Tube structure and original composition of Sinotubulites:

Blackwell Publishing Ltd

shelly fossils from the late Neoproterozoic in southern

Shaanxi, China
ZHE CHEN, STEFAN BENGTSON, CHUAN MING ZHOU, HONG HUA AND ZHAO YUE

LETHAIA Chen, Z. Bengtson, S. Zhou, C.-M., Hua, H. and Yue, Z. 2008: Tube structure and
original composition of Sinotubulites: shelly fossils from the late Neoproterozoic in
southern Shaanxi, China. Lethaia, Vol. 41, pp. 37–45.

Abundant well-preserved shelly fossils of Sinotubulites occur in the upper part of the
late Neoproterozoic Dengying Formation in Ningqiang County, Shaanxi Province,
China. Unlike ‘funnel-in-funnel’ structures of co-occurring Cloudina shells, the tube of
Sinotubulites is nearly cylindrical, composed of several thin layers and characterized by
‘tube-in-tube’ structure. The tube bears prominent longitudinal sculptures and/or
irregular annulations, which were formed by the wrinkles of tube layers, and weaken
gradually towards the inner layers. The irregular wrinkle ornament of the outer layer
shows plastic feature of the Sinotubulites tubes, which may further imply that the
organisms secrete the elastic and flexible organic-dominated walls with mineralization,
possibly aragonitic. Detailed thin section and SEM studies indicate that the silicification
occurred earlier than phosphatation. Specimens preserved in situ from Hubei and sur-
face ornamentation and polygonal shape of the cross-section suggest that Sinotubulites
probably lived as an epibenthos lying on the sea floor.
Aragonitic tube, Ediacaran,
Shaanxi. China, Sinotubulites, tube structure.

Zhe Chen [zhechen@nigpas.ac.cn], Chuan ming Zhou [cmzhou@nigpas.ac.cn], LPS,

Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing
210008, China; Stefan Bengtson [stefan.bengtson@nrm.se], Department of Palaeozoology,
Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden; Hong
Hua [huahong@nwu.edu.cn], Department of Geology, Northwest University, Xi’an
710069, China; Zhao Yue, [zhao_yue30@hotmail.com] Institute of Geology, Chinese
Academy of Geological Sciences, Beijing 100037, China; manuscript received on 25/01/
2007, manuscript accepted 02/06/2007.

A significant event in the history of life is the
acquisition and subsequent evolution of animal
skeletons. During the late Neoproterozoic, most
animals lacked skeletons (Conway Morris 1993; Sun
1994; Narbonne 2005). While the principal episode
of extensive acquisition of durable skeletons did not
start until the beginning of the Phanerozoic (Bengtson
& Runnegar 1992; Sun 1994 and others), there is
more evidence to indicate that biologically controlled
animal biomineralization began in the late Neoprot-
erozoic (Grotzinger et al. 2000; Watters & Grotzinger
2001; Amthor et al. 2003). The tubular fossils of
Cloudina (including Sinotubulites) are potential
index fossils for the terminal Proterzoic (Grant 1990).
Much research has been focused on Cloudina (e.g.
Germs 1972; Glaessner 1976; Hahn & Pflug 1985;
Conway Morris et al. 1990; Grant 1990; Chen & Sun
2001; Hua et al. 2003; Hua et al. 2005), while the
knowledge about Sinotubulites is limited.
Sinotubulites was first described from the base
of Baimatuo Member of the Ediacaran Dengying
Formation in the eastern Yangtze Gorges region,
western Hubei Province, China, and was referred to
as ?Cloudina (Chen & Wang 1977). Chen et al.
(1981) restudied the specimens and erected the new
genus Sinotubulites Chen et al. (1981). They placed
Sinotubulites in the Serpulidae, but expressed uncer-
tainty because of the absence of original structures
and the unknown growth mode. Similar fossils were
discovered from the La Ciénega Formation of Caborca
region of northwestern Sonora, Mexico (McMenamin
1985), from the Precambrian–Cambrian boundary
interval in the Deep Spring Formation of the White-
Inyo Mountains of eastern California and in Esmeralda
County, Nevada (Signor et al. 1983; Signor et al.
1987), and from the upper part of the Dengying
Formation in southern Shaanxi, China ( Zhang et al.
1992; Hua et al. 2000; Zhang & Hua 2000; Chen &
Sun 2001).
In Ningqiang County, Shaanxi Province, abundant
Sinotubulites shells associated with Cloudina occur
in the upper part of the Dengying Formation (the
Gaojiashan Member, Fig. 1). The well-preserved
structures of the Sinotubulites give us an opportunity
to advance the knowledge about this enigmatic group.
In this paper, we describe the construction of the

