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New Early Cretaceous clam shrimps (Spinicaudata) from uppermost

Bouhedma Formation of northern Chotts range, southern Tunisia: Taxonomy,
stratigraphy and palaeoenvironmental implic...

Article  in  Cretaceous Research · April 2017

DOI: 10.1016/j.cretres.2016.12.014

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 Cretaceous Research 72 (2017) 124e133

Contents lists available at ScienceDirect

Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

New Early Cretaceous clam shrimps (Spinicaudata) from uppermost

Bouhedma Formation of northern Chotts range, southern Tunisia:
Taxonomy, stratigraphy and palaeoenvironmental implications
Gang Li a, b, *, Kamel Boukhalfa c, d, Xiao Teng a, Mohamed Soussi d, Walid Ben Ali d,
Mohamed Ouaja e, Yassine Houla f
a
State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, East Beijing Road
39, 210008, Nanjing City, Jiangsu Province, PR China
b
University of Chinese Academy of Sciences, Beijing, 100049, PR China
c
University of Carthage, Faculty of Sciences of Bizerte, 7021, Jarzouna, Tunisia
d
University of Tunis El Manar, Faculty of Sciences of Tunis, UR11ES15, 2092, Tunis, Tunisia
e
University of Gabes, Faculty of Sciences of Gabes, 6072, Gabes, Tunisia
f
Geological Survey, National Office of Mines, 1080, Tunis, Tunisia

a r t i c l e i n f o a b s t r a c t

Article history: The fossil clam shrimp genus Ordosestheria was originally described from the Lower Cretaceous (Aptian)
Received 24 September 2016 Jingchuan Formation in Inner Mongolia, northwestern China. The species taxonomically described in this
Received in revised form paper as Ordosestheria chottsensis sp. nov. was recovered from the uppermost Bouhedma Formation in
1 December 2016
the northern Chotts range of southern Tunisia. The early Barremian occurrence of the genus in Tunisia
Accepted in revised form 20 December 2016
Available online 27 December 2016
suggests that ordosestheriids have most likely originated in North Africa and then migrated to East Asia
in the Aptian. Moreover, the occurrence of fossil spinicaudatans in two horizons respectively from the
Bouhedma and Sidi Aïch formations indicates the interaction of freshwater and marine conditions that
Keywords:
Fossil clam shrimps
characterized the northern African margin as part of widespread complex paralic environments over the
Palaeoenvironment Chotts and Gafsa areas during the Barremian.
Lower Cretaceous © 2016 Elsevier Ltd. All rights reserved.
Bouhedma and Sidi Aïch formations
Northern Chotts range
Southern Tunisia

1. Introduction
The Lower Cretaceous of North Africa is a particularly extensive
example of an Early Cretaceous terrestrial to coastal-shallow ma-
rine continental margin wedge that was subsequently flooded
during the Cenomanian transgression. The predominantly silici-
clastic Lower Cretaceous deposits, which are often referred to as the
“Continental Intercalaire” (CI) and Nubian Sandstone (Kilian, 1931;
Lefranc and Guiraud, 1990; Busson and Corne
e, 1991), extend from
Morocco to Egypt across the Saharan Africa and were deposited
along the southern shoreline of the Neotethyan Ocean (Kogbe and
Burollet, 1990; Lefranc and Guiraud, 1990; Guiraud, 1998; Guiraud
* Corresponding author. State Key Laboratory of Palaeobiology and Stratigraphy,
Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, East
Beijing Road 39, 210008, Nanjing City, Jiangsu Province, PR China.
E-mail address: gangli@nigpas.ac.cn (G. Li).
et al., 2005; Gallala et al., 2009; Aloui et al., 2012; Fanti et al.,
2012, 2014). Early interest in the CI of North Africa was driven by
searching for dinosaurs (Lapparent, 1951, 1960; Lapparent de Broin
and Taquet, 1966; Lapparent de Broin, 2000, 2002; Cuny et al., 2004,
2010; Fanti et al., 2012, 2014; Contessi and Fanti, 2012a, b; Contessi,
2013) whose evolution chart the breakup of Gondwana and the
progressive isolation of the African continent (Benton et al., 2000;
Anderson et al., 2007; Bodin et al., 2010; Raulin et al., 2011). The
Lower Cretaceous CI of North Africa is also of an important socio-
economic importance as one of the global largest freshwater
aquifer systems in one of the most arid regions (Castany, 1981;
Edmunds et al., 2003; Guendouz and Michelot, 2006; Elliot et al.,
2014; Newell et al., 2015).
The HauterivianeBarremian Bouhedma and Sidi Aïch forma-
tions, subject of this study, are part of the important Cretaceous
dominantly siliciclastic sequences (Fig. 1). The Hauterivianelower
Barremian Bouhedma Formation comprises several thick sandstone

