly defined in both valves. A prominent pitted pad dle Devonian, Hamilton Group; New York, USA.
84
Mediospirifer rerhohensis sp. n. lar, approximately as high as wide, with a distinct, flat and
Pl. 11, figs 1–21 thin delthyrial plate. Its dorsal edge is slightly concave and
the surface bears fine growth lines. The delthyrial chamber
H o l o t y p e : Dorsal valve (MaMe 0191), figured on pl. is deep. The sulcus is deep, subangular, with rounded bot-
11, figs 13, 16, 17, 19, housed in the Department of tom. The sulcus extends into moderate long, rounded, dor-
Palaeontology of the Museum of West Bohemia, Plzeň, sally directed tongue. Dental plates are extrasinal, thin,
Czech Republic. high, slightly thickened posteriorly in large shells. They rest
P a r a t y p e : Ventral valve (MaMe 0192), figured on pl. almost perpendicularly at the valve floor and their bases
11, fig. 2, housed in the Department of Palaeontology of the slightly diverge anteriorly, more in large than in small
Museum of West Bohemia, Plzeň, Czech Republic. shells. The plates are weakly thickened in large individuals.
Ty p e h o r i z o n : Middle Devonian, Eifelian. Teeth are large. The ventral muscle field is weakly im-
Ty p e l o c a l i t y : Algeria, Tamesna Basin, Tin Rerhoh pressed anteriorly, with poorly defined borders, posteriorly
area, section 5, sample H 1090. halved by a low and short myophragm.
E t y m o l o g y : After the type locality. The ornament consists of simple, low, rounded, straight
M a t e r i a l : Five ventral and four dorsal valves and plications, separated by narrow angular interspaces. Plica-
many fragments preserved as internal and external moulds tions regularly widen toward commissure. Each flank bears
in sandstone. 17 to 18 plications in large shells, all gradually decreasing
D i a g n o s i s : Mediospirifer with 17–18 low plications in size. Even the most lateral plications, already on cardinal
on each flank and short mucronations at the hinge line, with extremities, are distinct along the whole length. Concentric
high dorsal fold, short tongue and subcarinate dorsal valve; fine lamellose ornament is distinct on surface of large
ventral interarea strongly curved; delthyrial plate long, un- shells. Radial microornament has not been discovered even
divided; dental plates weakly divergent, long. in the best preserved specimens and its absence is probably
D e s c r i p t i o n : The shell is large (the maximum width primary.
is estimated 45–50 mm; based on fragments), subequally bi- R e m a r k s : Although there are numerous specimens,
convex, with highly uniplicate commissure and hinge line the material is highly fragmentary. It allows description but
extended into short mucronations. The shell wall is strong- for an illustration the material is poor. The species is re-
ly thickened posteriorly but thin anteriorly. ferred to Mediospirifer BUBLICHENKO, 1956 because of
The dorsal valve is semicircular in outline, with a shal- the general shell shape, presence of the delthyrial plate, lack
low, angular emargination in anterior margin. Anterolateral of radial microornament and widely spaced short growth
and lateral margins are evenly rounded. Transverse profile lamellae but differs from all thedescribed species by nu-
of the valve is subcarinate, with high fold and gently arched merous features. Mediospirifer is mostly American genus
flanks. The valve is highly and evenly convex in the lateral with range from the Eifelian to Givetian (Hall 1843,
profile. The dorsal interarea is very low, almost orthocline. Benedetto 1984). The species M. spinosum BENEDETTO,
Dental sockets are narrow, deepening and strongly expand- 1984 from the late Eifelian of Venezuela differs from the
ing anterolaterally, widely diverging from each other. The new species by shorter dental plates, wider ventral muscle
ctenophoridium is small, transversely rhomboidal, deeply field, longer mucronations and less numerous lateral plica-
concave, with some 20 high thin plates. It rests on a low, tions. Stratigraphically younger specimens from Venezuela,
short but clearly defined notothyrial platform. Crural bases by Benedetto (1984) referred to North American species M.
are thick, diverging at 100° from each other. The adductor audaculus (CONRAD, 1842) are similar to the new species
scars are deeply impressed in the posterior third of the by less transverse shell, higher number of lateral plications,
valve. The anterior adductor scars rest on the bottom and the and narrower sulcus and fold. The main difference between
slopes of the fold and have coarsely striated surface. Medi- these species is general absence of mucronations at the
an septum is absent, only weak short myophragm extends hinge line and more anteriorly divergent dental plates of M.
from the notothyrial platform between the anterior adduc- audaculus.
tors. The imprints of posterior adductors are weakly im- The genus is known from the Eifelian (upper part of the
pressed laterally to the fold. Their surface bears irregular Bir al Qasr Formation) of the Murzuq Basin in Libya.
oblique wrinkles. The dorsal valve interior shows weakly Havlíček et Röhlich (1987) described and figured mostly
impressed plications. broken shells referred to M. cf. audaculus (CONRAD,
The ventral valve is semicircular in outline, with evenly 1842). These shells, however, differ from our species by less
curved lateral margins. The umbo is strongly curved, direct- convex flanks of ventral valve, more widely divergent den-
ed almost dorsally in large specimens. The valve is highly tal plates and higher number of lateral plications that are
convex in transverse profile with remarkably arched flanks. narrower and higher.
Lateral profile is moderately convex. The maximum depth Of the spinocyrtiids described from the Givetian of the
lies at posterior one-third. The interarea is remarkably high, Ghadamis Basin (Mergl et Massa 1992), only Spinocyrtia
steeply apsacline, strongly concave in lateral profile. Its sur- sp. A has similar subparalell dental plates and rounded high
face bears fine transverse lines. The delthyrium is triangu- fold and deep sulcus.
85
The new species is known from samples H 1090, H resting at bottoms of third lateral plications. Their bases ex-
1105, H 1106 and H 333. Numerous specimens from the tend to one-third of the valve. The ventral muscle field is
first sample are mostly fragmentary and preserved as inter- large, weakly impressed, extending into shell midlength.
nal and external moulds in medium grained weathered sand- The diductors are weakly bordered from resting valve floor,
stone. These shells have slightly weaker plications that the adductor impressions are narrow, running along shell
differ from plications of the shells from other samples but axis. The posterolateral floor is never sealed with shells de-
despite this difference, all specimens are referred to the posits.
same species. Only the specimen from the sample H 1105 Each flank bears about 20 subangular, high and narrow
has preserved fine lamellose microornament. plications, separated by deep interspaces. The sizes of the
Distribution: Lower Eifelian. plications regularly decrease laterally and regularly increase
Occurrence: Tamesna Basin, Oued Felaou area (sample towards the shell periphery. The coarser concentric orna-
H 333), Tin Rerhoh area, section 5 (sample H 1090), In De- ment consists of short growth lamellae, commonly overlap-
birene section (samples H 1105, H 1106). ping, arranged in few concentric bands. The bands are
internally marked by strong concentric bands that make
Genus Subcuspidella MITTMEYER, 1965 prominent ridge in lateral shell profile. Fine concentric or-
nament consists of growth fila of uneven size.
Ty p e s p e c i e s : Spirifer subcuspidatus SCHNUR, R e m a r k s : In North Africa, the genus was already re-
1851; Lower Devonian, Emsian; Germany. ported by Le Maïtre (1952) from Algeria but this determi-
nation was questioned by Gourvennec (in Boumendjel et al.
Subcuspidella cf. subcuspidata (SCHNUR, 1851) 1997). Drot (1986) referred to Subcuspidella MITTMEYER
Pl. 12, figs 1–8 1965 a few poorly preserved specimens from the Upper
1952b ? Spirifer (Hysterolites) subcuspidatus SCHNUR; Le Emsian sandstones of the Tamesna Basin. The new materi-
Maïtre, p. 121, pl. 15, fig. 38. al is much more similar to the type species and the speci-
1986 Subcuspidella ? sp.; Drot, p. 513, pl. 1, figs 5, 6. mens determined as S. cf. subcuspidata (SCHNUR, 1851)
M a t e r i a l : Two dorsal valves and twelve, mostly in- by Godefroid (2001) from the Upper Emsian of Belgium.
complete ventral valves preserved as internal and external The genus is known only from the Emsian of Germany
moulds in sandstone. (Mittmeyer 1972), Belgium (Vandercammen 1963, Gode-
D e s c r i p t i o n : The shell is large, at least 50–55 mm froid 2001, Godefroid et al. 2002) and France (Drot 1962).
wide, ventri-biconvex, widest at the hinge line. The dorsal Its occurrences in other regions of Europe (Carls 1969,
valve is broadly triangular, with acute cardinal extremities, Gourvennec 1989) and reports of another age were subse-
without distinct mucronations. Smaller specimens are more quently questioned by Godefroid (2001). It is worth to note
transverse than the adult ones, in which DvL/DvW is about that some of our small sized specimens are remarkably sim-
0.60. The anterior margin is slightly emarginate, lateral ilar, mostly by transverse outline and apsacline ventral in-
margins are weakly curved anteromedianly, laterally be- terarea, to Tenuicostella MITTMEYER et GEIB, 1967. This
coming straight. The valve is gently convex in transverse genus is restricted to the Pragian and Lower Emsian and it
profile and slightly convex in lateral profile, with maximum is by some authors considered synonymous to Subcuspidel-
depth at the posterior third. The interarea is almost ortho- la (Racheboeuf et al. 2004). However, the procline ventral
cline, low, with fine transverse striation. The fold is narrow- interarea of other small specimens and the weak convexity
ly triangular, expanding anteriorly, depressed along its axis, of the ventral valve make the attribution of our specimens to
bordered by deep, slightly divergent and curved interspaces. Subcuspidella probable.
The notothyrial platform is high, short, unsupported by a D i s t r i b u t i o n : Upper Emsian.
median ridge. The dental sockets are moderately divergent, O c c u r r e n c e : Tamesna Basin, In Ateï section (sample
deep, short, rapidly expanding anterolaterally. The cardinal H 1003), In Rahir Tin Amzi section (samples H 1057, H
process is unknown. Crural bases are divergent, thick and 1058), Tin Rerhoh area, section 5 (sample H 1088).
short. Adductor scars are weakly impressed laterally to the
fold. Internal plications are more distinct anteriorly, posteri- Tenuicostella MITTMEYER et GIEB, 1967
orly obscured by shell deposits. Ty p e s p e c i e s : Spirifer subcuspidatus var. tenuicosta
The ventral valve is hemipyramidal in small specimens SCUPIN, 1900; Lower Devonian, Emsian; Germany.
with a remarkably high, catacline to almost procline inter-
area. The sulcus is deep, with flattened bottom, anteriorly
Tenuicostella ? sp.
extending into moderately long roundly triangular tongue.
Pl. 12, figs 9–15
The interarea of large valves is steeply apsacline, strongly
curved apically but weakly curved along the hinge line. The 1986 Brachyspirifer ? sp.; Drot, p. 513, pl. 1, figs 9, 16.
delthyrium is narrowly triangular, with sides enclosing 30° 1986 Tenuicostella ? sp.; Drot, p. 513, pl. 1, fig. 8.
angle. Its apical third is closed by slightly depressed trian- M a t e r i a l : Five ventral and one dorsal valve preserved
gular delthyrial plate. Ventral interior shows broadly diver- as internal and external moulds in sandstone.
gent, extrasinal, short and posteriorly obsolete dental plates, D e s c r i p t i o n : The shell is large, with width 45 to 50
86
mm, ventri-biconvex, transverse, about 50–60 % as long as the dorsal valve. Therefore, the generic position of
wide, moderately thick. Tenuicostella ? sp. is unclear and needs further study.