DOI 10.1111/j.1502-3931.2007.00040.x © 2007 The Authors, Journal compilation © 2007 The Lethaia Foundation
38 Chen et al.
Fig. 1. Geographical location and stratigraphic context of Sinotubulites.
A. Ediacaran outcrop distribution in Ningqiang County, Shaanxi
Province.
B. Generalized stratigraphy of Precambian/Cambrian boundary of the Lijiagou section in Ningqiang County, Shaanxi
Province. AD, Algal dolomite Member.
Sinotubulites tubes and discuss taphonomic and
diagenetic effects on the tube structures, as well as the
original composition, affinity and ecology of these
tubes.
Geological setting
The Precambrian–Cambrian boundary sections in
China are widespread around the Yangtze Platform,
including localities in Yunnan, Sichuan, Hubei, and
Shaanxi provinces. The stratigraphic successions at
all these localities are broadly similar, with a thick
Ediacaran succession dominated by dolomite and a
thin, basal Cambrian phosphatic interval (rich in
small shelly fossils) overlain by predominantly
siliciclastic units in which the first trilobites appear
(Xing et al. 1984).
The Southern Shaanxi region is located at the
northwestern margin of the Yangtze Platform, in
which the Ediacaran (including the Doushantuo and
Dengying formations) was extensively developed.
Here the Doushantuo Formation was deposited in
the Ningqiang-Zhenba area surrounding the Hannan
palaeo-landmass. Lithologically, the Doushantuo
Formation is divided into two parts, the lower part is
chiefly composed of siliciclastic rocks and the upper
one mainly of carbonates. The lower part of the
Dengying Formation consists of grey, medium to
thick-bedded dolomite with vadose structures. At the
bottom of the upper part (the Gaojiashan Member),
LETHAIA 41 (2008)
siltstone and sandy shale containing trace fossils and
metazoans predominate (Zhang et al. 1992; Hua et al.
2000), succeeded by interbedded chert and dolomite.
The top of the upper part of the Dengying Formation
(the lower Cambrian Kuanchuanpu Member) contains
the small shelly fossil assemblages Anabarites–
Circotheca–Protohertzina and Paragloborilus–Tiksitheca.
The lower Cambrian Guojiaba Formation, charac-
terized by black to grey-green siltstone and shale,
disconformably overlies the Dengying Formation.
The lower part of the Guojiaba Formation contains
hyolithids as well as the trilobites Parabadiella,
Wutingaspis, Eoredlichia, Yunnanocephalus, Zhenbaspis
and Hupeidiscus.
Sinotubulites was collected from dolomite of the
Gaojiashan Member of Dengying Formation at the
Lijiagou section (Fig. 1), about 3 km south of the
Kuanchuanpu village, Ningqiang County, Shaanxi
Province. At this horizon, Cloudina (Conway Morris
et al. 1990; Bengtson & Zhao 1992) was also found.
The fossil horizon is about 35 m below the lower
Cambrian Anabarites trisulcotus assemblage.
Tube structure and composition of
Sinotubulites
Tube structure
Sinotubulites baimatuoensis described from the
eastern Yangtze Gorges region occurs in silicified
LETHAIA 41 (2008)
dolomite. All fossils are preserved lying parallel to
bedding. The fossils are either silicified or dolomitized,
without original tube structures preserved. Chen
et al. (1981) described S. baimatuoensis as belonging
to three types, namely ‘large’, ‘medium’ and ‘small’
tubes. The large tubes are 4 –5 mm in diameter, have
an irregular polygonal cross-section and more or less
regular spaced annulations and troughs on the outer
surface. The medium-sized tubes (1.5–3 mm) have
either finely spaced annulations or high and irregular
