Revision of Fossil Clam Shrimp Taxonomy and a Case Study on
Palaobiogeography of Jurassic Clam Shrimps in China
Gang Li* State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, PR China
Received date: 30 Jul 2017; Accepted date: 20 Sep 2017; Published date: 26
*Corresponding author: Gang Li, State Key Laboratory of Palaeobiology and
Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy
of Sciences, Nanjing, PR China, E-mail: email@example.com;
Clam shrimp carapaces are normally preserved as the fossil records. This makes the morphological features of their carapaces important
for the taxonomy of fossil clam shrimps, such as the size and shape of the carapaces, the position and size of the umbo, the number and
frequency of growth lines, the ornamentation on growth bands, the presence and nature of radial carinae and costae. In this paper a short
review has been made on the Chinese systematic framework for fossil clam shrimps. And a case study was carried out for the Jurassic clam
shrimp biogeography, which may reflect the geographic barrier and/or palaeoclimate change.
Keywords: Fossil clam shrimp; Taxonomy; Palaeobiogeography
Clam shrimps are large branchiopod crustaceans with laterally
compressed bodies enclosed by a bivalved chitinous or complex chitinmineral
carapace [1,2]. They are widely distributed on all continents except
Antarctica [3,4], and inhabit seasonally astatic wetlands such as playas,
vernal (rain and snow-melt) pools, rice field or fishless lakes . These
habitats are normally dry for the most portion of the year, or several years
. This living environment is consistent with their relatively short life
cycles, such as in the species Eulimnadia texana individuals disappeared
two–three weeks after hatching from the egg . Clam shrimps are
important components of an ephemeral freshwater ecosystem [7-9] and
successful colonizers under a wet and dry alternating climate setting in
the earth history, as abundant fossils have been recorded worldwide in the
Mesozoic fine lacustrine deposits [10-21].
Taxonomy of Fossil Clam Shrimps
The fossilized partial soft parts of clam shrimps are very rare in the
geological records [22,23]. Clam shrimp fossils are commonly preserved
as calcium phosphate carapaces , or the external or internal moulds
of carapaces. This means that the classification theme of extant clam
shrimps, basing mainly on soft body structures, cannot be directly
applicable for fossil clam shrimp valves. Because the classification of the
latter is mainly based on the carapace characters, such as the carapace
outline, structure and the fine ornamentation patterns on growth bands
(Figure 1). Nowadays fossil clam shrimps have been accommodated in
four suborders: Laevicaudata Linder, 1945 , Spinicaudata Linder, 1945
, Estheriellina Shen, 2003  and Leaiina Kobayashi, 1972 .
Figure 1: General morphology of the fossil clam shrimp carapace. Left:
a carapace of a palaeolimnadiid; right: a carapace of a leaiid
Laevicaudata Linder, 1945
The fossil laevicaudatans, represented by the superfamily
Paleolynceioidea Tasch, 1956 , are characterized by their carapaces
that have one or no growth line. They have been recorded from the Upper
Jurassic and the Lower Cretaceous.
Spinicaudata Linder, 1945
The suborder Spinicaudata includes three fossil superfamilies, i.e. the
Lioestherioidea Raymond, 1946 , Eosestherioidea Zhang and Chen in
Zhang et al., 1976  and Estheriteoidea Zhang and Chen in Zhang et
al., 1976 .
Lioestherioidea Raymond, 1946: The superfamily Lioestherioidea was
first proposed by Kobayashi (1972) , and its authorship has been
credited to Kobayashi, 1954 (Kobayashi, 1973) [31,32], because he has
nominated the subfamily Lioestheriinae Kobayashi, 1954 . While
Zhang et al. (1976)  credited the authorship of the Lioestherioidea to
Raymond (1946) , because his nomination of the family Lioestheriidae
in 1946 antedated that of Kobayashi. Later, Kozur et al. (1981)  revised
the genus Lioestheria as forms with a semi-globular tubercle and a posteroventrally
directed radial rib on a relatively large umbo. Thus, Chen and
Shen (1985)  subsequently revised the Lioestherioidea (although they
used an –acea suffix) to include forms with a large umbo, on which spines,
nodules, a muscle scar or a carapace gland can occur; carapaces normally
with a few growth lines, which recurved at the dorsal margin, sometimes
they extend above the dorsal margin to form a jagged edge. The geological
range of the Lioestherioidea is from the Middle Devonian through the
Cenozoic. The Lioestherioidea includes seven families as following:
Palaeolimnadiidae Tasch, 1956 : The carapace has a large umbo and
fewer growth lines (Figure 1); ranging from the Late Devonian through
the Late Cretaceous.
