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Collections of the Natural History Museum of Lille. VIII. — Diplostraca (Types and figured specimens)

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The Diplostraca (also known as conchostracans; Arthropoda: Crustacea: Branchiopoda: Phyllopoda) are a group that were often collected in the coal basins of northern and eastern France, during the main period of activity of the coal industry (end of 19th Century to beginning of 20th Century); others were collected in Germany, Switzerland, Africa and Russia. Hence, the palaeontological collections of Lille (Natural History Museum of Lille and University of Lille) are especially rich in specimens. This paper presents a reasoned catalogue of the types and figured specimens of the collection of the Natural History Museum of Lille, associated with a brief review of the Palaeozoic diplostracans, and some new or updated data in geology and palaeontology. Fossil Diplostraca are a rather neglected group, which appears however to be important in biostratigraphy, sequential stratigraphy, palaeoclimatology, palaeobiology, palaeoecology and palaeobiogeography during both principal stages of the Late Carboniferous (Pennsylvanian); i.e., during Moscovian (traditionally called Westphalian in western Europe) and Kasimovian times (traditionally called Stephanian).
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Ann. Soc. Géol. du Nord.
T. 22 (2e série), p. 109-118, Décembre 2015.
COLLECTIONS OF THE NATURAL HISTORY MUSEUM OF LILLE. VIII. —
DIPLOSTRACA (TYPES AND FIGURED SPECIMENS)
Les collections du Musée d'Histoire Naturelle de Lille. - VIII. — Diplostraca (Types et gurés)
by Marie HENNION (*), Jessie CUVELIER (**), Oscar F. GALLEGO (***), Thierry OUDOIRE (****)
& Daniel VACHARD (*****)
Abstract. – The Diplostraca (also known as conchostracans; Arthropoda: Crustacea: Branchiopoda: Phyllopoda) are a group
that were often collected in the coal basins of northern and eastern France, during the main period of activity of the coal
industry (end of 19th Century to beginning of 20th Century); others were collected in Germany, Switzerland, Africa and
Russia. Hence, the palaeontological collections of Lille (Natural History Museum of Lille and University of Lille) are espe-
cially rich in specimens. This paper presents a reasoned catalogue of the types and gured specimens of the collection of the
Natural History Museum of Lille, associated with a brief review of the Palaeozoic diplostracans, and some new or updated
data in geology and palaeontology. Fossil Diplostraca are a rather neglected group, which appears however to be important
in biostratigraphy, sequential stratigraphy, palaeoclimatology, palaeobiology, palaeoecology and palaeobiogeography during
both principal stages of the Late Carboniferous (Pennsylvanian); i.e., during Moscovian (traditionally called Westphalian in
western Europe) and Kasimovian times (traditionally called Stephanian).
Résumé. – Les Diplostraca (autrefois appelés conchostracés ; Arthropoda : Crustacea : Branchiopoda : Phyllopoda) sont un
groupe qui a fait l’objet de nombreuses récoltes dans les bassins houillers du nord et de l’est de la France, durant la période
d’activité maximale des mines de charbon (c’est-à-dire de la n du XIXe siècle au début du XXe) ; d’autres exemplaires ont
été recueillis en Allemagne, en Suisse, en Afrique et en Russie. Ces très nombreux spécimens ont été rassemblés dans les
collections publiques lilloises de paléontologie (Musée d’Histoire Naturelle de Lille et Université de Lille). Cet article pré-
sente un catalogue raisonné des collections de types et gurés du Musée d’Histoire Naturelle de Lille ainsi qu’une révision
des données générales sur les Diplostraca paléozoïques, et un certain nombre de données géologiques et paléontologiques
nouvelles ou actualisées. En effet, ce groupe de fossiles s’avère important en biostratigraphie, stratigraphie séquentielle,
paléoclimatologie, paléobiologie, paléoécologie et paléobiogéographie pour les deux périodes principales du Carbonifère
supérieur (Pennsylvanien) : le Moscovien (classiquement Westphalien) et le Kasimovien (classiquement Stéphanien).
Keywords. – Diplostraca, conchostracans, Carboniferous, Nord – Pas-de-Calais Basin, Saar-Lorraine Basin.
Mots-clés. – Diplostraca, conchostracés, Carbonifère, Bassin du Nord – Pas-de-Calais, Bassin sarro-lorrain.
(*) Muséum National d’Histoire Naturelle – Département Biologie des Invertébrés Marins et Malacologie (BIMM), 55 rue Buffon, F-75005 Paris
(France); marie.h2p@hotmail.fr
(**) Université de Lille – Sciences et Technologies, UFR Sciences de la Terre (SN5), UMR 8198 Evo-Eco-Paléo, F-59655 Villeneuve d’Ascq
cedex (France); Jessie.Cuvelier@univ-lille1.fr
(***) Área Paleontología (Centro de Ecología Aplicada del Litoral, Centro Cientíco Tecnológico Nordeste, Consejo Nacional de Investigaciones
Cientícas y Técnicas), Casilla de Correo 128, 3400 Corrientes (Argentina); and Geología Histórica-Micropaleontología (Facultad de Ciencias
Exactas y Naturales y Agrimensura, Universidad Nacional del Nordeste), Corrientes (Argentina); ofgallego@live.com.ar
(****) Musée d’Histoire Naturelle de Lille, 19 rue de Bruxelles, F-59000 Lille (France); toudoire@mairie-lille.fr
(*****) 1 rue des Tilleuls, F-59152 Gruson (France); Daniel.Vachard@free.fr
I. — HISTORICAL BACKGROUND
The geological and palaeontological collections of the city
of Lille (northern France) are particularly rich, insofar as they
result from the merging of two collections, that of the Natural
History Museum and that of the University of Lille, which both
were constituted as early as the 19th century (Oudoire et al.,
2014). The Faculty of Sciences of Lille was founded in August
1854, but the department of geology was created 10 years later
with the appointment of Jules Gosselet to the chair of Geology
and Mineralogy (Thiébault, 2011). On the other hand, in the
same period, the Society of Sciences, Agriculture and Arts of
Lille bequeathed its collection of natural history specimens
to the city of Lille, which made it available for the Faculty
of Sciences. The city of Lille formalized the creation of the
Museum of Geology and Mineralogy on the 18th February,
1877. This museum was rst managed by a commission headed
by professors of the university, which included Jules Gosselet
(Blieck et al., 2013, 2014).
