Precambrian Research 191 (2011) 46–57

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Precambrian Research
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Morphology and paleoecology of the late Ediacaran tubular fossil Conotubus

hemiannulatus from the Gaojiashan Lagerstätte of southern Shaanxi Province,
South China
Yaoping Cai a , James D. Schiffbauer b,c , Hong Hua a,∗ , Shuhai Xiao c,∗
a
Early Life Institute, State Key Laboratory of Continental Dynamics, and Department of Geology, Northwest University, Xi’an 710069, China
b
Nanoscale Characterization and Fabrication Laboratory, Virginia Polytechnic Institute and State University Institute for Critical Technology and Applied Science,
Blacksburg, VA 24061, USA
c
Department of Geosciences, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061, USA

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

Article history: Conotubus hemiannulatus Zhang and Lin 1986 is a phylogenetically problematic tubular fossil that is only
Received 7 February 2011 known from the late Ediacaran (ca. 551–541 Ma) Gaojiashan Lagerstätte in southern Shaanxi Province,
Received in revised form 5 September 2011 South China. It is a cm-sized conical tube that tapers adapically from an aperture to a rounded apex.
Accepted 13 September 2011
The tube consists of a series of nested cylindrical-to-funnel-shaped tube walls (cylinders hereafter).
Available online 17 September 2011
This conotubular construction of nested cylinders is similar to that of the late Ediacaran fossil Cloudina.
Integrated morphological, taphonomic, and paleoecological data suggest that C. hemiannulatus occu-
Keywords:
pied an epibenthic life-mode, with the apex anchoring to muddy substrate and the aperture extending
Ediacaran
Gaojiashan
upwards into the water column. It was probably a suspension-feeding organism. This soft bottom dweller
South China employed specialized paleoecological strategies to rejuvenate and self-right the tubes after being sub-
Conotubus jected to sediment obrution, indicating strong burial-resistant capabilities. The tube likely provided space
Cloudina for the living organism, with successively larger cylinders added episodically to accommodate growth.
Currently available evidence suggests that C. hemiannulatus likely had non-biomineralized tubes, and,
although potentially closely related to Cloudina, its phylogenetic affinity remains unresolved.
© 2011 Elsevier B.V. All rights reserved.

1. Introduction phylogenetic affinities of these early tubular fossil assemblages is a

Ediacara-type fossils mark the evolution of complex macro-
scopic organisms, including early metazoans (Narbonne, 2005;
Xiao and Laflamme, 2009). Among all Ediacara-type biotas, the
549–542 Ma Nama assemblage (Grotzinger et al., 1995; Waggoner,
2003), best known from the Kuibis and Schwarzrand subgroups of
the Nama Group in Namibia (Fedonkin et al., 2007), has potential
to bridge the evolutionary gap between the faunas of the Ediacaran
and Cambrian periods. Arguably the most important elements of
the Nama assemblage are the earliest biomineralizing tubular fos-
sils such as Cloudina (Germs, 1972). Common in younger deposits
of the early Cambrian, biomineralized tubes also constitute a sig-
nificant fraction of small shelly fossil assemblages (e.g., Qian and
Bengtson, 1989); and non-biomineralized tubes have been iden-
tified from older assemblages, including the 560–550 Ma Flinders
biota (Droser et al., 2005; Sappenfield et al., 2011) and the >551 Ma
Miaohe biota (Xiao et al., 2002). Therefore, investigating the
∗ Corresponding authors.
E-mail addresses: huahong@nwu.edu.cn (H. Hua), xiao@vt.edu (S. Xiao).
critical step toward testing their evolutionary continuity across the
Precambrian–Cambrian boundary. However, such efforts would be
futile if the morphology and paleoecology of these tubular fos-
sils were not understood in detail. To contribute to this endeavor,
we chose to investigate the tubular fossil Conotubus from the
551–541 Ma Gaojiashan Lagerstätte of the Dengying Formation
(Figs. 1 and 2).
Conotubus is only known from the Gaojiashan Member of the
middle Dengying Formation in southern Shaanxi Province of South
China. The middle and upper Dengying Formation contains a wide
range of phylogenetically problematic forms, including several
tubular fossils such as Cloudina (Conway Morris et al., 1990; Hua et
al., 2003, 2005), Conotubus (Hua et al., 2007), Gaojiashania (Chen
et al., 2002), Shaanxilithes (Chen et al., 1975; Xing et al., 1984),
Sinotubulites (Chen and Sun, 2001; Chen et al., 2008), as well as
the vase-shaped fossil Protolagena (Hua et al., 2010). Of these
taxa, Cloudina is known to have a worldwide distribution (Cortijo
et al., 2010 and references therein), Sinotubulites has been reported
from the Ciénega Formation in the Caborca region of Mexico
(McMenamin, 1985), and Shaanxilithes has been recently reported
from the Yudoma Group of the Siberian Platform (Zhuravlev et al.,