The dorsal valve is moderately convex with semicircular D i s t r i b u t i o n : Late Upper Emsian and Eifelian.
shell outline. O c c u r r e n c e : Tamesna Basin, Oued Felaou area
The ventral valve is depressed hemipyramidal, of wide- (sample H 333), In Rahir Tin Amzi section (sample H
ly reverse trapezoidal outline, widest at hinge line. Cardinal 1058), Tin Rerhoh area, section 5 (sample H 1088), In De-
extremities are narrowly acute in young individuals, during birene section (sample H 1106).
the growth the angle rapidly increases. The interarea is ap-
sacline, high, strongly curved. The delthyrium is highly tri- Genus Spinocyrtia FREDERIKS, 1916
angular, apically with thick delthyrial plate. Transverse
profile of the valve is subcarinate, the flanks are steeply T y p e s p e c i e s : Delthyris granulosa CONRAD,
sloping almost straight or weakly arched. The sulcus is 1839; Middle Devonian, Hamilton Group; New York, USA.
broad, deep, bald, extending dorsally into long semiellipti-
cal tongue and occupying some 25–30 % of shell width at Spinocyrtia sp.
the commissure. Dental plates are broadly divergent, ex- Pl. 10, fig. 20
trasinal, subtending about 90–100° from each other, even in M a t e r i a l : Two dorsal valves preserved as internal and
the juvenile individuals. The dental plates are posteriorly external moulds in sandstone.
very thick, with obsolete inner sides meeting in a delthyrial D e s c r i p t i o n : The shell is some 36 mm wide, semiel-
chamber but their anterior edges are thin, acute and resting liptical, with extended cardinal extremities. The valve has
almost perpendicularly at valve floor. The ventral muscle evenly convex lateral profile, with weakly arched flanks.
field is large, 70 % as long and some 35–40 % as wide as The sulcus is prominent, deep, bald, with sides subtending
the valve, weakly impressed, anteriorly weakly bordered 20° angle. Ornamentation consists of some 20 high and
and laterally bounded by low, medianly converging ridges. rounded plication on each flank. Concentric ornamentation
The adductor scars are narrow, long, the diductor impres- consists of weak, densely spaced growth fila, forming short
sions are large, with distinct radial and oblique wrinkles. lamellae in surface of the sulcus.
Inner plication is distinct only along periphery of large R e m a r k s : The insufficient material makes the generic
valves, being obsolete by shells deposits in most internal affinity of both valves unclear. Both valves are similar to
surface. In small valves, however, the internal plication is spinocyrtiids but from the type species S. granulosa
distinct over the entire shell floor. (CONRAD, 1839) they differ by narrower sulcus and lack
Exterior bears low rounded plications separated by deep of short mucronations at cardinal extremities. The North
and narrow interspaces. Lateral plications rapidly decrease American species S. parvigranulata EHLERS et WRIGHT,
in size extremities. Each flank bears some 16 to 20 plica- 1955 has similarly shaped dorsal valves of young speci-
tions. Concentric ornament consists of growth lamellae of mens.
various strength. Some lamellae produce prominent edges D i s t r i b u t i o n : Givetian.
on shell surface, internally being marked by prominent O c c u r r e n c e : Tamesna Basin, In Ateï section (sam-
thickening of the shell and step-like edge. ples H 820, H 1006).
R e m a r k s : Although the microornament is unknown,
the species is surely a spinocyrtiid due to the presence of a
prominent delthyrial plate and general shell morphology. Order Terebratulida WAAGEN, 1883
However, its taxonomic position is ambiguous. Its morphol- Suborder Centronellidina STEHLI, 1965
ogy is similar especially to Tenuicostella MITTMEYER et
GIED, 1967 but this genus is restricted to the Pragian and Superfamily Stringocephaloidea KING, 1850
Lower Emsian and is substituted by Subcuspidella Family Rhipidothyrididae CLOUD, 1942
MITTMEYER, 1965 in the Upper Emsian in German and Subfamily Rhipidothyridinae CLOUD, 1942
Belgium Devonian sucession. The genus Spinocyrtia
FREDERICKS, 1916 comprises species with the larger sub- Genus Rhipidothyris COOPER et WILLIAMS, 1935
globose shells with prominent microornament that are pre-
sent in the Middle and Upper Devonian. Our specimens are Ty p e s p e c i e s : Rhipidothyris plicata COOPER et
by wide and long dental plates and broadly triangular shell WILLIAMS, 1935; Middle Devonian, Givetian; New York,
similar to the ventral valve of “Spinocyrtia” sp. (Boucot et USA.
al. 1983; p. 120, pl. 4, figs 1–6) from the Akara Formation. R e m a r k s : The genus Rhipidothyris COOPER et
Although the Givetian age of these shells has been suggest- WILLIAMS, 1935 is known from the Eifelian and Givetian
ed, the earlier Eifelian age consistent with our material can- of Bolivia, Europe, Libya, China, South Africa and North
not be excluded. The shells from the Tamesna Basin differ America (Wang et al. 1987, Isaacson 1993, Boucot et Wil-
from spinocyrtiid described from the Ghadamis Basin son 1994). The genus commonly occurs in monospecific
(Boucot et al. 1983, Mergl et Massa 1992) by a coarser pli- communities in shallow subtidal positions, with fossilized
cation, longer dental plates and absence of a myophragm in shells often in articulated conditions. Our occurrences of
87
Rhipidothyris in east margin of the Tamesna Basin are sim- scribed and figured from the Devonian basins on the territo-
ilar, with highly articulated specimens in fine sandstone ry of southwest Libya (Ghadamis and Murzuq Basins) and
beds generally without any other brachiopod fauna. the Tamesna Basin, Aïr Massif, Niger (Boucot et al. 1983,
Havlíček et Röhlich 1987, Mergl et Massa 1992). Great
Rhipidothyris africana BOUCOT, MASSA morphological variability of this species, especially the
et PERRY, 1983 number of ribs on specimens from the same bed, indicates
Pl. 13, figs 4–12 that species R. ensicostata HAVLÍČEK, 1984 can be re-
ferred to the species R. africana BOUCOT et al., 1983. The
1983 Rhipidothyris africana new species; Boucot et al., p. 117, pl.
specimens from the Tamesna Basin cannot be safely differ-
3, figs 1–27.
1984 Rhipidothyris ensicostata sp. n.; Havlíček, p. 59, figs 16, 17.
entiated from the specimens from the type locality (Give-
1987 Rhipidothyris ensicostata HAVLÍČEK; Havlíček et Röhlich, tian, Idri section, northern flank of the Murzuq Basin:
p. 163, pl. 1, figs 14–18. Boucot et al. 1983) although the ribbing is generally coars-
1992 Rhipidothyris africana BOUCOT, MASSA and PERRY, er and lateral ribs are more curved in the specimens from
1983; Mergl et Massa, p. 95, pl. 25, figs 1–15. the Tamesna Basin. The uniformity of shell morphology in
2000 Rhipidothyris ensicostata HAVLÍČEK, 1984; Mergl et Mas- particular localities and beds may be explained by pheno-
sa, pl. 1, figs 5, 6. typic variability of the local populations. The stratigraphical
M a t e r i a l : Some fifty specimens, mainly articulated, range comprises the Givetian (? Lower to Middle) but after
preserved as internal moulds and external moulds in fine- Havlíček (1984) the species appears already in the Eifelian
grained sandstone. (Bir al Qasr Formation of the Murzuq Basin, Libya).
D e s c r i p t i o n : The shell is biconvex, thin-shelled, The species is known also from the Amesgueur Sand-
with shell wall only 0.1 mm thick in the midlength, 12–14 stone, some 150 m thick unit of the suggested late Middle and
mm wide in adult specimens. The anterior margin is recti- Upper Devonian age of the Aïr Massif of northeast Niger.
marginate and strongly crenulate. The shell structure is ex- The species Rhipidothyris africana is strongly reminis-
opunctate, evidenced by well-preserved limonitic infillings cent and may be considered homoeomorphous to Derbyina
of exopunctae in external moulds. CLARKE, 1913 (family Mutationellidae CLOUD, 1942).
The dorsal valve is subcircular, widest at the midlength, Both genera have few coarse, subangular ribs on the shell
with evenly curved margins, moderate convex in transverse exterior. The differences lie in dorsal valve interior, with
and lateral profiles. The maximum depth is at the valve mi- discrete hinge plates not united into the septalium-like
dlength. The dorsal interior has narrow, widely divergent structure developed in Derbyina. Other species referred to
dental sockets. Inner socket ridges are subhorizontal, flat Rhipidothyris have much finer ornamentation, often with
and wide, distinctly bordered from outer hinge plates. Outer rounded costae (Cooper et Williams 1935, Cloud 1942,
hinge plates are narrower, longer, and clearly concave in the Johnson 1971, McKellar 1966, Isaacson 1993, Boucot et
transverse profile. The imperforate cardinal plate has a shal- Theron 2001). Therefore, we consider that the species
low septalium that is basally and anteriorly supported by a Rhipidothyris africana is only loosely related to the group
thin and long median septum, which may extend nearly to of roundly and weakly costate Rhipidothyris and may rep-
midlength of the valve. The crura anteriorly extend from the resent a separate genus.
subhorizontal inner hinge plates but were always broken in D i s t r i b u t i o n : Upper Eifelian or early Givetian.
the studied specimens. The dorsal muscle field is not im- O c c u r r e n c e : Tamesna Basin, In Ateï area (?) (sample
pressed. H 355) and area west of the Aïr Massif (sample G 1000).
The ventral valve is subpyriform in outline, with curved,
pointed apex. The pedicle foramen is submesothyridid, cir-
Rhipidothyris ? sp.
cular, with distinct pedicle collar. Deltidial plates are dis-
Pl. 13, figs 1–3, 13
crete. The dental plates are very thin, long, moderately
divergent anteriorly and resting perpendicularly on the M a t e r i a l : Two ventral valves and a single dorsal
valve floor. The ventral muscle field is not impressed in any valve, preserved in fine-grained sandstone. Some forty
specimens available. poorly preserved external moulds and shell fragments may
The shell bears coarse straight ribs, all extending from belong to the same species.
the umbo; exceptionally the new and weaker rib may appear D e s c r i p t i o n : The shell is minute, 8 mm wide, ven-
by an implantation. The ribs are subangular, about half as tribiconvex. The dorsal valve is subcircular, with flattened
high as wide, divided by deep, subangular interspaces of median sector and 12 strong rounded plications. The dorsal
identic size. There are 14 to 18 ribs on the exterior of the interior has a thin and long median septum.
large valves. The rib size rapidly and regularly decreases The ventral valve is elongate-oval, strongly convex, with
posterolaterally. The concentric ornamentation is restricted coarse rounded costae. Two median plications are coarser
to fine growth lines. The lines are coarser and more dense- and separated by wider interspaces from lateral plications,
ly spaced along the anterior periphery of the large valves, forming an incipient fold. The dental plates are thin and
forming short lamellae. gently divergent.