Palaeolimnadiopseidae Defretin-Lefranc, 1965 : The umbo is not
so large, the growth lines recurved at the postero-dorsal margin; ranging
from the Middle Devonian through the Late Cretaceous.
Sinoestheriidae Chen and Shen, 1982 : Carapace thin and large;
dorsal margin not thickened or ridge-like and serrated; growth lines stout
and convex, with a row of nodes, slightly recurved near the postero-dorsal
margin; growth bands broad, ornamented with veins, large polygonal
reticulation, fine and sparse radial lirae or bifurcated fibrous sculptures;
ranging from the Late Jurassic to the Cretaceous [36,37].
Ipsiloniidae Novojilov, 1958c : The carapace has a spine or wing-like
process on the dorsal margin behind (sometimes in front) of the umbo;
ranging from the Middle Devonian to the Early Cretaceous.
Lioestheriidae Raymond, 1946 : The carapace has a large umbo,
on which spines, nodules and radial ribs occur; ranging from Late
Carboniferous to Triassic.
Molinestheriidae Kozur, 1980 : The carapace has a large umbo; a long
spine extending from the dorsal ridge; occurring in the Lower Triassic.
Perilimnadiidae Chang and Chen in Chen, 1975 : The carapace has
a large umbo, on which carapace gland or muscle scar occurs, ranging
from the Late Permian through the Early Cenozoic.
Eosestherioidea Zhang and Chen in Zhang et al., 1976 : The
superfamily Eosestherioidea includes forms with small umbo and a large
number of growth lines. The umbo and dorsal margin have no special
structure. The Eosestherioidea ranges from the Middle Devonian through
the Cenozoic, and includes nine families as following:
Ulugkemiidae Novojilov, 1958a : The family is characterized by the
development of growth line indentation in the later stage; its geological
range is from the Middle Devonian to the Permian.
Euestheriidae Defretin-Lefranc, 1965 : The family is characterized
by the small-sized reticulation (mesh diameter normally smaller than
0.02 mm) on growth bands. It ranges from the Middle Devonian to the
Loxomegaglyptidae Novojilov, 1958a : Growth bands are
ornamented with large-sized reticulations. It ranges from the Late
Permian to the Cenozoic.
Orthothemosiidae Defretin-Lefranc, 1965 : Growth bands are
punctate or smooth. It ranges from Late Permian to Triassic.
Aquilonoglyptidae Novojilov, 1958b : Growth bands are ornamented
with scale-like ornamentation. It occurs in the Lower Triassic.
Eosestheriidae Zhang and Chen in Zhang et al., 1976 : The
ornamentation transitions from reticulation in upper and antero-ventral
part of the carapace to radial lirae in the ventral and postero-ventral part
of the carapace. It occurs in the Lower Cretaceous [45,46,47].
Diestheriidae Zhang and Chen in Zhang et al., 1976 :
Ornamentation on growth bands as that in Eosestheriidae, with large
transversely elongated reticulation superimposed over the radial lirae on
the ventral or postero-ventral part of the carapace. It occurs in the Lower
Nestoriidae Shen and Chen, 1984 : Carapace large; growth lines
stout and convex; broad growth bands ornamented with large and shallow
polygonal reticulations. Radial lirae may occur on growth bands near the
venter. It ranges from the Late Jurassic to the Early Cretaceous .