0301-9268/$ – see front matter © 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.precamres.2011.09.002
 Y. Cai et al. / Precambrian Research 191 (2011) 46–57
Fig. 1. Geological map (modified from Cai et al., 2010) showing Ediacaran outcrops in the Ningqiang area, southern Shaanxi Province, South China. Stars indicate occurrences
of Conotubus hemiannulatus. Rectangle in inset map shows the location of the Ningqiang area in the northwestern margin of the Yangtze Platform.
2009; Cai and Hua, 2011). Conotubus and Gaojiashania, on the other
hand, are restricted to and characteristic of the Gaojiashan biota.
In this report, we document the morphology and paleoecology of
Conotubus hemiannulatus, which is regarded as the senior synonym
of many Conotubus species described in the literature.
2. Geological and stratigraphic setting
Precambrian–Cambrian boundary sections crop out widely on
the Yangtze Platform of South China, and their stratigraphic suc-
cession is broadly similar, with a dolostone-dominated upper
Ediacaran sequence, a basal Cambrian phosphorite and chert
interval rich in small shelly fossils, and a predominately silici-
clastic series with Cambrian trilobites. Ediacaran outcrops are
well-exposed in the Ningqiang and Mianxian counties in southern
Shaanxi Province, located in the northwest margin of the Yangtze
Platform (Fig. 1, inset). In the study area, Ediacaran successions out-
crop in a series of approximately northwest–southeast trending
belts separated by faults (Fig. 1). The lower Ediacaran Doushantuo
Formation is characterized by slate, sandstone, and conglom-
erate in the lower part, and carbonate in the upper part. The
overlying Dengying Formation consists of thick-bedded peritidal
dolostone of the Algal Dolomite Member, thin-bedded calcareous
siltstone, mudstone, and limestone characterizing the Gaojiashan
Member, and a second thick-bedded peritidal dolostone unit com-
prising the Beiwan Member. Of particular interest, the Gaojiashan
Member contains abundant three-dimensionally pyritized fossils,
including C. hemiannulatus, which mainly occur in millimetric
beds of fine-grained calcisiltite-siltstone and calcilutite-mudstone
deposited below the storm wave base (Cai et al., 2010). The
Dengying Formation is overlain by phosphatic chert and bitumi-
nous limestone of the basal Cambrian Kuanchuanpu Formation
(Fig. 2).
47
These three lithostratigraphic members of the Dengying For-
mation are similar to, and can be correlated with, the tripartite
divisions of the Dengying Formation in the Yangtze Gorges area,
which is the type area of the Dengying Formation (Zhou et al.,
2007). In the Yangtze Gorges area, the Dengying Formation consists
of the lower Hamajing peritidal dolostone, middle Shibantan subti-
dal bituminous limestone, and upper Baimatuo peritidal dolostone.
This correlation is supported by the occurrence of Sinotubulites in
the correlative Baimatuo and Beiwan members (Chen et al., 2008).
Radiometric dates constrain the Dengying Formation in the Yangtze
Gorges area between 551 Ma (Condon et al., 2005) and 541 Ma, thus
the maximum age range of the Gaojiashan Member is 551–541 Ma
(Fig. 2).
All Conotubus fossils reported here were collected from the
middle Gaojiashan Member of the Dengying Formation at the Gao-
jiashan section (Fig. 2) and are reposited in the Early Life Institute
(ELI), Northwest University, Xi’an, China.
3. Systematic paleontology
Phylum, class, order, uncertain
Family Cloudinidae Hahn and Pflug 1985
The Cloudinidae is characterized by its tubular construction
with nested cylindrical or funnel-like units. Currently, this fam-
ily includes Cloudina, Conotubus, Sinotubulites, Wyattia, and their
synonyms (Grant, 1990; Conway Morris et al., 1990; Chen and
Sun, 2001). Other Ediacaran and Cambrian tubular fossils, both
non-biomineralized and biomineralized tubes, can be differenti-
ated from the Cloudinidae by their simple tube construction and
lack of nested cylinders. These include the worm-like tubular fossil
Sinospongia with closely spaced transverse annulations (Xiao et al.,
2002), the annulated tubular fossil Corumbella with fourfold radial