R e m a r k s : The shells are similar to specimens de- R e m a r k s : This species is referred to Rhipidothyris
88
COOPER et WILLIAMS, 1935 although it is also similar to evenly convex in the transverse profile, and moderately con-
Cydimia CHATTERTON, 1973 because of few rounded pli- vex in the lateral profile with the maximum of convexity
cations instead of subangular or angular ribs of the latter posteriorly. The depth maximum lies between the posterior
genus. Cydimia is an Upper Emsian genus from Australia one-third to one-fourth of the valve. The dental plates are
(Chatterton 1973). thin and short, weakly divergent anteriorly, never obsolete,
Two rocks samples with shells are labelled H 355. The resting almost perpendicularly on the valve floor. The ven-
first is ferruginous sandstone with poorly preserved external tral muscle field is poorly impressed with a weak
and internal moulds of disarticulated specimens that have myophragm posteriorly in the umbonal chamber.
size and ornamentation characteristic for the species R. The shell ornamentation consists of uniform rounded
africana BOUCOT et al., 1983. The second slab of almost costae, all extending from the umbo. The costae gradually
identical sandstone has the bedding plane covered by poor- grow anteriorly and regularly diminish laterally to become
ly preserved external moulds of smaller, more elongate and obscure posterolaterally. The costae are separated by nar-
convex shells that have less distinct costation and are very rower, deep interspaces. There are 50–55 costae in large
similar to the shells in sample H 1009. These smaller shells shells, numbering 4–5 per 5 mm anteromedianly. The large
may belong to R. sp. but it cannot be excluded that these valves have one or two strong concentric lamellae situated
shells are worn small shells of R. africana. anteromedianly.
D i s t r i b u t i o n : Late Eifelian ? and Givetian. R e m a r k s : As the type specimen of the species Rhipi-
O c c u r r e n c e : Tamesna Basin, In Ateï section (sam- dothyris orchas from the Bir al Qasr Formation (Eifelian of
ples H 1009), In Ateï area (?) (sample H 355). the Murzuq Basin, Libya; Havlíček 1984) has similar orna-
mentation and convexity, the specimens from the Tamesna
Subfamily Globithyridinae CLOUD, 1942 Basin are referred to the same species although the holotype
is somewhat smaller. The shape of the cardinalia and the
Genus Globithyris CLOUD, 1942 pedicle foramen of the holotype as well as of our specimens
are essentially unknown, other characters of the shells war-
T y p e s p e c i e s : Rensselaeria callida CLARKE, rant their attribution to Globithyris CLOUD, 1942. The
1907; Lower Devonian, Emsian; Maine, USA. species G. callida (CLARKE, 1907) from the Lower Devo-
nian (Tomhegan Formation) of Northern Maine (Boucot
Globithyris orchas (HAVLÍČEK, 1984) 1973) has coarser ornamentation, its outline is more elon-
Pl. 14, figs 1–8 gate and the crural plates are larger compared with the del-
1984 Rhipidothyris orchas sp. n; Havlíček, p. 58, figs 14, 15.
icate cardinalia of the species from the Tamesna Basin. The
? 1984 Globithyris sp.; Alvarez et Boucot in Boucot et al., p. 1199, species G. diania (CLARKE, 1907) has distinctly coarser
fig. 1 A–P. ornamentation and more erected ventral umbo.
1987 Rhipidothyris orchas HAVLÍČEK; Havlíček et Röhlich, p. Remarkably similar are two articulated specimens of
163, pl. 2, figs 3–6. Globithyris sp. from the Hammamiat member of the Jauf
1992 Rhipidothyris orchas HAVLÍČEK, 1984; Mergl et Massa, p. Formation (earlier Emsian age) of Saudi Arabia (Boucot et
96, pl. 25, figs 16, 17. al. 1989). These specimens have been recognized among
M a t e r i a l . Two conjoined shells, one dorsal and three fossils from a bryozoan thicket with dominant athyridid bra-
ventral valves. chiopod Sulcathyris ? arabica ALVAREZ et BOUCOT,
D e s c r i p t i o n : The shell is large, equally biconvex, 1984. The globithyridid shells have similar size, convexity
thin-walled, with obscure plicate and finely serrate anterior and ornamentation as the specimens from the Tamesna
commissure, 23–26 mm wide in adult specimens. Basin. The stratigraphic range of African globithyridid bio-
The dorsal valve is transversely oval, with evenly curved facies comprises early Emsian to Eifelian. Some athyridid
margins except for the less curved anterolateral part. The brachiopods (Anathyris, Sulcathyris) have similar extension
valve is regularly and moderately convex in the transverse indicating the similar environmental requirements. Athyrids
profile, with the deepest part in the posterior third and less- are known from Saudi Arabia (Boucot 1984, Boucot et al.
er convexity anteriorly. The maximum depth equals to 1989) and the Ghadames Basin of Libya (Mergl et Massa
20 %, the length equals to 80 % of the valve width. The dor- 1992). Although Isaacson (1974) referred to Globithyris
sal valve interior shows very thin median septum that ante- several specimens from the Emsian of Bolivia, they were
riorly extends to one-third of the valve. Dental sockets are later described as the new species of another genus (Rhipi-
small, long and widely diverging. The cardinal plate is un- dothyris ava ISAACSON, 1993) and their age has been re-
known. evaluated (Racheboeuf et al. 1993).
The ventral valve is broadly and roundly rhomboidal in The globithyridid biofacies ranges from the Siegenian to
the outline, with a prominent umbo. The palintrope is dis- Lower Emsian (Boucot 1963, 1975). The biofacies com-
tinct, high, anacline and weakly curved. The nature of the monly occurs in a nearshore, shallow subtidal position,
pedicle foramen is unknown. The valve is widest posterior probably in quiet waters. Globithyridids commonly form
to midlength, with rounded cardinal extremities. The hinge monospecific aggregations of shells. Our sample shows the
line occupies 30–40 % of the valve width. The valve is similar taphonomy. Several shells are complete and other
89
fauna is restricted (Tropidoleptus). The geographic exten- O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec-
sion of the globithyridid biofacies is very broad and com- tion (sample H 1053).
prises northern Appalachians, Rheinische Schiefergebirge,
Ardenne (Boucot 1963, Jux 1981), Saudi Arabia (Boucot
Family Centronellidae WAAGEN, 1882
1984) and Australia (McKellar 1966). If the Libyan species
G. orchas (Bir al Quasr Formation, Eifelian, Murzuq Basin
Subfamily Amphigeniinae CLOUD, 1942
of south Libya; Havlíček et Röhlich 1987), the Saudi Ara-
bian species G. sp. (Alvarez et Boucot in Boucot et al. Genus Amphigenia HALL, 1867
1984), and the specimens from the Tamesna Basin actually Ty p e s p e c i e s : Pentamerus elongata VANUXEM,
belong to the same taxon, then the extent of the globithyri- 1842; Middle Devonian, Eifelian; New York, USA.
did biofacies of the Emsian and Eifelian ages is remarkable,
covering the space from Western Sahara to North Arabian Amphigenia cf. elongata (VANUXEM, 1842)
Peninsula. In addition, Boucot et al. (1995) briefly dis- Pl. 14, figs 9–13, 15
cussed the systematic position of a weakly costate glo-
bithyridid from Libya and referred Neoglobithyris 1966 Amphigenia aff. elongata (VANUXEM, 1842); Drot, p. 373.
HAVLÍČEK, 1984 to genus Globithyris. This extends the 1986 Amphigenia cf. elongata (VANUXEM, 1842); Drot, p. 513,
pl. 1, figs 12–15.
vertical extension of the globithyridid biofacies into the
Frasnian. The patchy distribution of localities with glo- M a t e r i a l : Complete large dorsal valve and four small
bithyridid biofacies confirms specific ecological require- fragments of ventral and dorsal valves in sandstone.
ments of these terebratuloids. D e s c r i p t i o n : The dorsal valve is very large (77 mm
D i s t r i b u t i o n : Emsian to Lower Eifelian age. A small long), elongate-oval, with weakly truncate anterior margin
fragment of the ventral valve (pl. 14, fig. 5), which may be- and convergent anterolateral margins. The maximum width
long to the same or related species, is known in the Upper lies slightly posterior to midlength. The valve is strongly
Pragian. convex, with almost geniculate posterolateral margins and a
O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec- less convex central part. The anterior commissure is recti-
tion (sample H 1052), Anou Izileg area (sample H 823). marginate. The cardinal plate and crural plates are united in-
to a massive, sessile, block-like structure that is posteriorly
perforated. The anterior slope of the block extends into a
Family Mutationellidae CLOUD, 1942
pair of closely adjacent crural plates with parallel, anterior-
ly extended bases bordering a weak myophragm. The ven-
Genus Pleurothyrella BOUCOT, CASTER, IVES et
tral surface of the block is posterolaterally flat and
TALENT, 1963 anteromedianly weakly concave. The cardinal process is ab-
sent. Anterolateral edges of the block are acute and proba-
Pleurothyrella cf. knodi CLARKE, 1913 bly deeply excavated (this is obscured by preservation.).
Pl. 13, figs 14–16
The dental sockets are large, deep, diverging at approxi-
M a t e r i a l : One poorly preserved shell. mately 100°, with high and long socket ridges. The right
D e s c r i p t i o n : Shell is large, weakly ventri biconvex, canal of weakly divergent vascula media is poorly im-
rectimarginate, broadly subpentagonal in outline with max- pressed (Pl. 14, fig. 9) on the worn surface of the internal
imum width in the posterior third, 35 to 40 mm long (apex mould.
is broken). Ornament consists of 60 to 70 simple, straight Two fragments belong to small, about 10 mm long
and rounded costellae, gradually diminishing laterally and (based on two fragments) ventral valves (Pl. 14, fig. 10).
crossed by three prominent concentric lamellae. The dorsal The ventral valve interior shows narrow anteriorly tapering
valve interior has a weak median ridge. spondylium resting directly on the valve floor. The third
R e m a r k s : Although the shell interior is almost un- fragment belongs to a larger subcarinate ventral valve with
known, the shell size, shape and ornament are identical with a long and strong median septum (Pl. 14, fig. 15). Its deeply
the illustrated specimen of P. knodi CLARKE, 1913 from V-shaped spondylium is raised, at least anteriorly, above the
Bolivia (Boucot et al. 1963). The species originally de- valve floor.
scribed from Bolivia, has been by other authors later identi- R e m a r k s : The centre of distribution of Amphigenia
fied in Guinea-Bissau of West Africa (Drot et Villeneuve HALL, 1867 lies in the Eastern America Realm (Boucot et
1985) and more recently, almost identical shells were re- Wilson 1994). The ranges of formerly described species
ported in the Parecis Basin of Brazil (Boucot et al. 2001). cover the Upper Emsian to Eifelian (Esopus Formation and
The presence of Pleurothyrella cf. knodi in the Upper Pra- Kanouse Sandstone; Boucot 1959), with the earlier “small”
gian to Lower Emsian of the Tamesna Basin is of extraordi- and later “large” Amphigenia (Boucot et Wilson 1994).
nary palaeogeographic importance, indicating the extension Apart from North America, this genus has been reported
of the Malvinokaffric Realm brachiopod fauna as far as to from late Emsian and Early Eifelian of Venezuela (Benedet-
Central Sahara. to 1984), from Brazil (Boucot 1959) and from two sites in
D i s t r i b u t i o n : Upper Pragian to Lower Emsian. the western and southern Sahara. It is known from the
90
southeast border of the Taoudeni Basin (Drot 1966) and Maryland, USA. However, the lateral margins are not intro-
from the Tamesna Basin (Drot 1986). The specimens from verted; therefore it cannot be not excluded that the shell be-
the Tamesna basin, figured by Drot (1986) probably belong longs to a different genus.
to young individuals because all are about half size of our D i s t r i b u t i o n : Upper Pragian.
large specimen. Three newly figured fragments derive from O c c u r r e n c e : Tamesna Basin, In Rahir Tin Amzi sec-
the same sample (H 1088; collection of Lionel Lessard) as tion (sample H 1052).
the shells figured by Drot (1986).