Triglyptidae Wang, 2014 : Carapace ornamented with three kinds
of ornamentation: the dorsal part of the carapace is ornamented with
puncta; the middle part of the carapace with punctate small reticulations;
the lower part of the carapace with radial lirae intercalated with radially
aligned puncta. It occurs in the Middle and Upper Jurassic .
Estheriteoidea Zhang and Chen in Zhang et al., 1976 : The
superfamily Estheriteoidea is characterized by linear, dendritic, latticeshaped,
chain-like ornamentation on growth bands. It ranges from the
Devonian to the Cenozoic, and includes six families as following:
Asmussiidae Kobayashi, 1954 : Growth bands are ornamented with
fine fiber-like ornamentation. It ranges from the Middle Devonian to the
Fushunograptidae Wang in Hong et al., 1974 : Growth bands are
ornamented with simple radial lirae, which are straight, curving or with
intercalated cross bars. Fushunograptids range from the Late Permian to
the Cenozoic [54,55,56]
Jilinestheriidae Zhang and Chen in Zhang et al., 1976 : Growth
bands in the upper part of the carapace are ornamented with widely
spaced radial lirae with intercalated fine reticulations and short radial
lirae; growth bands in the lower part of the carapace are ornamented
with irregular reticulations, which are formed by adding short cross bars
between radial lirae. Members of this family occur in the Cretaceous.
Halysestheriidae Zhang and Chen in Zhang et al., 1976 : Growth
bands near the umbo are ornamented with widely spaced radial lirae
with intercalated fine reticulations and short radial lirae; growth bands in
the lower part of the carapace ornamented with chain-like reticulations.
Halysestheriids occur in the Cretaceous [58,59,60,61].
Estheriteidae Zhang and Chen in Zhang et al., 1976 : Growth
bands near the umbo are ornamented with widely spaced radial lirae,
intercalated with fine reticulations; growth bands in the lower part of the
carapace are ornamented with cavernous ornamentation, which appear as
isolated stout tubercles on the external mould. The members of the family
occur in the Upper Cretaceous .
Dimorphostracidae Chang and Chen, 1964 : Growth bands on the
upper part of the carapace are ornamented with branching, dendritic
radial lirae; growth bands on the lower part of the carapace with dense,
fine and curving radial lirae. It occurs in the Upper Cretaceous.
Estheriellina Shen, 2003 
The family Estheriellidae Kobayashi, 1954 was first elevated to the
superfamily Estherielloidea , including forms with dense radiating
costae (more than five) and were assigned to the suborder Leaiina
Kobayashi, 1972 [30,6]. Then, in considering that the multiple radiating
costae become obsolete near the umbonal area, it was further elevated
to the suborder Estheriellina Shen, 2003, including two superfamilies:
Estherielloidea and Afrograptioidea Novojilov, 1957 .
Estherielloidea Kobayashi, 1954 : Carapace has multiple radiating
costae that become obsolete near the umbonal area. It includes the family
Estheriellidae Kobayashi, 1954 , and occurs in the Lower Triassic.
Afrograptioidea Novojilov, 1957 : Chen and Shen (1985)  elevated
the family Afrograptidae Novojilov, 1957  to a superfamily, and
assigned it to the suborder Estheritina Kobayashi, 1972 . Shen (2003)
 revised the Afrograptidae to include forms with multiple radiating
costae or stout tubercles perpendicular to growth lines present from the
umbo through later growth bands, and suggested its close relationship
with Estherielloidea. The superfamily ranges from the Late Jurassic to the Early Cretaceous.
Leaiina Kobayashi, 1972 
The leaiids have a carapace with up to five radial carinae present from
the umbo through the later growth bands. They range from the Devonian
to the Late Permian, and are accommodated in the superfamily Leaioidea
Raymond, 1946 .
Leaioidea Raymond, 1946 : The superfamily is represented by four
families: i.e. Monoleiolophidae Novojilov, 1954 , Leaiidae Raymond,
1946 , Praeleaiidae Novojilov, 1956  and Rostroleaiidae Novojilov,
Monoleiolophidae Novojilov, 1954 : The carapace has one radial
carinae. The family occurs in the Upper Carboniferous.