annulations. The small tubes are 0.4 –1 mm in diameter,
have circular cross-section, and smooth outer surface.
Typical specimens from North America have a single,
rather thick wall, but some specimens appear to have
two or more concentric wall layers, revealing an
irregular but generally double-wall structure (McMe-
namin 1985).
Specimens from southern Shaanxi are similar to S.
baimatuoensis from the Yangtze Gorges in external
appearance and size. However, these specimens are
preserved in amorphous silica and/or phosphate
(Zhang et al. 1992; Chen & Sun 2001), and the well-
preserved specimens show clearly internal tube-in-
tube construction. Obviously, the difference between
two places’ specimens are result of different secondary
diagenesis. The specimens from Shaanxi are described
as S. baimatuoensis here. Sinotubulites baimatuoensis
from Shaanxi can be easily removed from the carbonate
matrix through maceration in an approximately 5 –8%
acetic acid solution.
The tubes of Sinotubulites are straight or slightly
curved, nearly cylindrical to weakly tapering in one
end, and with both ends open (Fig. 2A–C). The diameter
averages 1.6 mm (range from 0.3 to 4.2 mm). The
preserved length of the tube may reach up to 20 mm.
Fig. 2. Sinotubulites and Cloudina.
A–C. Isolated specimens of Sinotubulites from Shaanxi Province, both ends open, bearing transverse
high relief and longitudinal ridges.
A. Note regular transverse relief.
B, C. Note higher and irregular relief.
D. Cloudina from Shaanxi
Province, with one end closed and another open, and ‘funnel-in-funnel’ structure.
E. Sinotubulites baimatuoensis from Hubei Province,
showing irregular relief. Scale bar in C represents 1 mm for A; 1.2 mm for B; 500 µm for C; 280 µm for D and 2.5 mm for E.
Sinotubulites from Neoproterozoic in Shaanxi, China 39
The tube wall is composed of several layers,
normally four to seven in well-preserved specimens
(Figs 3A–F, 4A–G). Longitudinal cross-sections show
that the layers are parallel to each other, and fossils
have the ‘tube-in-tube’ construction (Fig. 3A).
The wall layers are very thin, about 30–50 µm thick,
and distinctly separated from each other by a broad
space (Fig. 4A, C). The multilayered structure may be
obscured by taphonomic and diagenetic alteration
(Figs 3D, 4G). It is circular or irregular polygonal in
transverse section, and the contiguous layers are
slightly eccentrically located (Figs 3E, F, 4A–G).
The tubes show irregular annulations, high relief
and/or longitudinal ridges (Figs 1A–C, 4G). These
surface ornaments, which were formed by wrinkling
of the tube surface layers, are prominent on the outer
wall and become gradually weaker towards the inner
layers (Fig. 3B–D). The innermost layer is straight
and smooth. The different degrees of folding seem to
suggest that the wall ornaments are not the result of
post-mortem deformation.
Tube preservation
Although Sinotubulites was first discovered 20 years
ago, knowledge about this genus is limited. The dif-
ferences in the description of the Sinotubulites tubes
are probably due to different degrees of taphonomic
and diagenetic effect. The understanding of the
taphonomic and diagenetic processes involved in the
preservation of the tube is critical to the interpreta-
tion of the structure.
Sinotubulites tubes from the eastern Yangtze Gorges
region are silicified or dolomitized. Chen et al. (1981)
inferred that these fossils may originally have been
40 Chen et al. LETHAIA 41 (2008)