http://dx.doi.org/10.1016/j.cretres.2016.12.014
0195-6671/© 2016 Elsevier Ltd. All rights reserved.
 G. Li et al. / Cretaceous Research 72 (2017) 124e133
Fig. 1. Schematic Lower Cretaceous stratigraphy of Tunisia from a southern, landward, position toward the marine north. The lithostratigraphical chart shows that the Bouhedma
Formation is intercalated between two main lower Cretaceous detrital deposits of the Continental Intercalaire (CI).
beds intercalated with carbonate and evaporites, while the Sidi
Aïch Formation is dominated by sandstone deposits, constituting
the most important detrital discharge recorded during the late
Barremian in central Tunisia.
The Bouhedma Formation was first defined by Burollet (1956) at
the type locality Jebel Bouhedma in southern Atlas Tunisia (the
Gafsa basin). In the study area, about 70 km to the south of Jebel
Bouhedma, this formation is made up of a very thick succession (up
to 900 m thickness) of sabkha to coastal deposits including bio-
clastic limestones, laminated dolomites, variegated shales, anhy-
drite, gypsum and fine grained sandstones (Abdeljaouad and
Zargouni, 1981). In central Tunisia, the Bouhedma Formation was
assigned to the Hauterivianelower Barremian based on ostracod
assemblages (Damotte et al., 1987; Chakhma et al., 1990; Damotte,
1990), and the overlying Sidi Aïch Formation is dominated by
channel-scoured base lags (rip up clasts) passing upward to fine- to
medium-grained sandstones covered in turn by thin claystone and
red palaeosol intercalations of flood plain setting. It is considered as
barren of fossil and has been speculated as being Barremian in age
(Ben Ferjani et al., 1990; Gallala et al., 2009; Aloui et al., 2012). The
depositional environments of this succession, organized in thick
fining upward sequences, showing important lateral facies varia-
tion from southern to northern Tunisia have been differently
interpreted, including coastal marine (M'Rabet, 1981), fluvial
(Abdeljaouad, 1983), deltaic fan (Lefranc and Guiraud, 1990),
“estran transgressif” (Ouaja et al., 2011) or semi-continental and
fluviomarine (Gallala et al., 2009; Aloui et al., 2012). Our previous
work (Boukhalfa et al., 2015) on this Formation cropping out at the
Zimlet El-Beidha Anticline Structure (ZBAS) in the northern Chotts
range, southern Tunisia, allowed identifying clam shrimp rich ho-
rizon in the middle part of the outcrop and to attribute this clam
shrimp (Cratostracus? tunisiaensis) fauna to the late Barremian. Our
recent investigation of the uppermost part of the Bouhedma For-
mation from the same section (Khanguet Aïcha) offers a new
finding of clam shrimps from a thin green shally horizon resting on
a bioclastic dolomite bed yielding well preserved dinosaur tracks
(Hamed et al., 2014). The present paper is a contribution to palae-
ontological and biostratigraphical analyses of the newly recovered
clam shrimp fauna in the uppermost Bouhedma Formation.
Moreover, this new fossil finding provides opportunities to describe
the taxonomy of the clam shrimps and to clarify the
125
palaeoenvironment context of the fossil-bearing horizons recorded
in both Bouhedma and Sidi Aïch formations from the Chotts
domain. Stratigraphic regional correlation will be attempted with
the Barremian deposits of central Tunisia where have been also
characterized for the first time new marker beds that can be
considered as correlative isochronous events.
2. Previous work and geological setting
The Lower Cretaceous deposits cropping out in the Chotts area
(northern and southern Chotts ranges) (Fig. 2A) have been studied
by numerous workers since 1980's. These studies have enhanced
our understanding on the stratigraphy (Ben Youssef and Peyberne s,
1986; Boukhalfa et al., 2015), sedimentology (M'Rabet, 1981;
Abdeljaouad, 1983; Aloui et al., 2012) and regional tectonic evolu-
tion (Zargouni et al., 1985; Fakraoui, 1990; Louhaïchi and Tlig, 1993;
Gharbi et al., 2013) of the area.
The Zimlet El-Beidha Anticline Structure (ZBAS) constitutes the
eastern part of the EeW trending northern Chotts range (Fig. 2A)
which is oriented N60 to N30 and limited in the SW and NE by two
major faults oriented EeW and NeS respectively (Louhaïchi and
Tlig, 1993). The lithostratigraphic successions of the ZBAS range
from Hauterivian to Miocene (M'Rabet, 1981; Abdeljaouad, 1983;
Ben Youssef and Peyberne s, 1986; Gharbi et al., 2013) (Fig. 2B).
The Bouhedma Formation cropping out in the ZBAS is charac-
terized by a thick (up to 900 m), well exposed carbonate-evaporite
succession with thin sandstone intercalations extending over 10 km
(Fig. 2B). The overlying Sidi Aïch Formation is 120 m thick and is
composed of sandstone deposits organized in thick fining upward
sequences. Each sequence is dominated by fine-grained sandstones
showing channelized base lags overlain by thin claystone and red
palaeosol intercalations (Fig. 3A). Two conchostracan bearing shally
horizons are discovered respectively in the uppermost part (BH,
BU) of the Bouhedma Formation and the middle part (SAH, SAU3) of
the Sidi Aïch Formation (Figs. 2 and 3). The fossil contents of the
middle part (SAH, SAU3) shally horizon of the Sidi Aïch Formation
has been the subject of our previous work (Boukhalfa et al., 2015),
while the clam shrimp bearing shally horizon of the uppermost
Bouhedma Formation constitutes the aims of the present work
(Fig. 3). This paper deals with the taxonomy of the clam shrimps,
the palaeoenvironments and regional stratigraphic relationship
126 G. Li et al. / Cretaceous Research 72 (2017) 124e133