Later, the museum and the faculty moved together to a single
new building, built in 1894. The Gosselet Museum of Geology
was inaugurated in 1902. Subsequently, the collections were
therefore enriched thanks to the investigations of the university
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professors and the eld excursions organized in the region for
the students. Charles Barrois, who was a former student of Jules
Gosselet, succeeded him as the head of the Chair of Geology
and Mineralogy and founded the Coal Museum in 1907, in
order to implement a collaboration between the mining industry
and university research. Numerous mining companies were
active in the mining area of Nord Pas-de-Calais (France)
during the rst part of the 20th century, and a great number
of samples were offered by these companies to the University
and Museum collections (Oudoire et al., 2014). That is why
the Natural History Museum and the University have a unique
collection of palaeobotanical specimens, as well as of all other
fossil forms of life encountered in the coal basin. In the 1960s,
the University’s buildings located within the city of Lille were
found too narrow and not suitable for teaching and housing
anymore. Therefore, the University moved to the outskirts of
Lille. The geology professors took away their specimens under
study, essentially fossil plants, to be received by the new campus
of Villeneuve d’Ascq. The rest of the collections (nearly 75%)
remained in the older buildings to form the Natural History
Museum, henceforth managed by the City of Lille itself. Thus
it happened that the palaeontological collection of Lille was
divided into two sets of objects (Oudoire et al., 2014).
II. — COLLECTIONS MANAGEMENT
The Natural History Museum is a multidisciplinar y museum
with 393,050 specimens including 120,000 in palaeontology
and 56,400 in geology, according to the observatory OCIM
in 2011 (Charon, 2014). The core collection encompasses
four groups of samples: regional collection (nearly all from
the geological Paris basin; 80,000 specimens), extra-regional
collection (20,000 specimens), palaeobotany (6,000 specimens)
and mineralogy (5,000 specimens). This core collection comes
from ndings of regional amateurs and scientic samplings
(Oudoire et al., 2014), like the Pierre Pruvost’s work (Pruvost,
1911, 1914, 1919) of the Carboniferous faunas, which provided
most of the diplostracan/conchostracan gured specimens of
the Museum. Other elements of the collections were provided
by Corsin (1932) in his palaeontogical guidebook on the Nord
Pas-de-Calais coal basins, and by Waterlot (1934), who studied
conchostracan faunas from the Saar-Lorraine coal basin.
Since the beginning of 21st century, the Museum began an
inventory policy with public funding; and several catalogues
have been published in the Annales de la Société Géologique
du Nord (Malvesy, 1999; Malvesy et al., 1999, 2000, 2002;
Blieck et al., 1999, 2013; Oudoire et al., 2008, 2011). In parallel,
the University proceeds with the inventory and photographs
of its collection, in the framework of the national program
Trans’Typal (http://transtypal.u-bourgogne.fr). Trans’Typal
is an online database listing the palaeontological types and
gured specimens housed in the French institutions such as
the universities and museums. This database was created in
1986 under the name Typal (Thierry, 1995), and was initially
coordinated by the University of Bourgogne (Dijon). In the near
future, it will be replaced by the great project E-Recolnat of
digitization of French taxonomic references. All the specimens
reported below are available on the website http://transtypal.u-
bourgogne.fr/ by implementing the following search criteria:
Order: “Diplostraca” / Establishment keeper: “Lille - Museum
d’Histoire Naturelle”.
III. — DIPLOSTRACA
Diplostraca, formerly named conchostracans, are small
branchiopod-phyllopod crustaceans, also called “clam shrimp”,
with a symmetrical, bivalved, chitinous carapace with growth
lines (Lecointre & Le Guyader, 2009). They differ from
the ostracods, which have a calcitic shell without preserved
growth lines (Meglitch, 1975; Vannier et al., 2003). Diplostraca
are present from the Devonian to Recent (Tasch, 1969;
Monfer ran et al., 2013). Their size varies from 1 millimetre to
a few centimetres. They inhabit temporary fresh water bodies
(Vannier et al., 2003) and consequently, provide good evidence
for lacustrine environments in sequential and biostratigraphic
analyses (Lucas & Milner, 2006). The geographical distribution
of extant Diplostraca is worldwide, except in Antarctica,
although Tasch (1987) described many fossil Diplostraca from
Antarctica. This very broad distribution can be explained,
because their eggs are time- and drought-resistant due to their
encystment, and possible, subsequent transportation by birds
and winds. These latter characters make diplostracans useful
for palaeogeographical reconstructions (Tasch, 1987).