As noted by Boucot (1959) the foramen perforating the
References
cardinal plate may be closed in large specimens. This is ap-
parent also in our large dorsal valve. Our specimens are by Balinski, A. (1979): Brachiopods and Conodonts from the Frasni-
the large size most similar to A. elongata (VANUXEM, an of the Debnik Anticline, Southern Poland. – Palaeontologia
1842). Other species A. chikasawensis BOUCOT 1959, A. Polonica, 39: 3–95.
preparva BOUCOT, 1959 and A. curta (MEEK et Bellini, E., Massa, D. (1980): A stratigraphic contribution to the
WORTHEN 1968) from New York, USA, are generally Palaeozoic of the southern basins of Libya. – In: Salem M. J.,
Busrewill, M. T. (eds.): The Geology of Libya. Academic
smaller and differ by morphology of the ventral valves and
Press. London, pp. 3–56.
shell convexity. Because of the fragmentary preservation of Bellion, Y. (1989): Histoire géodynamique post-paléozoique de
our ventral valves, it is difficult to compare reliably the l’Afrique de l’Ouest d’apres l’étude de quelques basins sédi-
Tamesna specimens with the American species. It seems mentaires. – Publications occasional CIFEG, 17: 302.
likely that the species from Tamesna Basin (Drot, 1986; pl, Benedetto, J. L. (1984): Les Brachiopodes dévoniens de la Sierra
1, figs 12–14) has high and strong ventral septum and a de Perija (Venezuela). Systématique et implications paléogéo-
raised spondylium similar to A. elongata. This distinguish- graphiques. – Biostratigraphie du Paléozoique, 1: 1–191.
es the African species from the earlier “small” Amphigenia Beuf, S., Biju-Duval, B., De Charpal, O., Rognon, P., Gariel, O.,
in the suggested Amphigenia phylogeny of New York, USA Bennacef, A. (1971): Les grés du Paléozoique inférier du Sa-
hara. – Science Technique Pétrole, 18: 464 pp.
(Boucot et Wilson 1994) and indicates the Middle Devoni-
Bigotte, G., Obeliane, J. M. (1968): Décourvertes de minéralisa-
an age. The Middle Devonian “large” Amphigenia has the tions Uraniumferes au Niger. – Mineralium Deposita, 3:
spondylium well supported by median septum. Therefore, 317–333.
the latest Emsian or earliest Eifelian age of our Amphigenia- Borghi, P. (1939): Fossili Devonici del Fezzan. – Annali Museo
bearing samples is suggested. This age is confirmed by oth- Libico, Storia Naturale, Tripoli, 1: 115–184.
er brachiopods, mainly spiriferids. Boucot, A. J. (1959): Brachiopods of the Lower Devonian rocks at
The late Emsian or early Eifelian occurrence of Amphi- Highland Mills, New York. – Journal of Paleontology, 33(5):
genia strongly supports the North American affinity of the 727–769.
brachiopod fauna in the Tamesna basin. Boucot, A. J. (1960): Lower Gedinnian Brachiopods of Belgium.
– Mémoires de l’Institut géologique de l’Universite de Lou-
D i s t r i b u t i o n : Upper Emsian to early Eifelian.
vain, 21: 280–324
L o c a l i t i e s : Tamesna Basin, In Rahir Tin Amzi sec- Boucot, A. J. (1963): The Globithyrid Facies of the Lower Devo-
tion (sample H 1058), Tin Rerhoh area, section 5 (sample H nian. – Senckenbergiana lethaea, 44 (1): 79–84.
1088). Boucot, A. J. (1973): Early Paleozoic Brachiopods of the Moose
River Synclinorium. Maine. – Geological Survey Professional
Subfamily Eurythyridinae CLOUD, 1942 Paper, 784: 1–81.
Boucot, A. J. (1975): Evolution and Extinction Rate Controls. –
Genus Eurythyris CLOUD, 1942 Elsevier. Amsterdam. 427 p.
Boucot, A. J. (1984): Old World Realm (Rhenish-Bohemian re-
Ty p e s p e c i e s : Oriskania lucerna SCHUCHERT, gion), shallow-water, early Devonian brachiopods from the
1913; Lower Devonian, Pragian; Maryland; USA. Jauf Formation of Saudi Arabia. – Journal of Paleontology, 58
(5): 1196–1202.
Eurythyris ? sp. Boucot, A. J., Wilson, R. A. (1994): Origin and early radiation of
Pl. 14, fig. 14 terebratuloid brachiopods: thoughts provoked by Prorensse-
laria and Nanothyris. – Journal of Paleontology, 68 (5):
M a t e r i a l : External mould of the dorsal valve in sand- 1002–1025.
stone. Boucot, A. J., Johnson, J. G. (1968): Brachiopods of the Bois
D e s c r i p t i o n : The dorsal valve is 20 mm wide, 110 % Blanc Formation in New York. – Geological Survey Profes-
as long as wide, subpentagonal in outline, almost flat be- sional Paper, 584–B: 1–27.
coming convex from the anterior one-fourth. The shell is Boucot, A. J., Harper, C. W. (1968): Silurian and Lower Devonian
Chonetacea. – Journal of Paleontology, 42(1): 1143–176.
smooth, without any sign of radial ornament.
Boucot, A. J., Theron, J. N. (2001): First Rhipidothyris (Brachio-
R e m a r k s : Is it impossible to make exact determina- pod) from southern Africa: Biostratigraphic, palaeoecological,
tion from the material but the presence of a terebratulid in biogeographical significance. – Journal of the Czech Geologi-
the sample H 1052 is noteworthy. The flat dorsal valve is cal Society, 46(3): 155–160.
strongly similar to the valve of Eurythyris lucerna Boucot, A. J., Caster, K. E., Ives, D., Talent, J. A. (1963): Rela-
(SCHUCHERT, 1913) from the Oriskany Sandstone of tionships of a new Lower Devonian Terebratuloid (Brachio-
91
pod) from Antarctica. – Bulletins of American Paleontology, Cloud, P. E. (1942): Terebratuloid Brachiopoda of the Silurian and
46(207): 81–151. Devonian. – Geological Society of America, Special Paper 38:
Boucot, A. J., Massa D., Perry, D. G. (1983): Stratigraphy, Bio- 1–182.
geography and Taxonomy of some Lower and Middle Devoni- Cocks, L. R. M., Rong, J. (2000): Strophomenida. – In: Williams,
an brachiopod-bearing beds of Libya and Northern Niger. – A., Brunton, C. H. C., Carlson, S. J. et al.: Treatise on Inverte-
Palaeontographica, A, 180(4–6): 91–125. brate Paleontology, part H, Brachiopoda, Revised, Volume 2.
Boucot, A. J., McClure, H. A., Alvarez, F., Ross, J. R. R., Taylor, Geological Society of America, Inc. and University of Kansas
D. W., Struve, W., Savage, N., Turner, S. (1989): New Devoni- Press, Boulder, Colorado, and Lawrence, pp. 216–349.
an fossils from Saudi Arabia and their biogeographical affini- Cooper, G. A., Dutro, J. T. (1982): Devonian brachiopods of New
tites. – Senckenbergiana lethaea 69(5/6): 535–597. Mexico. – Bulletins of American Paleontology, 82–83(313):
Boucot, A. J., Rowell, A. J., Racheboeuf, P., Pereira, E., Gonçalves 1–215.
de Melo, J. H., Peixoto de Siqueira, L. (2001): Position of the Cooper, G. A, Williams, J. S. (1935): Tully Formation of New
Malvinokaffric Realm’s northern boundary (Early Devonian) York. – Geological Society of America, Bulletin, 46: 781–868.
based on newly discovered brachiopods from the Parecis Basin Crickmay, C. H. (1950): Some Devonian Spiriferidae from Alber-
(Brazil). – Journal of the Czech Geological Society 46: ta. – Journal of Paleontology, 24: 219–225.
109–120. Deynoux, M., Sougy, J., Trompette, R. (1985): Lower Palaeozoic
Boucot, A. J., Bahlburg, H., Breitkreutz, C. Isaacson, P. A., rocks of West Africa and the western part of Central Africa. –
Niemeyer, H., Urzua, F. (1995): Devonian brachiopods from In: Holland, C. H. (ed): Lower Palaeozoic rocks of Northwest
Norther Chile. – Journal of Paleontology, 69(2): 257–263. and West Central Africa, John Willey. New York, pp. 337–495.
Boumendjel, K., Brice, D., Copper, P., Gourvennec, R., Jahnke, Drot, J. (1962): Nouvelles données sur la stratigraphique du Pri-
H., Lardeux, H., LeMenn, J., Melou, M., Morzadec, P., Paris, maire des environs d’Aïnhoa (Basses-Pyrénées). – Bulletin de
F., Plusquellec, Y., Racheboeuf, P. (1997): Les Faunes du la Société géologique de France, 7e série, 4: 431–435
Dévonien de l’Ougarta (Sahara Occidental, Algérie). – An- Drot, J. (1964): Rhynchonelloidea et spiriferoidea siluro-dévoni-
nales de la Sociéte geologique du Nord, 5(2eme série): ens du Maroc pré-Saharien. – Notes et Mémoires du Service
89–116. Géologique, 178: 1–287.
Brice D. (1971): Etude paléontologique et stratigraphique du Drot, J. (1967): Présence du genre Amphigenia (Brachiopode,
Dévonien de l’Afghanistan. Contribution a la connaissance des Centronellidae), dans le basin de Taoudeni (Maroc). – Compte
Brachiopodes et des polypiers rugueux. – Notes et Mémoires rendus Sommaire des séances de la Sociéte géologique de
sur le Moyen-Orient, 11: 1–364. France, 9: 373.
Brice, D. (1988): Brachiopodes du Dévonien de Ferques (Boulon- Drot, J. (1986): Un nouveau gisement Saharien á Amphigenia
nais, France). – In: Brice, D. (ed.) Le Dévonien de Ferques. (Brachiopode Terebratulida, Dévonien). Implications
Bas-Boulonnais (N. France). – Biostratigraphie du Paléo- paléogéographiques. – Geobios, 19(4): 511–516.
zoique, 7: 323–395. Drot, J., Villeneuve, M. (1985): Présence de Pleurothyrella knodi
Brice, D., Carls, P., Cocks, L. R. M., Copper, P., Garcia-Alcalde, J. (Brachiopoda, Terebratulida Dévoniens) en Guinée-Bissau.
L., Godefroid, J. Racheboeuf, P. (2000): Brachiopoda. – Implications Paléogéographiques. – Geobios, 18(4): 525–531.