Fig. 2. A. Digital Tunisian map showing the position of the Chotts Ranges (Northern and Southern Chotts Ranges). The dashed-line box shows the study area and the location of the
study section. B. Geological map (Abdeljaouad, 1983; Louhaïchi and Tlig, 1993; Gharbi et al., 2013) showing the main lithostratigraphic formations cropping out in the ZBAS and
particularly the location of the Khanguet Aïcha section. C. Close up of the well exposed Lower Cretaceous Bouhedma, Sidi Aïch and Orbata formations in the Khanguet Aïcha locality,
showing the stratigraphic position of the conchostracan-rich horizon (H1) in the uppermost Bouhedma Formation.

review of both Bouhedma and Sidi Aïch formations of Chotts and
central Tunisia based on previous studies (Damotte et al., 1987;
Chakhma et al., 1990; Boukhalfa et al., 2015) and new biostrati-
graphic data recovered in both Bouhedma and Sidi Aïch formations
of central Tunisia and the Chotts area.
3. Material and methods
The well-preserved clam shrimp specimens examined here are
fossilized carapaces from a thin greenish shally horizon in the up-
permost Bouhedma Formation. There are no soft parts recovered.
The figured specimens are deposited in the collection of the
Nanjing Institute of Geology and Palaeontology, Chinese Academy
of Sciences (NIGPCAS). Further specimens are deposited in the
collection of the Research Unit UR 11ES15 (Department of Geology,
Faculty of Sciences of Tunis). Here, the authors have relied on ex-
amination of specimens using a LEO 1530 VP scanning electron
microscope (SEM), and a Zeiss V20 Stereomicroscope.
4. Stratigraphy of Bouhedma and Sidi Aïch formations
The age assignment for the Bouhedma and Sidi Aïch formations
is controversy. Indeed, M'Rabet (1981) assigned the Bouhedma
Formation at Jebel Sidi Aïch (Gafsa Basin) to the Barre-
mianeBedoulian. However, Damotte et al. (1987) have assigned the
fossiliferous (ostracods and echinoids) claystones, considered to be
the equivalent lower part of the Bouhedma Formation at Jebel
Mrhila (central Tunisia), to the upper Hauterivian, and attributed it
to the Mrhila Formation at Jebel Mrhila (central Tunisia) (Fig. 1).
Chakhma et al. (1990) assigned the Bouhedma Formation at the Bir
El Hafay locality (central Tunisia) to the Hauterivianeupper Bar-
remian based on the age assignment of the ostracod and echinoid
faunal assemblages (F3: Veeniacythereis ghabounensis ghabounensis,
V. ghabounensis insueta, and V. ghabounensis variesculpta).
On the other hand, the Sidi Aïch Formation, described in the
type locality at Jebel Sidi Aïch (Gafsa Basin) as an unfossiliferous
sandy succession, has been attributed to the BarremianeBedoulian
(M'Rabet, 1981). However, Chakhma et al. (1990) assigned the Sidi
Aïch Formation at Bir El Hafay to the upper Barremian based on the
fossil evidence from ostracods and echinoids (F5: Veeniacythereis cf.
ghabounensis ghabounensis, Bairdia sp., and Cythererella sp.).
The Bouhedma Formation at the type locality is described
(Burollet, 1956; M'Rabet, 1981) as a thick succession (150e750 m)
of limestones, dolomites, evaporites and sandstones. Its uppermost
part is characterized in central Tunisia by the presence of a tran-
sitional interval composed of two thick bioclastic carbonate marker
beds (labelled as Mb1 and Mb2 in Figs. 4 and 5) intercalated by
sandstones and claystones (Damotte et al., 1987; Chakhma et al.,
1990). This transitional interval is also cropping out at the ZBAS
area, where it consists of a thick (20 m) interval dominated by fine-
grained sandstones comprising at the base and top two thick
(1e2 m) red carbonate beds (Mb1 and Mb2) (Fig. 3). In the previous
works (Abdeljaouad, 1983; Gharbi et al., 2013; Boukhalfa et al.,
2015) this transitional interval and associated carbonate marker
 G. Li et al. / Cretaceous Research 72 (2017) 124e133
Fig. 3. Lithostratigraphic column of the Barremian uppermost Bouhedma and Sidi Aïch
formations in the Khanguet Aïcha locality, showing five lithostratigraphic units with
two conchostracan-bearing horizons (H1 and H2) located at the base of the BHU1 and
SAU3, respectively.
beds were included within the Bouhedma Formation. Our detailed