Modern taxonomic descriptions of Diplostraca are
principally based on the anatomy of the living animal, not on
the carapace; so that, there are no cross checks between extant
and fossil specimens. On the other hand, several species were
based on only one fossil specimen, in spite of the fact that
recent papers suggest the comparison of fossil and extant taxa
(Martin & Davis, 2001; Astrop & Hegna, 2015). Molecular
and morphological studies have shown that the former
“Conchostraca” group was not monophyletic, and now this
group is part of the order Diplostraca, which include four extant
suborders, viz., Spinicaudata, Laevicaudata, Cyclestherida,
and Cladocera, and two extinct suborders, viz., Leaiina and
Estheriellina (Martin & Davis, 2001; Monferran et al., 2013).
The principal characters used to describe the fossil Diplostraca
(Fig. 1) are 1) the angle between dorsal and posterior margin;
2) the shape of the shell, which can be subovate, subcircular,
subelliptical, subquadrate or subtrapezoidal; 3) the presence
or absence and the number of growth lines on the carapace; 4)
the position of the umbo, which is placed between the median
point and the anterior end of the dorsal margin; 5) the presence
of different structures such as spines, nodes and ribs in the
umbonal region, the type of structure in the interspaces of
growth lines; and 6) the presence or absence and number of
radial ornaments, eventually of the carinae or ribs (Raymond,
1946; Kobayashi, 1954; Novozhilov, 1958a, b, c, d; Tasch, 1969;
Chen & Shen, 1985).
Fig. 1. – Sketch of a diplostracan (modied from Vannier et al., 2 003).
Fig. 1. – Schéma d’un diplostracé (modié d’après Vannier et al., 20 03).
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from 45° to almost vertical; “Voie de fond” – deepest mine
gallery); for others, we will use the proposed translations:
Assise – Formation; Faisceau – Member; Galerie – gallery;
Passée – parasequence; Puits, fosse – pit; Veine – coal seam.
4. Former numberings of samples
The Natural History Museum of Lille renumbered
specimens with the prex MGL and a number that can match
the historical collection (e.g., MGL 1886 for n°1886) or not (e.g.,
MGL 6341 for n°1884).
5. Editorial remark
We have transliterated the Russian names according to
the current phonetic rules based on the English phonetics; for
example, we used Novozhilov instead of Novojilov.
6. Systematic classication of Diplostraca
The classication followed in this work was established by one
of the authors (O.G.) using a compilation of previous established
classications: Novozhilov (1958a, b, c, d), Defretin-Lefranc
(1965, 1970), Tasch (1969, 1987), Chen & Shen (1985), Jones &
Chen (2000), and Martin & Davis (2001). Some of the specimens
mentioned in the catalogue have no clear systematic placement
because they lack the main diagnostic characters based on the
above mentioned literature. Further detailed studies in the future
Fig. 2. – Chronostratigraphic correlation in the Pennsylvanian Subsystem (modied from Heckel & Clayton, 2006).
Fig. 2. – Corrélation ch ronostratigraphique du sous-système Pennsylvanien (modié d’après Heckel & Clayton, 20 06).
IV. — PRELIMINARY DATA ABOUT THE
SYSTEMATIC CATALOGUE
1. Goals of this catalogue
This catalogue follows other publications about the
palaeontological collections of the Natural History Museum of
Lille. The main goal of a catalogue is to help further revisions
of a group of organisms, and the collection may be a good tool
for studies with modern methods like morphometric analyses,
SEM analyses or geochemical investigations.
2. Abbreviations
MGL, Musée Gosselet, Lille (Musée d’Histoire Naturelle
– Musée de Géologie); MHL, Musée Houiller, Lille (Musée
d’Histoire Naturelle – Musée de Géologie); Loc., Locality;
Lithost., Lithostratigraphy; Biost., Biostratigraphy; Chronost.,
Chronostratigraphy.
3. Adopted coal-units nomenclature
In the following list, we tried to translate, revise and re-
actualize some lithostratigraphic, geographic and toponymic
terms traditionally used in the French coal basins and/or coal
industry and mines. Names between inverted commas remain
directly used below in the text (“Bowette” – intersecting
horizontal gallery; “Dressant” – strongly dipped coal seam
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Fig. 3. – Table of stratigraphic correlations and stratigraphic distributions of the Diplostraca specimens of the Natural History Museum of Lille,
collected in the coalelds of Nord Pas-de- Calais and Saarland-Lorraine (modied from Laveine et al., 1977).
Fig. 3. – Tableau récapitulatif des concordances stratigraphiques et des extensions des spécimens de Diplostracés de la collection du Musée
d’Histoire Naturelle de Lille, récoltés dans les bassins houillers du Nord – Pas-de-Calais et de Sarre-Lorraine (modié d’après Laveine et al., 1977).
— 112 —
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could bring new information to the real systematic afnities of
these materials. The suborders Leaiina and Spinicaudata are
identied in the material analyzed in the present catalogue.
7. Revised chronostratigraphy
The chronostratigraphy is revised and re-actualized
according to the more recent classication of the Pennsylvanian
Subsystem proposed by Heckel & Clayton (2006) (Fig. 2).
8. Stratigraphic distribution of the principal species mentioned
in the catalogue
Among the 27 listed Diplostraca specimens for the Natural
History Museum, 24 come from the Nord – Pas-de-Calais
coaleld (Westphalian in age) and three from the Saarland-
Lorraine coaleld (Stephanian in age). The stratigraphic
distribution of the principal species of diplostracans is indicated
along a lithostratigraphic column of each basin (Fig. 3). This
catalogue shows the uniqueness of this collection because the
majority of the specimens come from the Nord Pas-de-Calais
coaleld, which has become inaccessible and has not been
exploited since the end of the 20th century.