Fig. 3. Tube structure of Sinotubulites.
A. Longitudinal sections of Sinotubulites, showing multilayered ‘tube-in-tube’ structure, b
(arrowed) points to fracture of the wall.
B,C and D. Multilayered structure and surface ornamentation, note that the surface ornamen-
tation gradually weaker to the inner layer.
B. Enlargement of the area a (arrowed) in A.
E, F. Transverse sections showing almost
centrically nested shell layers.
E. Thin section photograph.
F. Isolated specimen.
G. Irregular outer surface of Sinotubulites. Scale bar
in D represents 800 µm for A; 150 µm for B; 200 µm for C and D; 1 mm for E; 800 µm for F; 150 µm for G.

calcareous thick-walled tubular shells, later replaced Discussion

by silica. Sinotubulites cienegensis found in North
America preserved as calcareous tubes (McMenamin
1985; Signor et al. 1987). Some of the specimens have
their surfaces covered with blotchy, irregular silica
overgrowths, which appear to have filled voids
formed by diagenetic dissolution of the carbonate
matrix surrounding the outer shell surface (McMe-
namin 1985).
The tubular shells of S. baimatuoensis from
southern Shaanxi are preserved either as phosphate,
as silica and phosphate, or as silica only, within a
dolomite matrix (Fig. 5; Chen & Sun 2001). Speci-
mens with thick phosphatic wall show weakly layered
structures (Fig. 4G), while tubes with silica composi-
tion exhibit clear multilayered walls (Fig. 4A–B).
Specimens composed of silica and phosphate have
phosphate coating on the surfaces of silica layers
(Fig. 4C, D), replacing wall layers (Fig. 4E, F), and
filling the space between layers.
Original composition of Sinotubulites tube
Based on their occurrences in the Yangtze Gorges
region and America, Sinotubulites was inferred to
have a calcareous wall (Chen et al. 1981; McMe-
namin 1985), whereas silica and/or phosphate tubes
were found in Shaanxi Province. The Precambrian
has been proposed to be characterized by a ‘soda
ocean’, enriched in sodium carbonate and bicarbo-
nate (Kempe & Degens 1985; but see Hardie 2003;
Brennan et al. 2004), which may have contributed to
the preservation of Precambrian microfossils (e.g.
Martin 1999). Silica and CaCO3 have inverse trends:
as pH increases, silica dissolves and CaCO3 precipitates
(Carson 1991; Tucker 1991). Bacterial oxidation of
organic matter under aerobic conditions led to the
build-up of CO2 in porewaters, the concentration of
dissolved CO2 may have decreased pH and thus
LETHAIA 41 (2008) Sinotubulites from Neoproterozoic in Shaanxi, China 41

Fig. 4. Transverse thin section of Sinotubulites from Shaanxi Province, showing phosphatization processes.
A. Pre-phosphatization
stage, showing thin, constant separated siliceous layers.
B. Close-up of arrowed area in A.
C. Early phosphatization stage, note
phosphate coating on both sides of tube layer.
D. Late phosphatization stage, phosphate replaces siliceous layers and partly fills the space
between layers.
E and F. Enlargements of arrowed areas in D, showing remains of the siliceous layers, plane polarized light.
G. Post-
phosphatization stage, phosphate completely replaces siliceous layers and fills in-between space, multilayered structures are obscured.
Scale bar in G represent 1.2 mm for A; 300 µm for B; 1 mm for C; 500 µm for D; 120 µm for E and F; 800 µm for G.
42 Chen et al. LETHAIA 41 (2008)