study of this transitional interval confirmed this subdivision
scheme and allowed the finding of an Ordosestheria chottsensis sp.
nov. bearing horizon (H1) just above the top of the bioclastic
127
dolomite bed (Mb1) (Fig. 3). Although the genus Ordosestheria was
originally described from the non-marine Aptian Zhidan Formation
in Inner Mongolia of northeastern China. O. chottsensis sp. nov. is
the second described species of the genus, whose geological range
is not fully documented yet. We follow the early Barremian age
assignment for the uppermost part of the Bouhedma Formation
based on previous attribution (Chakhma et al., 1990) (Fig. 3).
The reinvestigation of the studied sections allowed the identi-
fication of new stratigraphic fossiliferous marker beds facilitating
the correlation of the uppermost part of the Bouhedma Formation
and the overlying Sidi Aïch Formation of the Chotts domain with
those equivalents cropping out in central Tunisia (Jebel Bir El Hafay
(JBH); Jebel Sidi Aïch (JSA) and Jebel Majoura (JM) outcrops)
(Figs. 2A, 4 and 5). Indeed, the fossiliferous clayey horizon (H1)
overlying the bioclastic Mb1 is recorded in both Chotts (Figs. 4A and
4B) and in central Tunisia (Fig. 4F). In the ZBAS area this horizon
yields a freshwater conchostracan fauna (F1: Ordosestheria chott-
sensis sp. nov.) (Figs. 6 and 7) while its equivalent in central Tunisia
(Jebel Majoura) shows a coastal marine fauna assemblage (F2:
Mytilus, bivalves, gastropods…) (Fig. 4G). The overlying marker bed
Mb2 at the top of the Bouhedma Formation in the Khanguet Aïcha
studied section is a thick (1.5 m) red dolocrete bed very bioturbated
at the base and rippled in the top with probably root traces (Figs. 4A
and 4C), albeit its equivalent in central Tunisia (Jebel Majoura)
(Fig. 4F) is composed of bioclastic and oolitic dolomite beds (Fig. 5)
(M'Rabet, 1981).
On the other hand, two additional marker beds (Mb3 and H2)
have been recorded in the Sidi Aïch Formation (Figs. 4 and 5). The
Mb3 marker bed, situated just few centimeters above the Mb2 (the
top of the Bouhedma Formation), shows lenticular coarse grained
sandstone encompassing at the base well preserved bone frag-
ments (Fig. 4D) in the ZBAS area and bone, tooth and bivalve as-
semblages at the Jebel Majoura outcrop (Fig. 4H). The fossiliferous
horizon H2 (Fig. 4E) (20 m thick lenticular claystone interval),
yielding faunal and floral assemblages (Boukhalfa et al., 2015), is
recorded only in the middle part of the Sidi Aïch Formation at the
Khanguet Aïcha outcrop (Figs. 4E and 5). Both in central Tunisia and
the Chotts area, the Sidi Aïch Formation is capped by the wide-
spread bioclastic carbonate bar of the Aptian Orbata Formation
(Fig. 5).
Although the distant (86 km) correlative Jebel Majoura-
Khanguet Aïcha sections (Fig. 5) cropping out respectively in cen-
tral Atlas Tunisia and the Chotts area (Fig. 2A), the Bouhedma and
Sidi Aïch formations show several marker beds (Mb1, Mb2 and
Mb3) and fossiliferous horizons (H1 and H2) which could be
considered as correlative isochronous events at the basin scale
(Fig. 5). In addition, the occurrence of a freshwater clam shrimp
fauna, dinosaur tracks, red dolocrete with root traces and palaeosol
levels recorded in the Chotts area indicate a coastal proximal
deposition under continental influences (emersion). However to
the North, in central Tunisia, the fauna assemblage (Mytilus, other
bivalves, teeth and others) indicates a deposition under coastal
environment influenced by tidal marine processes.
5. Palaeoecology and palaeoenvironmental significance
Recent clam shrimps can provide important ecological and
environmental information that may contribute to a thorough
understanding of the palaeoecology and palaeoenvironment
context of the fossil bearing horizons (Li, 2004; Li et al., 2009a,
2014, 2016; Li and Matsuoka, 2012; Gue
riau et al., 2016; Teng
et al., 2016). Clam shrimps are crustaceans (Class Branchiopoda)
with bivalved carapaces that completely enclose the body and
usually bear concentric growth bands in spinicaudatans (Tasch,
1969, 1987; Chen and Shen, 1977, 1982, 1983, 1985; Cohen, 2003;
128 G. Li et al. / Cretaceous Research 72 (2017) 124e133