Leaiidae Raymond, 1946 : The carapace has two radial carinae (Figure
1). The family ranges from the Middle Devonian to the Late Permian.
Praeleaiidae Novojilov, 1956 : The carapace has 3–5 radial carinae.
The family ranges from the Middle Devonian to the Late Permian.
Rostroleaiidae Novojilov, 1956 : The carapace has up to five radial
carinae; growth lines recurve at antero-dorsal and/or postero-dorsal
margins. The family ranges from the Middle Devonian to the Permian.
Spinicaudatans have carapaces with growth lines that reflect successive
moultings. During ecdysis spinicaudatans only cast off an inner skeletal
duplicature, but they do not shed their carapace, a growth line is added
peripherally to each valve . Thus, there would be the possibility to
draw ontogenetic information within an individual carapace. In the case
of the Middle Jurassic species Shizhuestheria truncata the first 10–14
growth bands are wide and relatively thin, while the later growth bands
are narrow. This indicates a rapid growth rate during the junior stage,
and a lower growth rate during the adult stage. The transition of growth
bands from wide to narrow is clearly exhibited. This shift in carapace
accretion rate could be attributed to the onset of the sexual maturity, and
to the transition in biological priorities from rapid juvenile growth to
reproductive activities .
Jurassic Clam Shrimp Biogeographic Provinces in China
Jurassic is a warm greenhouse period in the deep time of the earth
history. In China the Jurassic is mainly of non-marine origin. Abundant
fossil clam shrimps have been recorded in the Jurassic lacustrine deposits
[30,70,71]. This made it possible to discuss the palaeobiogeographic
During the Early Jurassic, in southern China, the palaeo-Yangtze
river, starting from the east, ran westwards along the Yunmeng, Ba-
Shu and Yunnan lakes into Tethys in southwestern Yunnan (Figure 2).
In northern China the biggest lake Qingyang occupied the Ordos basin,
and the water replenishment was supplied by three big rivers, e.g. the
Yan and Central Plains rivers in the east, and the Corridor river in the
west. East Tethys occupied Tibet and southwestern Yunnan Province. The
Guangdong-Jiangxi bay separated the southeastern costal region from
mainland China, and the Ussuri bay reached the eastern Heilongjiang
region . The Early and Middle Jurassic warm and humid climate
made clam shrimp faunas evenly distributed in China. The Early Jurassic
Palaeolimnadia baitianbaensis fauna and the Middle Jurassic Euestheria
ziliujingensis fauna flourished in the palaeo-Yangtze and the Qingyang
lake drainage systems, in the Junggar, Turpan and Qaidam basins, and
also occurred in the southeastern costal region. The late Early Jurassic
Eosolimnadiopsis fauna distributed mainly in the southeastern costal
region and the Qingyang lake .
Since the Late Jurassic with the gradual break-up of the Eurasian
continent, the warm, humid climate of the Early–Middle Jurassic period
 gave way to a warm, extremely arid climate in East Asia. Clam
shrimps differentiated into three faunas, e.g. the Tithonian Pseudograpta
fauna in northern China and the Kimmeridgian Qinghaiestheria-
Mangyalimnadia fauna in the Qaidam Basin, and the Kimmeridgian–
Tithonian Eosestheriopsis fauna in southwestern China  (Figure 3).