V. — SYSTEMATIC CATALOGUE
Authors of taxon names are indicated in small capital
letters followed by year of authorship. The corresponding
bibliographical references are not listed here in the bibliography
section; they can be found in the papers of Novozhilov (1958a,
b, c, d), Defretin-Lefranc (1965, 1970), Tasch (1969, 1987),
Chen & Shen (1985), Jones & Chen (2000), and Martin & Davis
(2001). All other cited papers are in the bibliography section.
Phylum ARTHROPODA
von Siebold
, 1848
Subphylum CRUSTACEA
brünnich
, 1772
Class BRANCHIOPODA
latreille
, 1817
Subclass PHYLLOPODA
PreuSS
, 1951
Order DIPLOSTRACA
GerStaecker
, 1866
Suborder LEAIINA
kobayaShi
, 1972
Superfamily LEAIOIDEA
raymond
, 194 6
Family LEAIIDAE
raymond
, 194 6
Genus HEMICYCLOLEAIA
raymond
, 194 6
Hemicycloleaia baentschiana (
beyrich
, 1864)
raymond
, 194 6
Comment. The assignment of this species to the genus
Hemicycloleaia follows Percy Raymond´s proposal and is based
on Jones & Chen (2000) and the revision of material (I-02824)
housed in the National History Museum of Madrid (Spain).
MGL 510 0 -1: Right valve Leaia leidyi var. baentschiana Beyrich,
1864, illustrated as Leaia baentschi in Wate rlot, 1934, pl. VI I, g. 5.
Loc.: Mining site of Göttelborn, B.I.X. Nördl. Hauptquersch. II
Tb. S., 41 metres north of the coal seam Wachlschied (Saarland,
Germany).
Lithost.: Sarrelouis Formation.
Biost.: Leaia beds.
Chronost.: Carboniferous, Lower Stephanian.
Hemicycloleaia minima (
PruvoSt
, 1914)
raymond
, 194 6
Comment. The assignment of this species to the genus
Hemicycloleaia follows Percy Raymond´s proposal and is based
on Jones & Chen (2000).
MGL 1888: Right valve illustrated as Leaia tricarinata va r.
minima Pruvost, 1914, pl. II, g. 9; Pruvost, 1919, pl. XXV, g.
4; Corsin, 1932, pl. XXXVIII, g. 3.
Loc.: Roof of a parasequence located at 12 metres in the lateral
gallery n°3, 0.80 m upon the coal seam, Pit 10, Concession of
Lens, Group of Lens (Pas-de-Calais, France).
Lithost.: Vicoigne Formation.
Chronost.: Carboniferous, Westphalian A.
Comment. The exact location of the type locality (i.e., its height
above roof) is most probably 12 m, as inscribed on the label
associated with the sample, rather than 11 m or 2 m, as indicated
in the publications of Pruvost (1914, 1919) or Corsin (1932).
MGL 60 35 -1: Right valve illustrated as Leaia tricarinata var.
minima in Pruvost, 1914, pl. II, g. 12; 1919, pl. XXV, g. 3.
Loc.: Roof of the coal seam Nord 4, Pit Déjardin, Concession of
Aniche, Group of Douai (Nord, France).
Lithost.: Vicoigne Formation, Modeste Member.
Chronost.: Carboniferous, Westphalian A.
MGL 6035 -2: Right valve illustrated as Leaia tricarinata var.
minima in Pruvost, 1914, pl. II, g. 11; 1919, pl. XXV, g. 2;
Corsin, 1932, pl. XXXVIII, g. 2.
Loc., lithost., chronost.: As for the specimen MGL 6035-1.
MGL 6035-3: Positive impression of a left valve and fragments
of the negative impression of a right valve illustrated as Leaia
tricarinata var. minima in Pruvost, 1914, pl. II, g. 10; 1919, pl.
XXV, g. 1.
Loc., lithost., chronost.: As for the specimen MGL 6035-1.
Suborder SPINICAUDATA
linder
, 1945
Superfamily EOSESTHERIOIDEA
ZhanG
&
chen
in
ZhanG
et al., 1976
Family EUESTHERIIDAE
defretin-lefr anc
, 1965
Genus EUESTHERIA
dePéret
&
maZera n
, 1912
Euestheria dawsoni (
JoneS
, 1870)
PruvoSt
, 1919
Comment. The species dawsoni was assigned to the genus
Estheria, Pseudestheria or Euestheria according to the authors.
The latter assignment currently prevails; however, due to the
absence of a distinctive ornamentation pattern, Raymond
(1946) mentioned that the species most probably belongs to the
Limnadiidae, as it has a large umbonal region / larval carapace,
conspicuous in the Jones’ specimens.
MGL 1899 -A: Right valve illustrated as Estheria dawsoni in
Pruvost, 1919, pl. XXIV, g. 27.
Loc.: “Dressant” at 523 m in the “bowette” 1303, Pit 13,
Concession of Lens, Group of Lens (Pas-de-Calais, France).
Lithost.: Anzin Formation, Meunière Member.
Chronost.: Carboniferous, Westphalian B.
MGL 1900: Right valve illustrated as Estheria dawsoni in
Pruvost, 1919, pl. XXIV, g. 25.
Loc.: Coal seam n° 6, Pit 4, Concession of LEscarpelle, Group
of Douai (Nord, France).
Lithost., chronost.: As for the specimen MGL 1899-A.
Comment. The sample MGL 1900, relocated by us in the
collections, had the same numbering and the same locality
as in the Pruvost’s publication. However, neither one of the
four fragments of mudstone, which comprise the sample,
corresponded to the specimen illustrated in this latter
publication.
MGL 1901-1: Right valve illustrated as Estheria dawsoni in
Pruvost, 1919, pl. XXIV, g. 26.