Fig. 5. Element distribution maps of thin section of Sinotubulites, showing also Sr, and the way it follows both the P (calcium phosphate)
and the Si (silica) distribution.
LETHAIA 41 (2008) Sinotubulites from Neoproterozoic in Shaanxi, China
resulting in the precipitation of silica (Martin 1999).
Organic decay in carbonate sediments may also be
sufficient to lower pH and cause dissolution of
CaCO3 and precipitation of silica (Knoll 1985).
Our specimens are sometimes preserved in silica
and sometimes in calcium phosphate. Based on thin
section and SEM studies, it is clear that the tubular
fossils underwent post-mortem phosphatization,
and observations on thin sections suggest that the
silicification occurred earlier than the phosphatization
(see Figs 4, 5). At any stage in the breakdown of cells
and tissues, mineral replication – especially silicifica-
tion and phosphatization – can intervene, and pre-
serve surviving cells and tissues in extraordinarily
fine details (Wilby 1993).
The process of phosphatization of Sinotubulites
can be divided into four stages:
1. Pre-phosphatization stage (Fig. 4A, B). The tubes
of Sinotubulites have a clear multilayer structure.
The layers, which are silicified, are thin and
uniform in thickness, and with a distinct space
between adjacent layers.
2. Early phosphatization stage (Figs 3E, 4C). The
tubes show multilayer structure. Phosphate replaced
silica and accreted on both sides of the layers.
The layer thickness increases unevenly.
3. Late phosphatization stage (Fig. 4D). The original
multilayered structure became obscured by phos-
phatization. Phosphate replaced the silica layer
and filled the spaces between the layers. The original
thin layers can only be observed in local areas.
4. Post-phosphatization stage (Fig. 4G). The
characteristic thin layers become less prominent.
Phosphate replaces silica and fills the space com-
pletely. The tubes show thick layered phosphatic wall.
The Sinotubulites fossils from southern Shaanxi have
very thin layers. The layers display both plastic and
brittle behaviour. The wrinkled surface of the tube
seen in both thin sections and isolated specimens
indicates flexibility of the tube layers during the early
secretion. The outer layers of the tubes show folding
in rough accordance with that of adjacent layers, but
the folding becomes gradually fainter towards the
smooth inner layers. This gradation in the degree
of folding shows that the ornaments are original
structure rather than the result of later taphonomic
distortion. Nonetheless, the sometimes brittle
behaviour (Fig. 3A) indicates that the walls were to
some extent mineralized. The trace element Sr is present
in roughly equal amounts (0.3–0.5 atomic percent) in
the phosphate and the silica of the walls, but is absent
in the matrix dolomite rock (Fig. 5). This is consistent
with an original aragonitic mineralization of the tube.
43
The co-occurring Cloudina shows predatory borings
on the calcareous tube (Bengtson & Zhao 1992; Hua
et al. 2003), but no Sinotubulites shells were affected.
The predators selectively chose Cloudina rather than
Sinotubulites as their prey. The great density of
concentric ridges on the shell surface, the tube
architecture of Sinotubulites, and secretion of toxins
or an exterior organic covering might have protected
Sinotubultes from being attacked (Hua et al. 2003).
The most easily recognizable factors that might influ-
ence a predator’s success are features of the prey’s
morphology (Leighton 2001), though many studies
of predatory drilling in the fossil record assume that
for a given predator, shell thickness correlates directly
with drilling time, and is independent of shell structure
(Kitchell et al. 1981). The organic-dominated shell
may also have an influence on predator’s success.
Affinity
Like most other Precambrian fossils, Sinotubulites is
of uncertain affinity. Sinotubulites was originally referred
to as ?Cloudina sp. (Chen & Wang 1977), or thought
to be a synonym of Cloudina that had undergone
alteration during transportation and erosion (Grant
1990). The well-preserved phosphatized specimens
give us an opportunity to distinguish these two genera.
Clearly, Sinotubulites has a tube-in-tube structure,