Qinghaiestheria is a common component of the latter two faunas, which
indicates that they belong to the same southern palaeobiogeographic
province. Thus, researchers should pay more attention on the detailed
distribution of fossil clam shrimps to reconstruct palaeo-drainage system
Figure 2: Distribution of Early and Middle Jurassic clam shrimp faunas in China (after Chen, 1979 ; Chen and Shen, 1983 ; Li and Matsuoka,2012 )
Figure 3: Distribution of Late Jurassic clam shrimp faunas in China (after Chen, 1979 ; Chen and Shen, 1983 ; Li and Matsuoka, 2012 ))
In China the non-marine deposits yield abundant clam shrimp fossils,
which would be a great help to propose a systematic framework. In
combining the recent clam shrimp classification theme, fossil clam shrimps
have been accommodated in four suborders: Laevicaudata Linder, 1945
, Spinicaudata Linder, 1945 , Estheriellina Shen, 2003  and
Leaiina Kobayashi, 1972 . The Chinese Jurassic clam shrimp data are
used to delineate the palaeobiogeographic provinces, and to reconstruc
the old drainage systems. The Early and Middle Jurassic warm and humid
climate made clam shrimp faunas evenly distributed in China. While
during the Late Jurassic clam shrimps have differentiated into the northern
and southern palaeobiogeographic provinces, i.e. the Pseudograpta fauna
in the north, and the Qinghaiestheria–Mangyalimnadia fauna and the
Eosestheriopsis fauna in the south.
This study was supported by the National Natural Science Foundation
of China (41572006, 91514302, 41688103) and by the Chinese Academy
of Sciences (XDPB05).
Tasch P (1969) Branchiopoda. In Moore RC (Ed) Treatise on invertebrate paleontology, part R, arthropoda 4 Lawrence, Geological Society of America, Boulder, and University of Kansas Press R128-eR191. Astrop TI, Hegna TA (2015) Phylogenetic relationships between living and fossil spinicaudatan taxa (Branchiopoda Spinicaudata): reconsidering the evidence. J Crustacean Bio 35: 339-354.Brendonck L, Rogers DC, Olesen J, Weeks SC, Hoeh R (2008) Global diversity of large branchiopods (Crustacea: Branchiopoda) in fresh water. Hydrobiologia 595: 167-176.Rogers DC, Rabet N, Weeks SC (2012) Revision of the extant genera of Limnadiidae (Branchiopoda: Spinicaudata). J Crustacean Bio 32: 827-842.Dumont H, Negrea S (2002) Introduction to the class Branchiopoda. Guides to the Identification of the Microinvertebrates of the Continental Waters of the World. Leiden, Backhuys, 1-398.Chen PJ, Shen YB (1985) An introduction to fossil Conchostraca. Science Press, Beijing, 241, 26 pls (in Chinese).Guériau P, Robet N, Clément G, Lagebro L, Vannier J, et al. (2016) A 365-Million-Year-Old Freshwater Community Reveals Morphological and Ecological Stasis in Branchiopod Crustaceans. Curr Bio 26: 383-390.Li G (2017) SEM morphological study of the type species of Ordosestheria. Wang, 1984 (Spinicaudata) from Ordos Basin of mid-west China. Cretaceous Res 75: 1-6.Li G (2017b) Morphological restudy of the type species of Xibeiestheria (Spinicaudata) from the lower Aptian, northwestern China. Cretaceous Res 80: 31-37.Li G (1993) Early Cretaceous fossil conchostracans from southern Shaanxi and western Henan. Palaeoworld 2: 57-72.Shen YB (1994) Jurassic conchostracans from Carapace Nunatak, southern Victoria Land, Antarctica. Antarct Sci 6: 105-113.Chen PJ, Li G, Batten DJ (2007) Evolution, migration and radiation of late Mesozoic conchostracans in East Asia. Geol J 42: 391-413.Gallego OF, Monferran MD, Astrop TI, Zacarias IA (2013) Reassignment of Lioestheria