Courier Forschungsinstitut Senckenberg, 220: 65–86. Garcia-Alcalde, J. L. (1997): North Gondwannan Emsian events.
Brice, D., Charriere, A., Drot, J., Regnault, S. (1983): Mise en év- – Episodes, 20(4): 241–246.
idence, par des faunes de Brachiopodes, de l’extension des for- Garcia-Alcalde, J. L. (2001): Paleobiogeographical relationships
mations dévoniennes dans la boutonniére d’Immouzer du between North Gondwana and South Baltica: The Ivanothyris
Kandar (Sud de Fés, Maroc). – Annales de la Sociéte havliceki fauna (Cantabrian Zone, latest Emsian). – Journal of
géologique du Nord, 103, 445–458. the Czech Geological Society, 46(3): 155–160.
Brice, D., Latréche, S. (1998): Brachiopodes du bassin d’Illizi (Sa- Garcia-Alcande, J. L., Racheboeuf, P. (1978): Nouveax brachiopodes
hara Algérien Oriental) prés de la limite Givétien-Frasnien). – Chonetacea du Dévonien de la Cordillère Cantabrique (Nord-
Geobios, 31(4): 437–454. Ouest de l’Espagne). – Géobios, 11(6): 835–865.
Brice, D., Milhau, B., Mistiaen, B. (1994): Affinités nord-améri- Garcia-Alcalde J. L., Truyóls-Massoni, M. (1994): Lower/Upper
caines de taxons dévoniens (Givétien-Frasnien) du Boulon- versus Zlichovian/Dalejan (Lower Devonian) boundary. –
nais, Nord de la France. Migrations et diachronismes. – Newsletter on Stratigraphy, 30: 83–89.
Bulletin de la Société géologique de France, 165: 291–306. Gatisky, Y, G., Klochko, V. P., Rozman, K. S., Trofimov, D. M.
Carls, P. (1969): Zur Einstufung des Devon der östlichen Guadar- (1966): More facts about the Paleozoic stratigraphy of the
rama (Spanien). – Senckenbergiana lethaea, 50(1): 67–79. Southern Sahara. – Doklady Akademii Nauk SSSR, 170 (5):
Carls, P. (1988): The Devonian of Celtiberia (Spain) and Devoni- 1154–1157.
an paleogeography of SW Europe. – In: McMillan, N. J., Em- Godefroid, J. (2001): Description de quelques brachiopodes
bry A. F., Glass, D. J. (eds.). Devonian of the World. Volume I: Spinocyrtiidae (Spiriferida) du Dévonien inférieur de Bel-
Regional Syntheses. Canadian Society of Petroleum Geolo- gique. – Bulletin de l’Institut Royal des Sciences Naturelles de
gists. Calgary, Alberta. pp. 421–466. Belgique, Sciences de la Terre, 71: 5–30.
Carls, P., Gandl, J., Groos-Uffenorde, H., Jahnke, H., Walliser, O. Godefroid, J., Fagerstrom, J. A. (1983): Le genre Paraspirifer
H. (1972): Neue Daten zur Grenze Unter/Mittel Devon. – Wedekind, R. 1926 dans le dévonien moyen de la partie orien-
Newsletter on Stratigraphy, 2(3): 115–147. tale de l’Amérique du Nord. – Bulletin de l’Institut Royal des
Chatterton, AB. D. E. (1973): Brachiopods of the Murrumbidgee Sciences Naturelles de Belgique, Sciences de la Terre, 55(2):
Group, Taemas, New South Wales. – Bureau of Mineral Re- 1–61.
sources, Geology and Geophysics, Bulletin 137: 1–146. Godefroid, J., Stainier, P., Trost, G. (2002): Two new spinocyrtiid
Claret, J., Tempere, C. (1968): Le Paleozoique du basin du Tames- brachiopods (Spiriferida) from the Pragian and Lower Emsian
na (Sud de l’Ahaggar). – Zeitschrift der Deutschen geo- of Belgium. – Bulletin de l’Institut Royal des Sciences Na-
logichen Gesselschaft, 117: 460–468. turelles de Belgique, Sciences de la Terre, 72: 25–41.
92
Gournennec, R. (1989): Brachiopodes Spiriferida du Dévonien in- Jahnke, H. (1971): Fauna und Alter der Erbslochgrauwacke. – Göt-
férieur du Massif Armoricain. – Biostratigraphie du Paléo- tinger Arbeiten zur Geologie und Paläontologie, 9: 1–105.
zoique, 9: 1–281. Jansen, U. (2001): Morphologie, Taxonomie und Phylogenie
Greigert, J., Pougnet, R. (1967a): Notice explicative sur la carte unter-devonischer Brachiopoden aus der Dra-Ebene
géologique de la République du Niger a l’échelle du (Marokko, Prä-Sahara) und dem Rheinischen Schiefergeborge
1/2.000.000. – Publication du Bureau de recherches géologi- (Deutschland). – Abhandlungen der Senckenbergischen Natur-
ques et minières. Paris. forschenden Gesselschaft, 554: 1–389.
Greigert, J., Pougnet, R. (1967b): Essai de descriptiondes forma- Johnson, J. G. (1970): Great Basin Lower Devonian Brachiopoda.
tions géologiques de la République du Niger. – Mémoires – The Geological Society of America, Inc., Memoir 121:
Bureau de recherches géologiques et minières. 48: 1–238. 1–421.
Paris. Johnson, J. G. (1971): Lower Givetian Brachiopods from Central
Hall, J. (1843): Natural History of New York. Geology. – 4: 1–525. Nevada. – Journal of Paleontology, 45(2): 301–326.
Harper, D. A. T., Boucot, A. J. (1978): The Stropheodontacea, Part Jux, U. (1981): Zur stratigraphischen Verbreitung Begischer Glo-
I (Leptostrophiidae, Eostropheodontidae, and Strophonelli- bithyridinen. – Sonderveröff entlichungen des Geologischen
dae). – Palaeontographica, A, 161(1–3): 55–118. Institutes der Universität Köln, 41: 93–107.
Havlíček, V. (1961): Rhynchonelloidea des böhmischen älteren Karpoff, R. (1960): Le Cambro-Ordovicen de l’Adrar des Iforas
Paläozoikums (Brachiopoda). – Rozpravy Ústředního ústavu (Sahara soudanais). – Comptes rendus Sommaire des Séances
geologického, 27: 1–211. de la Sociéte géologique de France, 16–17.
Havlíček, V. (1984): Diagnoses of new brachiopod genera and Kayser, E. (1892): Beitrage zur Kenntnis der Fauna der Siegenis-
species. Part 1. – In: Pařízek, A., Klein, L, et Röhlich, P. (eds.), chen Grauwacke. – Jahrbuch der köninglichen preussischen
Explantory Booklet, Geological Map of Libya, 1/250 000 (NG geologischen Landesanstalt, 2(8): 95–107.
33–1), Sheet Idri. Industrial Research Centre, Tripoli, Libya, Kilian, C. (1928): Sur la présence du Silurien a l’Est et a Sud de
57–59. l’Ahaggar. – Compte rendus de l’Academie des Sciences de
Havlíček, V., Röhlich, P. (1987): Devonian and Carboniferous bra- France, 186: 508–509.
chiopods from northern flank of Murzuk Basin (Libya). – Le Hérissé, A., Paris, F., Racheboeuf, P. R. (1992): Marine paly-
Sborník geologických věd, Paleontologie, 28: 117–177. nomorps from the Devonian of Bolivia. – Table Ronde
Heddebaut, C. (1981): Les Brachiopodes Spiriferacea et Reticu- Européene “Paléontologie et Stratigraphie d’Amerique latine”,
lariacea. – In: Morzadec, P., Paris, F., Racheboeuf, P.: La Lyon 7–9 juillet, Abstracts 32.
Tranchée de la Lezais. Emsien Supérieur du Massif Armori- Le Maïtre, D. (1952a): Contribution à l’étude des faunes paléo-
cain. Mémoires de la Société géologique and mineralogique de zoiques de l’Adrar Mauritanien (Sahara occidental). – Bulletin
Brétagne, 23 (313): 231–247. de la Direction des Mines d’Afrique occidentale francaise,
Herrera Z. A. (1995): The Lower Devonian Chonetoidea bra- 15(2): 299–283.
chiopods from the Argentina Precordillera. – Documentes des Le Maïtre, D. (1952b): La faune de Dévonien inférieur et moyen
Laboratoires de Géologique de la Faculté des Sciences de Ly- de la Saoura et des abords de l’Erg el Djemel (Sud Oranais). –
on, 136: 101–147. Matériaux pour la carte géologique de l’Algerie, 1, Paléon-
Herrera, Z. A, Racheboeuf, P. R. (2001): Les Eodevonariidés tologique, 12: 170 p.
(Chonetoidea, Brachiopoda) du Dévonien des Bassins de Le Maïtre, D. (1961): Découverte de nouveaux gisements africains
Taoudeni et du sud du Hoggar, Sahara. – Géobios, 34(5): de Pustulatia pustulosa (Hall). Répartition géographique et
493–503. stratigraphique de ce Brachiopode en Afrique. – Comptes ren-
Herrera, Z. A., Salas, M. J., Giolitti, J. A. (1998): Chilidiopsi- dus Sommaire des Seances de la Sociéte géologique de
doidea (Brachiopoda) del Devónico Inferior de la Precordillera France, 190–191.
Argentina. – Revista Española de Paleontologia, 13(2): Lessard, L. (1959): Note préliminaire sur la géologie des Tassilis
149–166. Oua-N’Ahaggar (Sahara méridional). – Comptes rendus Som-
Isaacson, P. E. (1974): First South American occurrence of Glo- maire des Seances de la Sociéte géologique de France, 6:
bithyris: its ecological and age significance in the Malvinokaf- 151–152.
fric realm. – Journal of Paleontology, 48(4): 778–784. Lessard, L. (1961): Les séries primaries des Tassilis Oua-n-Ahag-
Isaacson, P. E. (1977a): Devonian stratigraphy and brachiopod pa- gar au Sud du Hoggar entre l’Aïr et l’Adrar des Iforas (Sahara
leontology. Part A: Orthida and Strophomenida. – Palaeonto- méridional). – Bulletin de la Société géologique de France,
graphica, A, 155: 133–192. 7(3): 501–513.
Isaacson, P. E. (1977b): Devonian stratigraphy and brachiopod pa- Lessard, L. (1962): Couverture sédimentaire (series primaries des
leontology. Part B: Spiriferida and Terebratulida. – Palaeonto- Tassilis Oua-n-Ahaggar). Régions d In Azaoua, Ti-m-Missaou,
graphica, A, 156: 168–217. In Guezzam. – In: Notice Explicante de la Carte géologique du
Isaacson, P. E. (1993): Devonian brachiopoda of Bolivia. – In: Hoggar. 81–98. Bureau de recherches géologiques et minières.
Suarez-Soruce, R. (ed.): Fosiles y facies de Bolivia – vol. II. Paris.