Loc.: Coal seam Bernicourt, Pit Notre-Dame, Concession of
Aniche, Group of Douai (Nord, France).
Lithost., chronost.: As for the specimen MGL 1899-A.
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— 115 —
MGL 60 28-1: Left valve illustrated as Estheria dawsoni in
Pruvost, 1919, pl. XXIV, g. 24.
Loc.: Thirth parasequence in lower part of the Désirée coal
seam, Pit 4, Concession of Meurchin, Group of Lens (Pas-de-
Calais, France).
Lithost.: Flines Formation, Olympe Member.
Chronost.: Carboniferous, Westphalian A.
? Euestheria simoni (
PruvoSt
, 1911)
PruvoSt
, 1919
Comment. The species E. simoni, similarly to E. dawsoni, was
rst assigned to the genus Estheria or Pseudestheria, due to
the absence of a conspicuous ornamentation. Defretin-Lefranc
(1970) described then the reticular ornamentation of this
species. Based on our newly investigated specimens, we suggest
that the taxon most probably belongs to the Palaeolimnadiidae,
because of its large and smooth umbonal regions. However, a
more accurate and detailed study is necessary for modifying the
assignment, to Palaeolimnadiidae or to Euestheriidae.
MGL 1892: Group of valves (Paratype of Estheria simoni in
Pruvost, 1919, pl. XXIV, gs. 30-32; Corsin, 1932, pl. XXXIX,
gs. 1-5).
Loc.: Deepest mine gallery 2001, roof of the coal seam
Beaumont, Pit 8, Concession of Lens, Group of Lens (Pas-de-
Calais, France).
Lithost.: Bruay Formation, Dusouich Member.
Chronost.: Carboniferous, Westphalian D.
MGL 6029: Left valve (Holotype of Estheria simoni designed
by Pruvost, 1911, pl. I, gs. 4-8; Pruvost, 1919, pl. XXIV, g. 29;
Piveteau, 1953, p. 264, pl. I).
Loc.: “Bowette” 811, “voie de fond” n° 2187, roof of the coal
seam Beaumont, Pit 8, Concession of Lens, Group of Lens (Pas-
de-Calais, France).
Chronost.: Carboniferous, Westphalian C.
MGL 6030: Right valve (Paratype of Estheria simoni in
Pruvost, 1911, pl. I, g. 5).
Loc.: Roof of the coal seam Beaumont, Pit 3, Concession of
Liévin, Group of Liévin (Pas-de-Calais, France).
Chronost.: Carboniferous, Westphalian C.
MGL 6031: Left valve (Paratype of Estheria simoni in Pruvost,
1911, pl. I, g. 6).
Loc.: Roof of the coal seam Marie, Pit 9, Concession of
Courrières, Group of Hénin-Liétard (Pas-de-Calais, France).
Chronost.: As for the specimen MGL 6030.
MGL 6032: Left valve (Paratype of Estheria simoni in Pruvost,
1919, pl. XXIV, g. 31).
Loc.: Coal seam Beaumont, Concession of Lens, Group of Lens
(Pas-de-Calais, France).
Chronost.: Carboniferous, Westphalian?
MGL 60 33 -1: Left valve (Paratype of Estheria simoni in
Pruvost, 1919, pl. XXIV, g. 33).
Loc.: Roof of coal seam Beaumont, Pit 4, Concession of Liévin,
Group of Liévin (Pas-de-Calais, France).
Lithost.: Bruay Formation, Dusouich Member.
Chronost.: Carboniferous, Westphalian D.
“Estheria” rimosa (
GoldenberG
, 1877)
Comment. Raymond (1946), who has been the principal revisor
of this taxon, indicated, 1) that both species, “Estheri a” limbata
and “E.” rimosa, were erected by Goldenberg (1877), and 2)
that Pruvost and Waterlot speculated that E. rimosa was only
a form of E. limbata, because they are often found together
among the fossil assemblages. Raymond (1946) maintained
the individuality of two different species, but assigned them
to the genus Pseudestheria. Kobayashi (1954, p. 160, 165)
renamed both taxa Cornia (?) limbata and Euestheria rimosa,
respectively, but he provided no taxonomic details for justifying
these taxonomic changes. Our specimens are totally devoted
of structures in the umbonal area. That indicates that they
differ from Cornia (?) limbata, the genus assignment of which
remains however doubtful, as suggested by previous authors.
MGL 5101-1: Group of valves illustrated as Estheria
limbata Goldenberg, 1877 in Waterlot, 1934, pl. VI, gs.
14-15-15a .
Loc.: Schiffweiler (Saarland, Germany).
Lithost.: Sarrelouis Formation in Göttelborn area.
Chronost.: Carboniferous, Lower Stephanian.
Superfamily ESTHERITEOIDEA
ZhanG
&
chen
in
ZhanG
et al., 1976
Family FUSHUNOGRAPTIDAE
WanG
in
honG
et al.,
1974 Genus indet.
“Lioestheria” striata
münSter
in
GoldfuSS
, 1826
emend.
PruvoSt
, 1919
Comment. As discussed above, the taxonomy of the fossil
Spinicaudata remains questionable at different hierarchical
levels. One of these problematic taxa is the family Asmussiidae,
which is an interesting group, dened on the straight, long dorsal
margin and the strong angles at both ends and subcentral umbo.