Fig. 4. A. The Bouhedma and Sidi Aïch formations cropping out in the ZBAS (AeE; Chotts area) showing the main marker beds (Mb1, Mb2, Mb3) and fossiliferous horizons (H1, H2).
B. Bioclastic dolomite marker bed (Mb1) overlain by the conchostracan clayey horizon (H1). C. Red dolocrete marker bed (Mb2) at the top of the Bouhedma Formation showing
bioturbations and root traces. D. Mb2 overlaid by the sandstone with bone and silicified wood fragments. E. Conchostracan clayey horizon (H2) recorded in the middle part of the
Sidi Aïch Formation. F. The Bouhedma and Sidi Aïch formations cropping out at Jebel Majoura (FeH; Central Tunisia) showing the main marker beds (Mb1, Mb2, Mb3) and
fossiliferous horizon (H1). G. Coastal marine fauna recorded in the clayey horizon (H1) just above the bioclastic dolomite marker bed (Mb1). H. Main fossil fragments found in the
sandstone marker bed (Mb3). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

Vannier et al., 2003; Zhou et al., 2003; Li and Batten, 2004a, b,
2005; Li et al., 2007, 2009b, c, 2010; Astrop and Hegna, 2015).
Recent clam shrimps mostly inhabit freshwater oxygenated small
temporary or permanent inland ponds, flood-plain pools and
margins of large lakes, but they can also occur in more saline en-

n,
vironments such as large playa lakes and coastal salt flats (Gisle
1936; Tasch and Zimmerman, 1961; Rzoska, 1961; Webb, 1979;
Frank, 1988; Kozur and Mock, 1993; Lana and Carvalho, 2002;
 G. Li et al. / Cretaceous Research 72 (2017) 124e133 129

Fig. 5. Stratigraphic correlative scheme showing the main marker beds and fossiliferous horizons recorded in both Bouhedma and Sidi Aïch formations across the Chotts-central
Tunisia area based on previous and new biostratigraphic data. F1: conchostracan fauna (Ordosestheria chottsensis sp. nov.). F2: Mytilus sp., bivalves, gastropods fauna. F3: ostracods
and echinoids fauna (Veeniacythereis cf. ghabounensis ghabounensis, V. ghabounensis variescupta, V. ghabounensis insueta; Bairdia sp., and Cythererella sp.) (Chakhma et al., 1990). F4:
conchostracan fauna (Cratostracus? tunisiaensis) (Boukhalfa et al., 2015). F5: ostracods and echinoids fauna (Veeniacythereis cf. ghabounensis ghabounensis, Bairdia sp., and
Cythererella sp.) (Chakhma et al., 1990).

Rohn et al., 2005). Some species are slow swimmers whereas others in the same environment. Indeed, we can state that dense vegeta-
crawl on, or burrow in, bottom sediments for relatively long periods tion protects the clam shrimps from active predators. Hence, it is
for feeding. possible to infer that there were coastal perennial vegetated ponds/
The faunal assemblages recorded in two fossiliferous horizons in oxbow lakes around the study area during the Barremian.