Invertebrados y paleobotanica. – Revista Técnica de YPFB, Mailieux, E. (1940): Contribution a la connaissance du frasnien
13–14(1–4): 5–33. moyen (Assise de Frasnes) de la Belgique. – Bulletin du
Isaacson, P. E., Chlupáč, I. (1984): Significance of a Tropidoleptus Musée royal d’Histoire naturelle de Belgique, 16: 1–44.
assemblage from the Devonian of the Moravo-Silesian region, Massa, D. (1988): Paleozoique du Libye occidentale. Stratigraphie
Czechoslovakia. – Časopis pro mineralogii a geologii, 29(2): et Paléogeographie. – Thése doctoire d’Etat, Nice, 2 volumes,
141–154. 1–514. Nice (unpublished).
Isaacson, P. E., Perry, D. G. (1977): Biogeography and morpho- McKellar, R. G. (1966): Additional brachiopods and bivalves from
logical conservatism of Tropidoleptus (Brachiopoda, Orthida) the Etonvale Formation, Adavale Basin, Queensland. –
during the Devonian. – Journal of Paleontology, 51(6): Queensland Geological Survey, Publication 332: 11–17.
1108–1122. Mergl, M., Massa, D. (1992): Devonian and Lower Carboniferous
93
Brachiopods and Bivalves from Western Libya. – Biostratigra- der Zone des Monograptus hercynicus. – Geologische
phie du Paléozoique, 12: 1–115. Jahrbuch, 81: 171–220.
Mergl, M., Massa, D. (2000): A Palaeontological review of the Solle, G. (1971): Brachyspirifer und Paraspirifer im Rheinischen
Devonian and Carboniferous Succession of the Murzuq Basin Devon. – Abhandlungen des Hessischen Landesdamtes für
and the Djado Sub-Basin. – In. Sola, M. A., Worsley, D. (eds.): Bodenforschung, 59: 1–163.
Geological Exploration in Murzuq basin. Elsevier Science B. Sougy, J. (1964): Les formations paléozoiques du Zemmour Noir
V., pp. 41–88. (Mauritanie Septentrionale): Étude stratigraphique, petro-
Mergl, M., Massa, D. (in print): A new giant discinoid brachiopod graphique et paléontologique. – Dakar, Louis-jean Gap,
genus from the Lower Devonian of Algeria. – Acta Palaeonto- 1–695.
logica Polonica. Su, Y. Z. (1976): Brachiopods from Cambrian to Devonian. – In:
Mittmeyer, H.G. (1972): Delthyrididae und Spinocyrtiidae (Bra- Atlas of fossils of North China, Nei-Monggol Volume, Pt. 1,
chiopoda) des tiefsten Ober-Ems im Mosel-Gebiet (Ems- Paleozoic. Geological Publishing House, Beijing, pp.159–227.
Quartzit, Rheinisches Schiefergebirge). – Mainzer Thomas, I. (1905): Neue Beiträge zur Kenntnis der devonischen
geowissenschaftliche Mitteilungen, 1: 82–121. Fauna Argentinies. – Zeitschrift der deutschen geologischen
Pardo, M. V., Garcia-Alcalde, J. L. (1984): Biostratigrafía del Gesselschaft, 57: 233–290.
Devónico de la región de Almadén (Ciudad Real, Espana) – Valsardieu, C. (1970): Cadres géologiques et paléogéographiques
Trabajos de Geología, 14: 79–120. des minéralisations de Carbon, de Cuivre et d Uranium de la
Perrodon, A. (1971): Esquisse tectonique du bassin du Tamesna region d’Agades (République du Niger). – Doctoral Sciences
(Algerie, Niger et Mali). – Tectonique de l’Afrique, Sciences Thesis. Université de Nice, 1–750 (unpublished).
de la Terre, 6: 390-UNESCO. Paris. Vandercammen, A. (1963): Spiriferidea du Dévonien de la Bel-
Racheboeuf, P. R. (1981): Chonetacés (Brachiopodes) siluriens et gique. – Institut Royal des Sciences naturelles de Belgique,
dévoniens du Sud-Ouest de l’ Europe. – Mémoires de la So- Mémoires, 150: 1–177.
ciété géologique et minéralogique de Bretagne, 27: 1–294. Villemur, J. R., Drot, J. (1957): Contribution a la faune Dévoni-
Racheboeuf, P. R. (1990a): Silurian to Middle Devonian chone- enne du bassin de Taoudeni. – Bulletin de la société
tacean brachiopods from the northwestern Gondwanaland géologique de France, 6(7): 1077–1082.
margin: A review in time and space. – In: MacKinnon, D. I., Walmsley, V. G., Boucot, A. J., Harper, C. W. (1969): Silurian and
Lee, D. E., Campbell, J. D. (eds): Brachiopods through time, Lower Devonian salopinid brachiopods. – Journal od Paleon-
319–325. tology, 43(2): 492–516.
Racheboeuf, P. R. (1990b): Paléobiographie de la marge nord- Wang, Y., Boucot, A. J., Rong, J.-Y., Yang, X.-Ch. (1987): Com-
gondwanienne au Dévonien inférieur et moyen: nouvelles don- munity Paleoecology as a Geologic Tool: The Chinese
nées déduites de l’étude des Brachiopodes Chonetacés. – Ashgillian–Eifelian (latest Ordovician through nearly Middle
Compte rendus de l’Academie des Sciences de Paris, 310, Devonian) as an example. – Geological Society of America
série II: 1481–1486. Inc., Special Paper, 211: 1–100.
Racheboeuf, P. R., Feldman, H., R. (1990): Chonetacean Bra- Williams, H. S. (1913): Recurrent Tropidoleptus Zones in the Up-
chiopods of the “pink Chonetes” Zone, Onondaga Limestone per Devonian in New York. – United States Geological Survey,
(Devonian, Eifelian), Central New York. – Novitates, 2982, Professional Paper, 79: 1–103.
1–16. Williams, A., Brunton, C. H. C. (2000): Orthotetidina. – In: Wil-
Racheboeuf, P. R., Gourvennec, R., Deynoux, M., Brice, D., liams, A., Brunton, C. H. C., Carlson, S. J. et al.: Treatise on
(2004): The Devonian of the Hodh area (Islamic Republic of Invertebrate Paleontology, part H, Brachiopoda, Revised, Vol-
Mauritania): paleontology and stratigraphy. – Journal of Pale- ume 3. Geological Society of America, Inc. and University of
ontology, 78(1): 98–110. Kansas Press, Boulder, Colorado, and Lawrence, pp. 644–681.
Racheboeuf, P. R., Isaacson, P. E. (1993): Los Chonetoidos Williams, A., Harper, D. A. T. (2000): Orthida. – In: Williams, A.,
(Braquiópodos) Silúricos y Devónicos de Bolivia. – Revista Brunton, C. H. C., Carlson, S. J. et al.: Treatise on Invertebrate
Técnica de YPFB, 13–14(1–4): 99–119. Paleontology, part H, Brachiopoda, Revised, Volume 3. Geo-
Racheboeuf, P. R., Le Herisse, A., Paris, F., Babin, C., Guil- logical Society of America, Inc. and University of Kansas
locheau, F., Truyols-Massoni, M. (1993a): El Dévonico de Bo- Press, Boulder, Colorado, and Lawrence, pp. 714–782.
livia: Bio y Cronoestratigrafía. – Bulletin de l’Institut Français Williams, A., Wright, A. D. (1965): Orthida. – In: Moore, R. C.:
d’ Études Andines, 22(3): 645–655. Treatise on Invertebrate Paleontology, part H, Brachiopoda,
Racheboeuf, P., R., Le Herissé, A., Paris, F., Babin, C., Guil- Volume 1, Geological Society of America, Inc. and University
locheau, F., Truyols-Masssoni, M., Súarez-Soruco, R. (1993b): of Kansas Press, Lawrence, pp. 299–359.
Le Dévonien de Bolivie: biostratigraphie et chronostratigra- Weddige, K., Werner, R., Ziegler, W. (1979): The Emsian-Eifelian
phie. – Compte rendus de l’Academie des Sciences de Paris, Boundary. – Newsletter on Stratigraphy, 8(2): 159–169.
317, série II: 795–802.
Rathbun, R. (1874): On the Devonian Brachiopods of Ereré,
province de Pará, Brazil. – Bulletin of Buffalo Society of nat- Explanation to the plates
ural Sciences, 1(23): 236–261. All photos by M. M.
Reed, F. R. (1903): Brachiopoda from the Bokkeveld beds. – An-
nals of the South African Museum, 4(3): 165–200. PLATE 1
Solle, G. (1953): Die spiriferen der gruppe arduenensis-inter- Mesoleptostrophia (M.) explanata (SOWERBY, 1842)
medius im Rheinischen Devon. – Abhandlungen des Hessis- A g e a n d l o c a l i t i e s : Emsian to Eifelian; Tamesna Basin, In
chen Landesdamtes für Bodenforschung, 5: 1–156. Rahir Tin Amzi section (1, 6), In Ateï section (2, 4), Kori-Ide-
Solle, G. (1963): Hysterolites hystericus (Schlotheim) (Brachiopo- meg – Touaret section (3), and Tin Rerhoh section (5, 7).
da; Unterdevon), die Einstufung der oberen Graptoli- 1, 6. Dorsal valve, internal mould and latex cast of interior; sam-
ten–Schiefer in Thüringen und die stratigraphische Stellung ple H 1058; MaMe 0048, × 1.2, × 1.7.
94
2. Ventral valves, internal moulds; sample H 1003; MaMe 0043A 11, 14. Ventral valve, internal mould in ventral and posterior
and MaMe 0043B, × 1.1. views; sample H 884; MaMe 0044B, × 2.0 and × 3.8.
3. Ventral valve, internal mould; sample H 884; MaMe 0047, × 1.1. 12. Dorsal valve, latex cast of exterior; sample H 826; MaMe
4. Ventral valve, detail of interarea, internal mould; sample H 0152B, × 2.0.
1003; MaMe 0039, × 6.0. 13. Ventral valve, internal mould; sample H 1004; MaMe 0145, × 2.0.
5. Dorsal valve, internal mould; sample H 1087; MaMe 0024A, × 2.0. 15. Detail of spines on left side, latex cast of ventral valve exteri-
7. Ventral valve with original shell, interior; sample H 1086; or; sample H 884; MaMe 0154, × 7.0.
MaMe 0014, × 1.1. 16. Detail of spines on right side, latex cast of ventral valve exte-
rior; sample H 1018; MaMe 0153B, × 7.0.
PLATE 2
Arcuaminetes racheboeufi sp. n.
Amziella rahirensis sp. n.
A g e a n d l o c a l i t y : Eifelian; Tamesna Basin, Tin Rerhoh section.
A g e a n d l o c a l i t y : Upper Pragian; Tamesna Basin, In Rahir
Tin Amzi section. 17, 21. Holotype, ventral valve, internal mould in ventral and pos-
terior views; sample H 1090; MaMe 0183A, × 6.0.
1. Dorsal valve, internal mould; sample H 1052; MaMe 0011C, × 3.5.
18. Ventral valve, internal mould; sample H 1090; MaMe 0183B, × 6.0.
2. Ventral valve, internal mould; sample H 1052; MaMe 0026B, × 3.5.
19. Ventral valve, latex cast of exterior; sample H 1090; MaMe
3. Ventral valve, internal mould; sample H 1052; MaMe 0074, × 3.5.
0179, × 6.0.
4. Ventral valve, internal mould; sample H 1052; MaMe 0049, × 3.5.
20, 24. Ventral valve, latex cast of exterior in dorsal and posterior
5. Dorsal valve, internal mould; sample H 1052; MaMe 0026F, × 3.5.
views; sample H 1090; MaMe 0183C, × 6.0.