whereas Cloudina has a funnel-in-funnel structure
(Fig. 2D, Hua et al. 2005). The outer surface ornaments
are also different. Sinotubulites is characterized by
irregular wrinkles on the surface of the outer layers,
and smooth inner layers. Sinotubulites has high irregular
relief and longitudinal ridges on its surface, whereas
Cloudina has regular fine annulations and lacks
longitudinal structure.
Some specimens of Sinotubulites, found in northwest
Sonora, Mexico, appear to have two or more concentric
wall layers, and have been compared with Rugatotheca
He, Platysolenites, and Coleolella (McMenamin 1985).
Well-preserved specimens of Sinotubulites, however,
are multilayered, with normally four to seven layers.
The multilayered structure and irregular surface
ornaments can be used to distinguish Sinotubulites
from these other genera.
Chen et al. (1981) inferred that Sinotubulites probably
belongs to the Serpulidae. Superficially tubes of both
taxa may resemble each other in their longitudinal
and annular sculptures, as well as polygonal shape in
cross-section. The serpulid tube structure, dominated
by chevron-shaped calcified growth layers formed by
an aperturally situated collar gland (Weedon 1994),
gives little cause for comparison with the Sinotubulites
tube structure characterized by thin smooth inner layers
and strongly wrinkled outer ones. Thus, in the
44 Chen et al.
absence of uniquely serpulid characters or soft part in
Sinotubulites, such comparisons are obviously tenuous.
Ecology
Sinotubulites found in Shaanxi has been transported,
and no ecological information can be obtained from
the host rocks. In the eastern Yangtze Gorges region,
however, Sinotubulites in the middle Dengying
Formation is considered to be probably preserved in
situ (Zhao et al. 1985). There, Sinotubulites tubes lie
parallel to the bedding-plane. The tube morphological
characters can also give some ecological clues. For
examples, some annelids and molluscs (such as the
serpulids Eupomatus and Pomatoceros) have polygonal
tube bearing longitudinal sculpture. They are often
attached to various types of substrates including
algae, bryozoans, corals, shells, rocks, and even hulls
of ships. In cross-section, the tubes of serpulids are
usually incomplete in the area of attachment, and if
the tube grows away from the substratum, it becomes
complete (Simkiss & Wilbur 1989). Sinotubulites
may share a similar life habit, but almost all the tubes
of Sinotubulites are complete in transverse section,
which probably suggests that Sinotubulites did not
live the same way as some modern serpulids attaching
on the hard substratum.
All tubes are open at both ends and no attachment
structures have ever been found on either end,
indicating that neither end was attached to a substrate.
Furthermore, the polygonal cross-section and longi-
tudinal ridges may stabilize a tube lying on the sub-
strate, preventing it from rolling in currents. Thus,
we infer that Sinotubulites may be epifaunal, with its
longitudinal ridges and polygonal cross-section
stabilizing the tubes lying procumbently on the
sediment surface during life.
Conclusion
The latest Neoproterozoic tubular fossils Sinotubulites
bears a characteristic tube-in-tube structure. The original
composition of the tube laminae was probably
organic with an uncertain admixture of mineral,
probably aragonite. Both the phosphate and the silica
composition of the walls are of secondary diagenetic
origin. Later phosphatizations may cause an increase
of the layer thickness and obscure the multilayered
structure. Despite its unique morphological charac-
teristics, the affinity of Sinotubulites is still uncertain.
However, the size and morphological complexity
suggest that Sinotubulites shares some similarities
with worm tubes. Sinotubulites appears to have lived
a procumbent epifaunal life.
LETHAIA 41 (2008)
Acknowledgements. – Financial support by the National Natural
Science Foundation of China (40502002), the Chinese Academy
of Science (KZCX-SW-141) and the Chinese Ministry of Science
and Technology (2006CB806400 and 2003CB716805) is greatly
appreciated. We thank Shuhai Xiao and another anonymous
reviewer for their comments.
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