the uppermost Bouhedma Formation (H1) and the middle part (H2)
of the Sidi Aïch Formation (Boukhalfa et al., 2015) in the Khanguet
Aïcha section (northern Chotts range) (Fig. 3) allow us to clarify the
6. Systematic palaeontology
depositional conditions of the uppermost Bouhedma and Sidi Aïch
formations. Indeed, the thin clayey layer (H1) overlying the bio-
Order: Diplostraca Gerstaecker, 1866
clastic dolomite bed with dinosaur tracks (Mb1) yielded only a clam
Suborder: Spinicaudata Linder, 1945
shrimp fauna (Ordosestheria). This clam shrimp genus, previously
Superfamily: Estheriteoidea Zhang and Chen, in Zhang et al., 1976
recovered in northern China, could be temporarily interpreted that
Family: Fushunograptidae Wang, in Hong et al., 1974
it inhabits a freshwater palaeoenvironment. However, the fossilif-
erous horizon (H2) recorded in the middle part of the Sidi Aïch Genus Ordosestheria Wang, 1984
Formation yields an association of both animal (clam shrimps,
1984 Ordosestheria Wang, p. 733.
fishes, shrimps) and plant fossils (Boukhalfa et al., 2015). The as-
sociation of fossil clam shrimps with marine faunas could be Type species: Ordosestheria wujiamiaoensis Wang, 1984 from the
attributed to the existence of temporary pools close to ancient Lower Cretaceous Jingchuan Formation at Wujiamiao, Hanggin
fluctuating shorelines or lagoons, so that episodic invasion of Banner, Ih Ju League, Inner Mongolia, China.
seawater over such areas could have mixed the faunas (Tasch and
Remarks. The original description of the genus Ordosestheria was
Zimmerman, 1961). Other possibilities include the dispersal of
based on external moulds. Thus, the originally described nodules
clam shrimp eggs to nearshore marine or estuarine areas, and
along the lower margin of growth bands should be small pits in the
torrential flooding that covered pools temporarily, forming a
ventral part of the carapace. On the other hand, the pores on
widespread sheet of water that drained towards the sea, mingling
growth lines and the serrated lower margins have not been
the faunas (Lana and Carvalho, 2002). The presence of fish and
described in the original description.
plants in the H2 of the Sidi Aïch Formation (Fig. 3) is interesting
Occurrence. Lower Cretaceous Jingchuan Formation, Inner
because, many modern clam shrimp species are assumed not to
Mongolia, China; lower Barremian uppermost Bouhedma Forma-
persist in environments with a resident fish fauna or are very rare
tion, southern Tunisia.
in permanent water bodies that contain fishes because they are
easy prey for active predators (Olsen, 1984; Rohn et al., 2005). Ordosestheria chottsensis sp. nov.
However, in our study case clam shrimps, fishes and plants coexist (Figs. 6 and 7).
130 G. Li et al. / Cretaceous Research 72 (2017) 124e133