6. Dorsal valve, internal mould; sample H 1052; MaMe 0026E, × 3.5.
22. Paratype, dorsal valve, internal mould; sample H 1090; MaMe
7. Dorsal valve, external mould; sample H 1052; MaMe 0026C, × 3.5.
0185, × 6.0.
8. Ventral valve, internal mould; sample H 1052; MaMe 0070, × 3.5.
23. Dorsal valve, internal mould; sample H 1090; MaMe 0184, × 6.0.
9. Paratype, dorsal valve, internal mould; sample H 1052; MaMe
0062, × 6.0. PLATE 4
10. Holotype, ventral valve, internal mould; sample H 1052;
Monsenetes ? drotae sp. n.
MaMe 0011A, × 3.5.
A g e a n d l o c a l i t i e s : Emsian; Tamesna Basin, In Ateï section
11. Ventral valve, internal mould; sample H 1052; MaMe 0026D, × 3.5.
(1, 2), In Rahir Tin Amzi section (4, 8), and Oued Felaou area
12. Dorsal valve, latex cast internal mould showing median sep-
(3, 5–7, 10, 11).
tum, anderidia and cardinal process with shallow cardinal pit;
sample H 1052; MaMe 00050B, × 6.0. 1. Ventral valve, internal mould; sample H 1003; MaMe 0042A, × 2.2.
2. Ventral valve, internal mould; sample H 1003; MaMe 0042B, × 2.2.
Lomaella amziensis HERRERA et RACHEBOEUF, 2001 3. Holotype, ventral valve, internal mould; sample H 328; MaMe
A g e a n d l o c a l i t y : Upper Pragian and Emsian; Tamesna 0086A, × 2.0.
Basin, In Rahir Tin Amzi section. 4. Dorsal valve, latex cast of cardinal process; sample H 1057;
13. Ventral valve, internal mould; sample H 1052; MaMe 0023, × 2.0. MaMe 0025, × 5.0.
14. Incomplete dorsal valve, internal mould; sample H 1052; 5. Paratype, dorsal valve, interior with cardinal process, anderidia
MaMe 0022A, × 2.0. and median septum; sample H 328; MaMe 0032B, × 2.0.
16. Incomplete dorsal valve; sample H 1086; MaMe 0015, × 2.0. 6, 9. Ventral valve, exterior in ventral and anterior views; sample
17. Ventral valve, internal mould; sample H 1052; MaMe 0031, × 2.0. H 328; MaMe 0086B, both × 2.0.
18. Ventral valve, latex cast of external mould; sample H 1052; 7. Ventral valve, internal mould; sample H 328; MaMe 0030, × 2.0.
MaMe 0050A, × 2.0. 8. Dorsal valve, latex cast of interior; sample H 1057; MaMe
Montsenetes sp. 0027, × 2.3.
A g e a n d l o c a l i t y : Emsian; Tamesna Basin, Tin Rerhoh section. 10. Ventral valve, interior; sample H 328; MaMe 0029A, × 2.3.
11. Dorsal valve, exterior; sample H 328; MaMe 0029B, × 2.3.
15. Ventral valve, internal mould; sample H 1087; MaMe 0024B,
× 4.0. Devonochonetes cf. salemi MERGL et MASSA, 1992
A g e a n d l o c a l i t y : Givetian; Tamesna Basin, In Ateï section.
PLATE 3
12. Ventral valve, internal mould; sample H 820; MaMe 0083, × 2.0.
Monsenetes pervulgatus sp. n. 13. Ventral valve, latex cast of exterior; sample H 1006; MaMe
A g e a n d l o c a l i t i e s : Eifelian; Tamesna Basin, In Ateï section 0073A, × 2.0.
(1–9, 13, 16), and Kori-Idemeg – Touaret area (10–12, 14, 15). 14. Dentral valve, exterior; sample H 820; MaMe 0082, × 2.0.
1, 5. Holotype, ventral valve in ventral and anterior views; sample Longispina sp.
H 1017; MaMe 0120, both × 2.0. A g e a n d l o c a l i t y : Eifelian; Tamesna Basin, Oued Felaou area.
2, 6. Ventral valve in ventral and oblique views; sample H 1017;
MaMe 0121, both × 2.0. 15, 21. Ventral valve, internal mould in ventral and oblique views;
3. Ventral valve, latex cast of exterior; sample H 826; MaMe sample H 1405; MaMe 0135, both × 5.5.
152C, × 2.0. 16. Ventral valve, internal mould; sample H 1405; MaMe 0130, × 5.5.
4. Dorsal valve, latex cast of exterior with cardinal process; the 17. Ventral valve, internal mould; sample H 1405; MaMe 0134A, × 5.5.
same level as sample H 826; MaMe 0178, × 2.0. 18. Dorsal valve, external mould; sample H 1405; MaMe 0133, × 5.5.
7, 8. Two dorsal valves, latex casts of interior with cardinal pro- 19. Ventral valve showing spines, internal mould; sample H 1405;
cess and detail of left valve; sample H 1018; MaMe 0153A and MaMe 0180, × 5.5.
MaMe 0153C, × 2.0 and × 5.0. 20. Ventral valve showing spines, internal mould; sample H 1405;
9. Complete shell showing interareas, sample H 358; MaMe 0186, MaMe 0134B, × 5.5.
× 2.0. anopliid sp.
10. Ventral valve, internal mould; sample H 884; MaMe 0044A, × 2.0. A g e a n d l o c a l i t y : Givetian; Tamesna Basin, In Ateï section.
95
22. Dorsal valve, internal mould; sample H 1006; MaMe 0073B, × 5.5. 16. Ventral valve, internal mould; sample H 1009; MaMe 0017,
× 2.0.
PLATE 5 17. Ventral valve, internal mould; sample H 823; MaMe 0101,
Roemerella sp. × 2.5.
A g e a n d l o c a l i t y : Upper Pragian; Tamesna Basin, Tin Rer- 19. Ventral valve, internal mould; sample H 1052; MaMe 0052A, × 5.5.
hoh section. 22. Ventral valve, internal mould; sample H 1106; MaMe 0140C, × 1.5.
1. Ventral valve, internal mould; sample H 1079; MaMe 0046, × 1.6. Athyris sp.
Salopina sp. A g e a n d l o c a l i t y : Eifelian, In Debirene section.
A g e a n d l o c a l i t y : Emsian; Tamesna Basin, Tin Rerhoh section. 18, 21. Ventral valve, internal mould in ventral and anterior views;
2. Ventral valve, internal mould; sample H 1087; MaMe 0024D, × 5.5. sample H 1106; MaMe 0140A, both × 2.0.
96
Pustulatia tamesnaensis sp. n. 10, 18. Ventral valve, posterior and ventral views; sample H 1017;
A g e a n d l o c a l i t i e s : Emsian to Eifelian; Tamesna Basin, In De- MaMe 0124, both × 1.3.
birene section (13, 14, 16, 20–22), and Tin Rerhoh section (18, 19). 11, 12. Ventral valve, internal mould in ventral and anterior views;
13, 16, 17. Holotype, dorsal valve, internal mould, latex cast of ex- sample H 1090; MaMe 0126, both × 1.3.
terior and external mould, sample H 1107; MaMe 0128, all × 6.0. 13, 17. Incomplete shell, dorsal and anterior views; sample H
14. Dorsal valve, latex cast of interior; sample H 1107; MaMe 1017; MaMe 0122, both × 1.3.
0127A, × 6.0. 14. Ventral valve, anterior view; sample H 1016; MaMe 0131, × 1.3.
18. Small ventral valve, internal mould; sample H 1087; MaMe 15, 19. Ventral valve, latex cast of exterior and detail of microor-
0024F, × 6.0. nament; sample H 1105; MaMe 0129, × 1.3, × 9.0.
19. Small ventral valve, latex cast of exterior; sample H 1087; 16. Small ventral valve, latex cast of exterior; sample H 1106;
MaMe 0024G, × 6.0. MaMe 0141, × 1.3.
20. Ventral valve, latex cast of exterior; sample H 1107; MaMe
0127B, × 6.0. PLATE 10
21, 22. Paratype, ventral valve, internal mould and latex cast of Eleutherokomma mutabilis sp. n.
exterior; sample H 1107; MaMe 0127C, both × 6.0. A g e a n d l o c a l i t i e s : Eifelian; Tamesna Basin, Oued Felaou
area (1–5, 8, 9, 12–14, 17), In Ateï section (7, 15, 18), Tin Rer-
PLATE 8 hoh section (16), In Debirene section (10, 11) and Kori-Ide-
Hysterolites hystericus SCHLOTHEIM, 1820 meg – Touaret area (19).
A g e a n d l o c a l i t i e s : Upper Pragian; Tamesna Basin, In Rahir 1. Ventral valve, internal mould; aequivalent of sample H 333;
Tin Amzi section (1), and Tin Rerhoh section (2). MaMe 0150, × 1.7.
1. Dorsal valve, internal mould; sample H 1052; MaMe 0053, × 4.0. 2, 5. Ventral valve, internal mould and latex cast of exterior; sam-
2. Ventral valve, internal mould; sample H 1080; MaMe 0139, × 1.7. ple H 333; MaMe 0149A, both × 1.7.
3, 8. Ventral valve, latex cast of interior and internal mould; sam-
Filispirifer merzakhsaiensis JANSEN, 2001
ple H 333; MaMe 0149B, × 1.4, × 1.7.
A g e a n d l o c a l i t i e s : Upper Pragian; Tamesna Basin, In Rahir
4. Ventral valve, internal mould; sample H 333; MaMe 0151B, × 1.7.
Tin Amzi section (3, 8–17), and Tin Rerhoh section (4–7).
6. Dorsal valve, latex cast of exterior; sample H 828; MaMe
3. Ventral valve, internal mould; sample H 1052; MaMe 0069A, × 1.7. 0152A, × 1.7.
4. Ventral valve, internal mould; sample H 1080; MaMe 0137, × 1.7. 7. Dorsal valve with uneven mucronations, internal mould; sam-
5. Ventral valve, internal mould; sample H 1080; MaMe 0136, × 1.7. ple H 1004; MaMe 0144, × 1.7.
6. Ventral valve, internal mould; sample H 1080; MaMe 0138, × 1.7. 9. Dorsal valve, internal mould; sample H 1404; MaMe 0146A, × 1.8.
7. Ventral valve, internal mould; sample H 1080; MaMe 0132, × 1.7. 10, 11. Small dorsal valve, internal mould and latex cast of exte-
8. Ventral valve, internal mould; sample H 1052; MaMe 0026H, × 1.7. rior; sample H 1106; MaMe 0140B, both × 1.7.
9,14. Dorsal valve, internal mould; sample H 1052; MaMe 0060, 12. Ventral valve, anteriior view to internal mould; sample H 333;
× 2.5, × 4.0. MaMe 0151A, × 1.7.
10, 13. Dorsal valve, internal mould and details of cardinalia; sample 13. Dorsal valve, internal mould; sample H 1404; MaMe 0147, × 1.8.