The original diagnosis (Kobayashi, 1954, as emended by Tasch,
1969) emphasized that it showed an ornamentation ranging from
smooth, punctuate, radial lirae to alveolar. Chen & Shen (1985)
assigned this family to the superfamily Estheriteoidea, due to its
radial ornamentation, nevertheless, the type genus (Asmussia) and
its type species (A. membranacea Pacht) both display a reticulate
ornamentat ion. Furthermore, the sp ecies L. striata was included by
Defretin-Lefranc (1970) in the genus Pseudoasmussia (the name
of which was changed by Tasch (1987) into Pseudoasmussiata,
due to the homonymy with Pseudoasmussia (Novozhilov, 1954));
nevertheless, many of the species of genus, including the type
species (P. grassmücki Defretin-Lefranc, 1969), show a reticulate
ornamentation. However, as the species L. striata exhibits a radial
lirae ornamentation, its assignment to the genus Lioestheria
Depéret & Mazeran (sensu Tasch 1969, neither in the sense of
Kozur et al., 1981 or Holub & Kozur, 1981) mi g ht be ju s tie d . Th i s
possibility was already suggested by Pruvost (1919), Raymond
(1946) and Kobayashi (1954). Further studies are necessary to
understand to which genus belongs this species.
MGL 189 6 -1: Left valve illustrated as Estheria striata in
Pruvost, 1919, pl. XXIV, g. 20-20a; Corsin, 1932, pl. XXXVIII,
g. 6.
Loc.: “Bowette” 1302, second parasequence at the bottom of
the coal seam n° 11, Pit 13, Concession of Lens, Group of Lens
(Pas-de-Calais, France).
Lithost.: Vicoigne Formation, Modeste Member.
Chronost.: Carboniferous, Westphalian A.
MGL 6034-5: Right valve illustrated as Estheria striata in
Pruvost, 1919, pl. XXIV, g. 21; Corsin, 1932, pl. XXXVIII,
g. 7.
Loc.: Coal seam Saint-Charles, Pit 3, Concession of Meurchin,
Group of Lens (Pas-de-Calais, France).
Lithost.: Vicoigne Formation, Modeste Member.
Chronost.: Carboniferous, Westphalian A.
MGL 603 4-7: Left valve illustrat ed as Estheria striata in Pruvost,
1919, pl. XXIV, g. 23; Corsin, 1932, pl. XXXVIII, g. 9.
Loc., lithost., chronost.: As for the specimen 6034-5.
— 114 —
— 115 —
MGL 60 34-1 2: Left valve illustrated as Estheria striata in
Pruvost, 1919, pl. XXIV, g. 22; Corsin, 1932, pl. XXXVIII,
g. 8.
Loc., lithost., chronost.: As for the specimen 6034-5.
Superfamily VERTEXIOIDEA Kobayashi, 1954
(sensu
ZhanG
et al., 1976)
Family LIOESTHERIIDAE (
raymond
, 1946) emend.
holub
&
koZur
, 1981
Subfamily VERTEXIINAE
kobayaShi,
1954
Genus CORNIA
lyutkevich
, 1937
Cornia limbata (
GoldenberG
, 1877)
kobayaShi
, 1954
Comment. The revision of this specimen shows the presence
of nodes in the umbonal area that support Kobayashi´s (1954)
assignment of it to the genus Cornia.
MGL 5099: Group of valves illustrated as Estheria limbata
Goldenberg, 1877 in Waterlot, 1934, pl. VI, gs. 13-13a-13b.
Loc.: “Bowette” of Hirschbach at 766.50 m (Saarland,
Germany).
Lithost.: Sarrelouis Formation.
Chronost.: Carboniferous, Lower Stephanian.
Family PALAEOLIMNADIOPSEIDAE
defretin-lefr anc
, 1965
Subfamily ANOMALONEMATINAE
novoZhilov
, 1958d
Genus ANOMALONEMA
raymond
, 194 6
Anomalonema reumauxi (
PruvoSt
, 1911)
raymond
, 194 6
MGL 188 6: Right valve (Paratype of Estheriella reumauxi
Pruvost, 1911 in Pruvost, 1919, pl. XXIV, g. 34; Corsin, 1932,
pl. XXXIX, g. 6; Tasch, 1960, pl. 42, gs. 2a-2b) (Fig. 4A).
Loc.: “Bowette” 167, Coal seam Arago, Pit 1, Concession of
Liévin, Group of Liévin (Pas-de-Calais, France).
Lithost.: Bruay Formation.
Chronost.: Carboniferous, Westphalian C-D.
Comment. Defretin-Lefranc (1970) has indicated that the
valve illustrated on her pl. 13, g. 10 is identical to that of
Pruvost (1919, pl. 24, g. 34). Pruvost’s valve corresponds to
a specimen of our collections numbered MGL 1886 (Fig. 4A).
That is in contradiction to the legend of plate 13 of Defretin-
Lefranc (1970), the published number of which is 1884 (then,
re-numbered MGL 6341). Evidently, the specimen of Defretin-
Lefranc is the specimen number 1884 (= MGL 6341) (Fig. 4B)
and differs from the specimen number 1886 (= MGL 1886) (Fig.
4A). Unfortunately, the specimen number 1886 was designated
as the holotype of a new genus by Novozhilov (1958d) and
renamed Pierrepruvostia defretinae. Subsequently, Chen
& Shen (1985, p. 78-79) synonymized the genus dened by
Novozhilov (1958d) with the genus Anomalonema.
MGL 1895: Right valve (Paratype of Estheriella reumauxi in
Pruvost, 1911, pl. I, g. 3).
Loc.: Roof of the coal seam Arago, Pit 9, Concession of Lens,
Group of Lens (Pas-de-Calais, France).
Chronost.: Carboniferous, Westphalian C.
MGL 6 340 -2: Right valve (Paratype of Estheriella reumauxi in
Pruvost, 1911, pl. I, g. 2).