Fig. 6. Ordosestheria chottsensis sp. nov. All are SEM images, except for AeC (light microscopy images). A. NIGPCAS 164814, holotype, a right vavle, male (?). B. NIGPCAS 164815, a
right valve, female (?). C. NIGPCAS 164816, a left valve, female (?). D. NIGPCAS 164817, a broken left valve. E. small polygonal reticulation on growth band near the umbo of the
specimen in Fig. 6A. F. polygonal small reticulation on growth band near the umbo of the specimen in Fig. 6B. G. polygonal small reticulation on growth band near the umbo of the
specimen in Fig. 6C. H. transversally extended short and fine ridges and fine radial lirae on growth bands in the posteroventral part of the carapace.

Derivation of name. The figured clam shrimps specimens are from
the northern Chotts range, southern Tunisia.
Type material. Holotype, NIGPCAS 164814, a right valve, male (?).
Dimensions. In order: specimen no. (prefixed NIGPCAS), number of
growth lines, length of carapace (mm), height of carapace (mm),
height/length ratio: 164814, 12, 4.6, 2.9, 0.63; 164815, 10, 4.1, 3.3,
0.80; 164816, 10, 3.9, 2.9, 0.76; 164818, >10, 4.7, 3.1, 0.65; 164819,
14, 5.9, 3.9, 0.66.
Type locality and horizon. The Khanguet Aïcha locality at the ZBAS in
the eastern part of the northern Chotts range, southern Tunisia;
lower Barremian uppermost Bouhedma Formation.
Diagnosis. Carapace small in size, elliptical or oval in outline, with
10e14 growth bands, which near the umbo are ornamented with
small polygonal reticulation; growth bands in the middle part of
the carapace are smooth, while those near the ventral margin are
ornamented with a row of small pits along their lower margins;
 G. Li et al. / Cretaceous Research 72 (2017) 124e133 131

Fig. 7. Ordosestheria chottsensis sp. nov. All are SEM images, except for BeD (light microscopy images). A. NIGPCAS 164818, internal mould of a right valve, male. B. NIGPCAS 164819,
external mould of a left valve, male. C. NIGPCAS 164820, a right vavle, male. D. NIGPCAS 164821, a broken right valve. E. ornamentation on growth bands near the posterior margin
of the carapace of the specimen in Fig. 7C: a row of small pits along the lower margin of each growth band; a row of pores on each growth line; growth lines are serrated in their
lower margins. F. small reticulation on internal surface of the carapace of the specimen in Fig. 7B. G. ornamentation on growth bands near the ventral margin of the broken
specimen in Fig. 7D: a row of small pits along the lower margin of each growth band; a row of small pores on each growth line; growth lines are serrated in their lower margins. H.
ornamentation on growth bands near the posterior margin of the broken specimen in Fig. 7D: a row of small pits along the lower margin of each growth band; a row of small pores
on each growth line; growth lines are serrated in their lower margins.