H 1052; MaMe 0069B, × 1.7, × 3.0. 14. Dorsal valve, internal mould showing details of cardinalia;
11. Ventral valve, internal mould; sample H 1052; MaMe 0076, × 1.7. sample H 1404; MaMe 0146B, × 3.0.
12. Ventral valve, internal mould; sample H 1052; MaMe 0026I, × 1.7. 15, 18. Holotype, complete shell,, dorsal valve and anterior view;
15, 16. Dorsal valve, internal mould and details of cardinalia; sample sample H 1017; MaMe 0125, both × 1.7.
H 1052; MaMe 0064, × 1.7, × 3.0. 16. Ventral valve, internal mould; sample H 1090; MaMe 0148, × 1.7.
17. Ventral valve, external mould; sample H 1052; MaMe 0065, × 1.7. 17. Dorsal valve, internal mould showing muscle field; sample H
1404; MaMe 0200, × 3.0.
PLATE 9
19. Ventral valve with median sulcus in dorsal fold, probably the
Arduspirifer cf. mosellanus (SOLLE, 1953) same species, internal mould; sample H 884; MaMe 0045, × 1.7.
A g e a n d l o c a l i t i e s : Emsian; Tamesna Basin, In Rahir Tin
Amzi section (1–3, 5–8), and Tin Rerhoh section (4). Spinocyrtia sp.
A g e a n d l o c a l i t y : Givetian; Tamesna Basin, In Ateï section.
1. Small ventral valve, internal mould; sample H 1058; MaMe 163, × 1.7.
2. Ventral valve, internal mould; sample H 1058; MaMe 0165a, × 1.7. 20. Dorsal valve, internal; sample H 820; MaMe 0077, × 1.3.
3. Ventral valve, internal mould; sample H 1058; MaMe 0166b, × 1.7.
4. Dorsal valve, latex cast of exterior; sample H 1088; MaMe PLATE 11
0092B, × 2.5. Mediospirifer rerhohensis sp. n.
5. Ventral valve, latex cast of exterior; sample H 1058; MaMe A g e a n d l o c a l i t i e s : Eifelian; Tamesna Basin, Tin Rerhoh
0164A, × 1.7. section (1–6, 8, 10, 12, 13, 15–21), Oued Felaou area (7, 11,
6, 7, 9. Complete shell valve, latex cast of ventral valve exterior, 14), and In Debirene section (9).
posterior view and deatil of microornament in external mould; 1. Ventral valve, latex cast of exterior; sample H 1090; MaMe
sample H 1058; MaMe 0166A, × 1.7, × 9.0, × 1.7. 0174, × 1.7.
8. Incomplete ventral and dorsal valves, internal moulds; sample H 2. Paratype, ventral valve, latex cast of exterior; sample H 1090;
1058; MaMe 0165B, × 1.7. MaMe 0192, × 1.7.
Paraspirifer aff. cultrijugatus (ROEMER, 1844) 3. Ventral valve, latex cast of interarea; sample H 1090; MaMe
A g e a n d l o c a l i t i e s : Eifelian; Tamesna Basin, In Ateï section 0173, × 1.7.
(10, 13, 14, 17, 18), Tin Rerhoh section (11, 12), and In De- 4. Ventral valve, latex cast of interarea; sample H 1090; MaMe
birene section (15, 16, 19). 0193, × 1.7.
97
5. Dorsal valve, latex cast of exterior; sample H 1090; MaMe Rhipidothyris africana BOUCOT, MASSA et PERRY, 1983
0188, × 1.7. A g e a n d l o c a l i t i e s : Givetian; Tamesna Basin, area W to the
6, 10. Ventral valve, latex cast of exterior in ventral and anterior Air Massif (4–8, 10–12), and In Ateï section (9).
views; sample H 1090; MaMe 0170, both × 1.7. 4. Complete shells, external moulds showing pedicle foramen;
7. Incomplete ventral valve, internal mould; sample H 333; MaMe sample G 1000; MaMe 0113A (left) and MaMe 0113D, × 3.5.
0151D, × 1.7. 5, 10. Ventral valve, latex cast of external mould showing exop-
8, 15. Dorsal valve, internal mould; sample H 1090; MaMe 0189, unctation, and latex cast of four complete shells; sample
both × 1.7. G 1000; MaMe 0113C, × 9.0, × 3.5.
9. Dorsal valve, latex cast of exterior; sample H 1105; MaMe 6. Ventral valve, internal mould showing dental plates; sample
0143, × 1.7. G 1000; MaMe 0114, × 3.5.
11, 14. Ventral valve, internal mould in ventral and anterior views; 7. Dorsal valve, internal mould showing dental plates; sample
sample H 333; MaMe 0151C, both × 1.7. G 1000; MaMe 0113B, × 3.5.
12. Ventral valve, latex cast of exterior; sample H 1090; MaMe 8. Dorsal valve, internal mould showing crural plates; sample
0190, × 1.7. G 1000; MaMe 0118, × 3.5.
13, 16, 17, 19. Holotype, dorsal valve, internal mould in dorsal, 9. Bedding plane of sandstone with external and internal moulds
posterior and posterodorsal views, and detail of ctenophoridi- of Rhipidothyris africana BOUCOT et al. 1983; sample H 355,
um; sample H 1090; MaMe 0191, × 1.7, × 1.7, × 1.7, × 6.5. MaMe 0111, × 1.5.
18, 21. Dorsal valve, internal mould in anterior and dorsal views; 11. Dorsal valve of complete shell, internal mould; sample
sample H 1090; MaMe 0187, both × 1.7. G 1000; MaMe 0116, × 3.5.
20. Dorsal valve, internal mould showing muscle scars; sample H 12. Ventral valve of complete shell, internal mould; sample
1090; MaMe 0186, × 1.5. G 1000; MaMe 0117, × 3.5.
PLATE 12 Pleurothyrella cf. knodi (CLARKE, 1913)
Subcuspidella cf. subcuspidata (SCHNUR, 1851) A g e a n d l o c a l i t y : Upper Pragian; Tamesna Basin, In Rahir
A g e a n d l o c a l i t i e s : Emsian; Tamesna Basin, Tin Rerhoh Tin Amzi section.
section (1–3, 5, 6, 8), and In Rahir Tin Amzi section (4, 7). 14–16. Complete shell, internal mould in anterior, ventral and dor-
1. Ventral valve, internal mould; sample H 1088; MaMe 0094, × 2.0. sal views; sample H 1053; MaMe 0119, × 1.3.
2, 5. Ventral valve, latex cast of exterior and ventral interarea;
sample H 1088; MaMe 0171, × 2.0, × 2.0. PLATE 14
3, 6. Ventral valve, latex cast of exterior and ventral interarea; Globithyris orchas (HAVLÍČEK, 1984)
sample H 1088; MaMe 0095, × 2.0, × 2.0. A g e a n d l o c a l i t i e s : Pragian; Tamesna Basin, Anou Izileg
4. Dorsal valve, latex cast of exterior; sample H 1058; MaMe area (1–4, 6–8), and In Rahir Tin Amzi section (5).
0164B, × 2.0.
7. Dorsal valve, latex cast of exterior; sample H 1057; MaMe 1. Dorsal valve, internal mould; sample H 823; MaMe 0104, × 2.0.
0168, × 2.0. 2, 4. Complete shell valve, internal mould in ventral and anterior
8. Dorsal valve, internal mould; sample H 1088; MaMe 0096, × 2.0. views; sample H 823; MaMe 0108, × 2.0.
3, 6. Ventral valve, internal mould in lateral and ventral views;
Tenuicostella ? sp. sample H 823; MaMe 0106, × 2.0.
A g e a n d l o c a l i t i e s : Emsian to Eifelian; Tamesna Basin, In 5. Incomplete ventral valve, internal mould; sample H 1052;
Debirene section (9–12, 15), Tin Rerhoh section (13), and In MaMe 0063, × 7.5.
Rahir Tin Amzi section (14). 7, 8. Ventral valve, internal mould in ventral and posterior views;
9, 12. Ventral valve, internal mould and latex cast of exterior; sam- sample H 823; MaMe 0112, × 2.0.
ple H 1106; MaMe 0140D, both × 1.5.
Amphigenia cf. elongata (VANUXEM, 1842)
10, 15. Ventral valve, internal mould in anterior and ventral views; A g e a n d l o c a l i t i e s : Emsian; Tamesna Basin, In Rahir Tin
sample H 1106; MaMe 0091, both × 1.5. Amzi section (9, 12, 13), and Tin Rerhoh section (10, 11, 15).
11. Small ventral valve, internal mould; sample H 1106; MaMe
0142, × 1.5. 9, 12, 13. Dorsal valve, internal mould, in dorsal and posterodor-
13. Ventral valve, internal mould; sample H 1088; MaMe 0172, × 1.5. sal views and latex cast of cardinalia (12); sample H 1058;
14. Dorsal valve, internal mould; sample H 1058; MaMe 0167A, × 1.2. MaMe 0160, × 0.8, × 2.5, × 1.0.
10. Incomplete small ventral valve, internal mould; sample H
PLATE 13 1088; MaMe 0092A, × 3.5.
Rhipidothyris ? sp. 11. Incomplete small dorsal valve, internal mould; sample H
A g e a n d l o c a l i t y : Givetian; Tamesna Basin, In Ateï section 1088; MaMe 0093A, × 3.5.
(1–3, 13) 15. Incomplete ventral valve, internal mould; sample H 1088;
MaMe 0099, × 1.5.
1. Ventral valve, latex cast of exterior; sample H 355; MaMe
0019, × 2.5. Etymothyris sp.
2. Ventral valve, internal mould; sample H 1009; MaMe 0016A, × 2.5. A g e a n d l o c a l i t y : Upper Pragian; Tamesna Basin, In Rahir
3. Ventral valve, internal mould; sample H 1009; MaMe 0016B, × 2.5. Tin Amzi section.
13. Bedding plane of sandstone with external and internal moulds 14. Dorsal valve, latex cast of exterior; sample H 1052; MaMe
of Rhipidothyris ? sp; sample H 355, MaMe 0019, × 1.5. 0161, × 1.7.
98
Plate 1
5
3
6 7
99
Plate 2
1 2
3
4
8
9
12
10 11
14 15
13
16 17 18
100
Plate 3
4
1 2 3
5 6
7 8
9
11
12
10
15
14
13
16
19
18
17
20
24
21 22 23
101
Plate 4
4
1 2
3 5
6 8
10 11
12 13 14
15 16 17 18
19 20 21 22
102
Plate 5
2 3 4
1
5
6
7
12
11
13
10
15
14
16
17
18 19
20 21 22
103
Plate 6
1 2
3
4
7
6
5
8 10 11
104
Plate 7
1 2 3
4
5 6 7 8
9
10 11 12
13 14
15
16 17 18 19
20 21 22
105
Plate 8
1 2 3
4 5 6
7 8
10
11 12
15
13
14
16 17
106
Plate 9
1 2
4 5
9
7 8
10 12
11
15
13 14
16
17 18
19
107
Plate 10
1 2 3
5 6
10
8
9 11
12 14
13
15 16
17
18 19 20
108
Plate 11
3
5 6
11
10
14
12
15
13
16
18
17
20
19 21
109
Plate 12
1 2 3
6
4 5
7 8
10
13
11
12
14 15
110
Plate 13
1 2
3
4
6 7
5
11
10
14 12
13 15 16
111
Plate 14
1 2
3
4
5
10
11
12
8
9
15
13 14
112