Loc.: Roof of the coal seam Arago, Pit 1, Concession of Liévin,
Group of Liévin (Pas-de-Calais, France).
Chronost.: Carboniferous, Westphalian C.
MGL 63 41: Right valve (Holotype of Estheriella reumauxi in
Pruvost, 1911, pl. I, gs. 1-1a-1b; Pruvost, 1919, pl. XXIV, gs.
35-35a; Corsin, 1932, pl. XXXIX, gs. 7-7a; Tasch, 1960, pl. 42,
gs. 3-4a-4b; Defretin-Lefranc, 1970, pl. XIII, g. 10) (Fig. 4B).
Loc.: Roof of the coal seam Arago, Pit 12, Concession of Lens,
Group of Lens-Liévin (Pas-de-Calais, France).
Lithost.: Bruay Formation, Dusouich Member.
Chronost.: Carboniferous, Westphalian D.
Comment. Defretin-Lefranc (1970) was mistaken about the
number of the valve; she indicated n°1886 in her text, whereas
this number is in reality 1884 (= MGL 6341). On the other hand,
Tasch (1960) suggested that this specimen was identical to the
specimen illustrated in pl. 1, g. 16 of Pruvost (1911); this is
probably another mistake, because this plate does not include 16
photos; and the identied gure is most probably 1a, and not 16.
Anomalonema pruvosti (
raymond
, 194 6)
MGL 1902-A: Right valve illustrated as Estheria dawsoni in
Pruvost, 1919, pl. XXIV, g. 28.
Loc.: Terris, Pit 6, Concession of Ostricourt, Group of Oignies
(Nord, France).
Lithost.: Anzin Formation, Meunière Member.
Chronost.: Carboniferous, Westphalian B.
MGL 1902bis-A: Right valve illustrated as Estheria dawsoni
in Pruvost, 1919, pl. XXIV, g. 28bis.
Loc.: Roof of the 0.55 m-thick coal seam, Pit 5, Concession of
Ostricourt, Group of Oignies (Nord, France).
Lithost., chronost.: As for the specimen MGL 1902-A.
We think that ve other illustrated specimens were probably
deposited in the Natural History Museum collection because
this museum was the repository of the majority of the specimens
of these publications. Up to now, these specimens have not
been re-found. These ve specimens encompass one specimen
gured by Pruvost in 1911 (Estheria simoni, pl. I, g. 7), two
specimens gured by Pruvost in 1914 (Leaia tricarinata, pl. II,
gs. 4-5; pl. II, gs. 6-7) and two specimens gured by Corsin
in 1932 (as Estheria mathieui, pl. XXXIX, gs. 8-9; and Leaia
tricarinata, pl. XXXVIII, g. 1).
VI. — DISCUSSION AND CONCLUSION
This catalogue allows us to investigate the validity of
many of the mentioned species. The investigated collection
is constituted by nine species included in two suborders, four
superfamilies, ve families and ve genera; among them, we
revised two species of Anomalonema, one species of Cornia,
one species of “Estheria”, two species of Euestheria, two
species of Hemicycloleaia, and a species of “Lioestheria”.
Our numerous collections, from different stratigraphic and
geographical localities, demonstrate that several species,
e.g., Hemicycloleaia baentschiana and Euestheria simoni,
provide additional possibilities to develop detailed population
(autoecological) studies, for dening intraspecic variations. On
the other hand, as these type localities are closely related with
the coal seams, the use of the Diplostraca in the parasequences
provides new tools for the correlation and interpretation of the
beari ng sequences. This investigation also led to the re-discovery
of Diplostraca with internal structures interpreted as “eggs” by
Pruvost in 1919 (see MGL 1896-1 (Fig. 4C), MGL 6034-7 (Fig.
4D)). Moreover, our collections have yielded more specimens
with “eggs”, not yet illustrated, which might give additional
informations about these types of enigmatic structures. It is
completely possible that this collection is still concealing several
specimens with other well-preserved soft parts (e.g., legs,
claspers, furcas, shell gland, muscle scars and digestive tract; the
previous global records of which were summarized by Zhang et
al., 1990 and Shen & Schram, 2014). Fossil-Lagerstätten with
well-preserved Diplostraca are known in Ireland (Orr & Briggs,
1999), from localities, the geology of which differs from that of
our outcrops and their specimens with possible eggs.
— 116 —
— 117 —
It must be noted that this catalogue only inventories a
small portion of the Diplostraca of the collection, i.e., the types
and illustrated specimens. Many of the specimens have been
collected by the palaeontologists of Lille during the 20th century
and could be useful in the study of morphological variability via
morphometric tools. Diplostraca have been used as tools for the
correlation and interpretation of the fossil-bearing sequences
since the beginning of the mining industry. There is a large
palaeontological collection of different groups from the Nord
Pas-de-Calais coaleld, with well-known type localities in the
mine galleries. This large collection could permit further new
biostratigraphical, palaeoecological and palaeoenvironmental
investigations of the Pennsylvanian Diplostraca.
Aknowledgements. — We were helped during our
research by Mrs E. Locatelli (UMR 8198 Evo-Eco-Paleo)
for the bibliography and by Mr A. Blieck for advice about
the presentation of the catalogue. O.F. Gallego´s research is
supported by the Universidad Nacional del Nordest e (Cor rientes,
Argentina) and the Consejo Nacional de Investigaciones
Cientícas y Tecnológicas (CONICET, Argentina). We
thank all those who participated in the collection inventory
at the museum for several years. The reviews of Drs Astrop
and Monferran were very useful as well as the remarks of
Dr S. Lucas about our English.