growth lines, bearing a row of pores, are serrated along their lower
margins.
Description. Carapace small in size, elliptical (male?) or ovate (fe-
male?) in outline. Umbo small, located in the anterior one-third of
the dorsal margin. Growth lines 10e14 in total or a little more,
which are serrated in the lower margins, with a row of pores in
each growth line (Figs. 7E, 7G, 7H). Growth bands narrow near the
umbo, wide in the middle part of the carapace, and narrow again
near the ventral margin (Figs. 6C, 7A, 7D). Growth bands near the
umbo are ornamented with polygonal small reticulations (mesh
132 G. Li et al. / Cretaceous Research 72 (2017) 124e133
diameter 10e15 mm) (Figs. 6E, 6F) with very thin mesh wall. Growth
bands in the middle part of the carapace are smooth. Growth bands
in ventral and posterior parts of the carapace are ornamented with
a row of small pits along their lower margins (Figs. 7E, 7G, 7H). Very
fine, transversely extended short ridges and fine radial lirae are
observed in the posteroventral part of a broken specimen (Fig. 6H).
Small reticulation is observed on the carapace internal surface
(Fig. 6F, mesh diameter about 5 mm).
Discussion. The new species is similar to the type species in having
serrated and pored growth lines, and a row of small pits along the
lower margin of each growth band in the ventral part of the cara-
pace. The new species is easily differentiated from the type species
by having less growth lines (10e14 in total), while the type species
has 26e28 growth lines. Although we have found polygonal small
reticulation on growth bands near the umbo in the new species, we
do not know if this feature is also occurred in the type species. It is
needed to re-study the type species in the near future.
With respect for the sex dimorphism of fossil clam shrimps, it is
normally difficult to precisely distinguish the sex without pre-
served soft parts, such as eggs for female or claspers for male spi-
nicaudatans. But fossil clam shrimp worker has speculated that the
fossil carapace shape does carry a sex-based signal (Tasch, 1969; Li
and Batten, 2004a; Gallego, 2005; Li et al., 2006, 2007, 2010; Chen
et al., 2007; Gallego et al., 2013). As in living spinicaudatans the
male carapaces are slightly longer than those of females and have
somewhat straighter dorsal margin (Shen, 1994; Astrop et al.,
2012). Thus, we doubtfully identify the elliptical carapace as a
male, and an ovate one as a female for the Tunisian specimens.
7. Conclusion
The newly recovered well preserved clam shrimps from the
uppermost Bouhedma Formation is represented by a monospecies
Ordosestheria chottsensis sp. nov. fauna, which is the second species
described in the genus Ordosestheria. Its occurrence in the Chotts
area during early Barremian indicates that ordosestheriids have
originated in northern Africa and then migrated to eastern Asia in
the Aptian. Although some recent spinicaudatans can withstand
low salinity, common clam shrimps are good indicators for fresh-
water conditions. Thus, the two spinicaudatan bearing horizons
(H1 and H2, Fig. 3) in the uppermost Bouhedma and middle Sidi
Aïch formations represent freshwater conditions. The occurrence of
fish and fossil plants in the horizon H2 indicates that a perennial
vegetated pond/oxbow lake has occurred around the study area in a
seasonally humid climate during Barremian.
Acknowledgements
This research is supported by Tunisian Ministry of Scientific
Research (UR 11 ES 15), Major Basic Research Projects of the Min-
istry of Science and Technology, China (973 Project 2012CB822004),
and National Natural Science Foundation of China (41572006,
91514302, 41172010). Many thanks go to anonymous reviewers for
the critical review of the manuscript.
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