codoensis Cardoso (Spinicaudata, Anthronestheriidae) from the Lower Cretaceous of Brazil: systematics and paleoecology. Revista Brasileira de Paleontologia 16: 47-60.Boukhalfa K, Li G, Ben Ali W, Soussi M (2015) Early Cretaceous spinicaudatans (“conchostracans”) from lacustrine strata of the Sidi Aïch Formation in the northern Chotts range, southern Tunisia: taxonomy, biostratigraphy and stratigraphic implication. Cretaceous Res 56: 482-490.Li G, Matsuoka A (2015) Searching for a non-marine Jurassic/Cretaceous boundary in northeastern China. J Geological Society of Japan 121: 109-122.Scholze F, Golubev VK, Niedźwiedzki G, Sennikov AG, Schneider JW, et al. (2015) Early Triassic Conchostracans (Crustacea: Branchiopoda) from the terrestrial Permian–Triassic boundary sections in the Moscow syncline. Palaeogeogr Palaeoclimatology Palaeocl 429: 22-40.Schneider JW, Scholze F (2016) Late Pennsylvanian–Early Triassic conchostracan biostratigraphy: a preliminary approach. Geological Society London, Special Publications 450, SP450-SP456.Teng X, Xiao JN, Zhang YZ, Matsuoka A, Li G (2016) Nestoria sikeshuensis (spinicaudatan), a new clam shrimp species from the Tugulu Group in Junggar Basin, northwestern China. Sci Rep Niigata Univ (Geology) 31: 75-81.Li G, Boukhalfa K, Teng X, Soussi M, Ben Ali W, et al. (2017a) New Early Cretaceous clam shrimps (Spinicaudata) from uppermost Bouhedma Formation of northern Chotts range, southern Tunisia: Taxonomy, stratigraphy and palaeoenvironmental implications. Cretaceous Res 72: 124-133.Li YL, Teng X, Matsuoka A, Li G (2017b) SEM morphological study of clam shrimp Diestheria (spinicaudatan) of the Jehol Biota of China. Sci Rep Niigata Univ (Geology) 32: 15-23.Teng X, Li G (2017) Clam shrimp genus Ordosestheria from the Lower Cretaceous Dalazi Formation in Jilin Province, north-eastern China. Cretaceous Res 78: 196-205.Zhang WT, Shen YB, Niu SW (1990) Discovery of Jurassic conchostracans with well-preserved soft parts and notes on its biological significance. Palaeontologia Cathayana 311-351.Orr PJ, Briggs DEG (1999) Exceptionally Preserved Conchostracans and Other Crustaceans from the Upper Carboniferous of Ireland. Special Papers in Palaeontology 62: 5-68.Stigall AL, Babcock LE, Briggs DEG, Leslie SA (2008) Taphonomy of Lacustrine Interbeds in the Kirkpatrick Basalt (Jurassic), Antarctica. Palaios 23: 344-355.Linder F (1945) Affinities within the Branchiopoda with notes on some dubious fossils. Arkiv för Zoologi 37A: 1-28. Shen YB (2003) Review of the classification of the family Afrograptidae (Crustacea: Conchostraca). Acta Palaeontologica Sinica 42: 590-597.Kobayashi T (1972) On the two discontinuities in the history of the Order Conchostraca. P Jpn Acad 48: 725-729.Tasch P (1956) Three general principles for a system of classification of fossil conchostracans. J Paleontol 30: 1248-1257.Raymond PE (1946) The genera of fossil Conchostraca-an order of bivalved Crustacea. B Museum Compactive Zoöl 96: 217-307.Zhang WT, Chen PJ, Shen YB (1976) Fossil Conchostraca of China. Science Press, Beijing, 1-325 (in Chinese).Kobayashi T (1954) Fossil Estherians and allied fossils. : 1-192.Kobayashi T (1973) On the classification of the fossil Conchostraca and the discovery of Estheriids in the Cretaceous of Borneo. Contributions to the geology and palaeontology of southeast Asia, 130. Geology and palaeontology of Southeast Asia 13: 47-72.Kozur H, Martens T, Pacaud G (1981) Revision von "Estheria" (Lioestheria) Lallyensis Depéret & Mazeran, 1912 und "Euestheria" autunensis Raymond, 1946. Zeitschrift für geologische Wissenschaften 9: 1437-1445.Defretin-Lafranc S (1965) Étude et révision de phyllopodes conchostracés en provenance d’U.R.S.S. Annales de la Société Géologique du Nord 85: 15-48.Chen PJ, Shen YB (1982) Late Mesozoic conchostracans from Zhejiang, Anhui and Jiangsu provinces. Palaeontologia Sinica Ser. B. 17, 1–116 (in Chinese, English summary).Chen PJ, Shen YB (2014) On the family Sinoestheriidae (Crustacea: Diplostraca: Spinicaudata). Acta Palaeontologica Sinica 53: 443-451.Li G, Teng X, Matsuoka A (2016a) SEM morphological study of clam shrimp Ganestheria (spinicaudatan) from Upper Cretaceous of Jiangxi, southeastern China. Sci Rep Niigata Univ (Geology) 31: 69-74.Novojilov NI (1958c) Conchostraca de la super famille des Limnadiopseidea superfam. nov. Annales du Service d’Information Géologique du Bureau de Recherches Géologique, Géophysiques et Miniéres 26: 95-128.Kozur H (1980) . Zeitschrift für Geologische Wissenschaften 8: 885-903.Chen PJ (1975) Tertiary conchostracans of China. Scientia Sinica 6: 618-630. (1958a) Nouveaux Conchostraca fossils. Recueil D’Articles sur les Phyllopodes Conchostraces. Annales du Service d’Information Gologique du Bureau de Recherches Géologiques, ophysiques et Miniéres 26: 7-13.Li G, Wan XQ, Willems H, Batten DJ (2007b) Revision of the Conchostracan Genus Tenuestheria from the Upper Cretaceous Lanxi Formation in Zhejiang and Its Biostratigraphic Significance in Southeast China. Acta Geologica Sinica 81: 925-930.Li G, Hirano H, Batten DJ, Wan XQ, Willems H, et al. (2010) Biostratigraphic significance of spinicaudatans from the Upper Cretaceous Nanxiong Group in Guangdong, South China. Cretaceous Res 31: 387-395.Novojilov NI (1958b) Conchostraca du Permien et du trias du littoral de la mer des Laptev et de la Toungounzka inférieure. Annales du Service d’Information Géologique du Bureau de Recherches Géologiques, Géophysiques et Miniéres 26: 15-80.Li G, Wang SE, Shen YB (2006) Revision of the genus Abrestheria (Crustacea: Conchostraca) from the Dabeigou Formation of northern Hebei, China. Prog Nat Sci 16(Special Issue) 284-291.Li G, Shen YB, Batten DJ (2007a) Yanjiestheria, Yanshaniaand the development of the Eosestheria conchostracan fauna of the Jehol Biota in China. Cretaceous Res 28: 225-234.Li, G., Matsuoka, A., Willems, H., 2015. SEM morphological study of the clam shrimp type specimens of Eosestheria sihetunensis from the Lower Cretaceous Yixian Formation in western Liaoning, northeastern China. Sci Rep Niigata Univ (Geology) 30: 27-37.Li G, Ohta T, Batten DJ, Sakai T, Kozai T (2016b) Morphology and phylogenetic origin of the spinicaudatan Neodiestheria from the Lower Cretaceous Dalazi Formation, Yanji Basin, north-eastern China. Cretaceous Res 62: 183-193.Shen YB, Chen PJ (1984) Late Middle Jurassic conchostracans from the Tuchengzi Formation of W. Liaoning, NE China. Bulletin of the Nanjing Institute of Geology and Paleontology, Academia Sinica 9: 309-326.Li G, Shen YB, Liu YQ, Bengtson DJ, Willems H,et al. (2009a) Revision of the clam shrimp genus Magumbonia from the Upper Jurassic of the Luanping Basin, Hebei, Northern China. Acta Geologica Sinica 83: 46-51.Wang SE (2014) Triglyptidae Fam. Nov. and its significance in evolution and biostratigraphy. Acta Paleontologica Sinica 53: 486-496.Li G, Ando H, Hasegawa H, Yamanoto M, Hasegaea T, at al. (2014a) Confirmation of a Middle Jurassic age for the Eedemt Formation in Dundgobi Province, southeast Mongolia: constraints from the