Fig. 4. - A - Anomalonema reumauxi, MGL 1886 (former number MHL 1886), Westphalian C-D. B - Anomalonema reumauxi, MGL 6341
(former number MHL 1884). Westphalian D. C - “Lioestheriastriata with internal structures interpreted by Pruvost as “eggs”, MGL 1896-
1, Westphalian A. D - “Lioestheriastriata with internal structures interpreted by Pruvost as “eggs”, MGL 6034-7. Westphalian A. Scale bar
represents 5 mm.
Fig. 4. - A - Anomalonema reumauxi, MGL 1886 (ancien numéro MHL 1886), Westphalien C-D. B - Anomalonema reumauxi, MGL 6341
(ancien numéro MHL 1884). Westphalien D. C - “Lioestheriastriata avec des structures internes interprétées par Pruvost com me des “oeufs”,
MGL 1896 -1, Westphalien A. D - “Lioestheriastriata avec des structures internes interprétées pa r Pruvost comme des “oeufs”, MGL 6034-7.
Westphalien A. L’échelle représente 5 mm.
— 116 —
— 117 —
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— 118 —
ANNEX: SYSTEMATIC INDEX
Anomalonema pruvosti (
raymond
, 1946) p. 115
Anomalonema reumauxi (
PruvoSt
, 1911)
raymond
, 1946 p. 115
Cornia limbata (
GoldenberG
, 1877)
kobayaShi
, 1954 p. 115
Estheria dawsoni
JoneS
, 1870 p. 113, 114, 115
Estheria limbata
GoldenberG
, 1877 p. 114, 115
Estheria simoni
PruvoSt
, 1911 p. 114
Estheria striata
münSter
, 1826 p. 114, 115
“Estheria” rimosa (
GoldenberG
, 1877) p. 114
Estheriella reumauxi
PruvoSt
, 1911 p. 115
Euestheria dawsoni (
JoneS
, 1870)
PruvoSt
, 1919 p. 113
? Euestheria simoni (
PruvoSt
, 1911)
PruvoSt
, 1919 p. 114
Hemicycloleaia baentschiana
(
beyrich
, 1864)
raymond
, 1946 p. 113
Hemicycloleaia minima (
PruvoSt
, 1914)
raymond
, 194 6 p. 113
Leaia baentschi
beyrich, 1864
p. 113
Leaia leidyi var. baentschiana
beyrich
, 1864 p. 113
Leaia tricarinata var. minima
PruvoSt
, 1914 p. 113
“Lioestheria” striata
münSter
in
GoldfuSS
, 1826
PruvoSt
, 1919 p. 114
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Chapter
Biostratigraphic and taxonomic studies were carried out to determine whether the fossil conchostracan record of the southern continents contained credible evidence of nonmarine dispersal between them during portions of Paleozoic and Mesozoic time. Fossil collections were made toward that end in the following: Africa, Cave Sandstone, Triassic of the Republic of South Africa and Lesotho; Australia, Newcastle Coal Measures, Permian (Tartarian); Antarctica, Ohio Range, Permian, and Queen Alexandra Range and southern Victoria Land, Lower Jurassic; India, Raniganj Formation, Upper Permian, Panchet Formation, Triassic, and Kot Formation, Lower Jurassic; South America, Brazil, Rio do Rasto Formation, Upper Permian. Other conchostracan taxa were contributed and/or loaned by colleagues or museums, or came from the writer's collections from Africa, Antarctica, Australia, India, and South America. These included conchostracan fossils from Morocco and Algeria, Carboniferous and Cretaceous; Angola, Triassic; Devonian and Jurassic of southern and northern Victoria Land, Antarctica, respectively; upper Paleozoic and Triassic of western and eastern Australian basins; the Cretaceous of Victoria, Australia, and the Tasmanian Triassic; the Mesozoic of Brazil, Chile, Argentina, Bolivia, Colombia, and Venezuela; and conchostracan-bearing cores from the Triassic of India. For all conchostracan taxa studied (where preservation permitted) a standardized series of measurements and ratios were used to facilitate inter- and intra-continental comparisons between species. Mitchell's Australian conchostracan types are here revised and refigured, as are T. R. Jones conchostracan types from the southern continents (exclusive of Antarctica), as well as other taxa described in the older and modern literature.
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Conchostracans with well-preserved soft parts were found from the Middle Jurassic Jiulongshan formation at Zhouyingzi of Luanping, northern Hebei. Four types of preserved patterns in the collections show the carapace valve, carapace valve with soft parts, soft parts alone and post-larvae respectively. The cephalon, mandibula, antennae, pinnate setae, claspers, eggs, appendages, digestive tube, caudal segment (telson) and caudal furca are clearly visible. All the specimens are considered to represent a new species, Euestheria luanpingensis sp. nov., which occurs in association with fossil insects and plant remains. As compared with living concostracans, the new species should be referred to the suborder Spinicaudata. In consideration of the similarities in the cephalon and telson between the living and fossil materials, it is possible that the new species represents an ancestral form of living family Cyzicidae. Following the study of the higher classification for the living and fossil conchostracans, a unified scheme proposed here comprises 3 orders: Laeviscaudata Linder, 1945; Spinicaudata Linder, 1945; and Leaiina Kobayashi, 1972. Based on the sedimentary characters, the occurrence of minor kaolinite and the associated insects, the waters in which the conchostracans lived seemed to be a small, shallow and quiet pond in the Jurassic time. It might have existed for at least one month or more, and the new species could reproduce from generation to generation. Besides, in the waters there were no carnivorous and saprophagous animals.