discovery of new spinicaudatans (clam shrimps). Alcheringa 38: 305-316., et al. (1974) Stratigraphy and palaeontology of Fushun coal-field, Liaoning Province. Acta Geologica Sinica 2: 113-149.Li G (2004) Discovery of Qinghaiestheria from the Upper Jurassic Penglaizhen Formation in Sichuan, southwestern China. J Asian Earth Sci 24: 361-365.Li G, Batten DJ (2004a) Cratostracus? cheni, a new conchostracan species from the Yixian Formation in western Liaoning, north-east China, and its age implications. Cretaceous Res 25: 577-584.Li G, Batten DJ (2004b) Revision of the conchostracan genera Cratostracus and Porostracus from Cretaceous deposits in north-east China. Cretaceous Res 25: 919-926.Li G, Wang SE, Chen PJ, Willems H (2014b) Morphological study of the type species of Fengninggrapta (Crustacea: Spinicaudata) from the Xiguayuan Formation of northern Hebei Province, northern China. Acta Palaeologica Sinica 53: 527-532.Li G (2005) Halysestheria yui from the Nenjiangian Taipinglinchang Formation of Heilongjiang Province, China. Acta Palaeontologica Sinica 44: 322-324.Li G, Chen PJ, Wang DY, Batten DJ (2009b) The spinicaudatan Tylestheria and biostratigraphic significance for the age of dinosaur eggs in the Upper Cretaceous Majiacun Formation, Xixia Basin, Henan Province, China. Cretaceous Res 30: 477-482.Li G, Wan XQ, Batten DJ, Bengtson P, Xi DP, et al. (2009c) Spinicaudatans from the Upper Cretaceous Nenjiang Formation of the Songliao Basin, northeast China: taxonomy and biostratigraphy. Cretaceous Res 30: 687-698.Li G, Matsuoka A (2013) Revision of clam shrimp (“conchostracan”) genus Tylestheria from Late Cretaceous deposits of China. Sci Rep Niigata Univ (Geology) 28: 51-63.Li G, Batten DJ (2005) Revision of the conchostracan genus Estherites from the Upper Cretaceous Nenjiang Formation of the Songliao Basin and its biogeographic significance in China. Cretaceous Res 26: 920-929. Chang WT, Chen PC (1964) New Cretaceous Conchostraca from Jilin and Heilongjiang. Acta Palaeontologica Sinica 12: 1-15.Novojilov NI (1957) Crustacés bivalves de l’ordre des conchostracés du Crétacé inférieur Chinois et Africain. Annales de la Société Géologique du Nord 77: 235-243.Novojilov NI (1954) Upper Jurassic and Cretaceous conchostracans from Mongolia. Trudy Paleontologicheskogo Instituta 48: 7-124.Novojilov NI (1956) Crustaces phyllopodes bivalves 1 Leaiidae: Adad Nauk SSSR, Inst. Paeont, Travaux 61: 1-128.Li G, Hirano H, Kozai T, Sakai T, Pan YH (2009d) Middle Jurassic spinicaudatan Shizhuestheria from the Sichuan Basin and its ontogenetic implication. Science in China Series D: Earth Sciences 52: 1962-1968.Chen PJ (1979) An outline of palaeogeography during the Jurassic and Cretaceous Periods of China-with a discussion on the origin of Yangtze River. Acta Scientiarum Naturalium Universitatis Pekinensis : 519-524. Chen PJ, Shen YB (1983) Jurassic and Cretaceous conchostracan biogeographic provinces of China. In: Lu YH, Mu EZ, Zhou MZ, Yang ZY, Wang Y, et al. (Eds) Palaeobiogeographic Provinces of China, Science Press, Beijing, 131-141. Zhang YZ, Li G, Teng X, Wang LH, Cheng XS (2017) New Jurassic spinicaudatans from Xinjiang Uygur Autonomous Region of northwestern China and their evolutionary implications. Palaeoworld.Li G, Matsuoka A (2012) Jurassic clam shrimp (“conchostracan”) faunas in China. Sci Rep Niigata Univ (Geology) 27: 73-88. Chen PJ, Norling E (2002) Postpalaeozoic stratigraphy, palaeogeography and tectonic evolution of the Tarim Basin, Xinjiang, NW China. Palaeoworld 14: 1-48.