48 Y. Cai et al. / Precambrian Research 191 (2011) 46–57
Fig. 2. Biostratigraphy of the Ediacaran Gaojiashan Member of the Dengying For-
mation at the Gaojiashan section (see Fig. 1 for location).
symmetry (Hahn et al., 1982; Babcock et al., 2005), cross-walled
tubular microfossils such as Ramitubus, Sinocyclocyclicus, Quadrati-
tubus, and Crassitubus (Liu et al., 2008), giant sinuous tubular fossils
from the Flinders Range (Droser et al., 2005; Sappenfield et al.,
2011), and early Cambrian tubular fossils such as hyoliths, anabar-
itids, and conulariids. Gaojiashania does not have nested cylinders
either, and it can be further distinguished from the Cloudinidae by
its O-ring elements that are often disarticulated (Cai et al., 2010).
Genus Conotubus Zhang and Lin, 1986
Type species: C. hemiannulatus Zhang and Lin in Lin et al., 1986.
non C. varius Zhang, Li and Dong in Li et al., 1992, p. 92, pl. XI, figs.
1, 3–8, pl. IX, figs. 3–4, 6, 9.
Original diagnosis: Straight or curved conical tube preserved
parallel to bedding surface. Tube wall with parabolic folds whose
convex end points posteriorly [toward tube apex]; this feature
is particularly prominent on the dorsal surface. Ventral surface
uneven. Tube interior empty. Tube wall consists of double or multi-
ple layers, which are always pyritized. Tube length 3–80 mm. Tube
diameter 0.3–12 mm at anterior [apertural] end, and 0.1–4 mm at
posterior [apical] end. Transverse cross section is oblate or sub-
circular in shape. [Translated from Lin et al. (1986), with current
authors’ annotations in brackets. The diagnosis provided in Zhang
(1986) is similar.]
Emended diagnosis: Straight or sinuous conotubular tube pre-
served with a closed apex and an open aperture. Tube demarcated
into a series of sections by constriction, abrupt bending, or change
in orientation. Tube wall does not appear to be biomineralized, and
it consists of nested funnel-shaped cylinders that are successively
added at the apertural end. Tube does not branch or split. Tube
length 3–80 mm, aperture diameter 0.3–12 mm, and apex diameter
0.1–4 mm.
Discussion: Conotubus and its type species C. hemiannulatus
were established by Zhang and Lin in Lin et al., 1986. Eight Cono-
tubus species have been described since, including C. hemiannulatus
Zhang and Lin 1986, C. gaojiashanensis Zhang and Lin, 1986, C. mimi-
cus Zhang and Lin, 1986, C. impolitus Zhang and Lin, 1986, C. hamatus
Zhang, 1986, C. cephalotus Zhang, 1986, C. adpressus Zhang, 1986,
and C. varius Zhang, Li, and Dong in Li et al., 1992. Unfortunately,
no holotype was designated for the type species C. hemiannula-
tus, and thus Conotubus and its type species C. hemiannulatus were
not validly published. Also, the original diagnosis of Conotubus and
the distinction of many Conotubus species were largely based on
taphonomic variations, such as the degree of compression and the
thickness of pyritized tube wall which is susceptible to pyrite over-
growth. All described Conotubus species, except C. varius, can be
regarded as taphonomic and ontogenetic variants of a single taxon.
Therefore, to correct the nomenclature errors and to delineate the
morphological limit of Conotubus, this genus and its type species
are redescribed here.
The original diagnosis of Conotubus was partly based on tapho-
nomic features such as pyritization and preservation on bedding
surfaces. The dorsal–ventral differentiation in the original diagno-
sis implies an affinity with bilaterian animals (worms). However,
our observation suggests that this differentiation is a preservational
artifact due to compaction, because less compacted specimens are
circular in cross section and thus lack a dorsal–ventral differentia-
tion. Finally, compaction is also responsible for the imbrication of
the cylinders (Cai et al., 2010), which gives the parabolic appear-
ance of the annulations. All of these features have been removed in
the emended diagnosis.
Among all Ediacaran and Cambrian tubular fossils, Cloudina is
perhaps most similar to Conotubus, as both genera are character-
ized by nested funnels (Hua et al., 2005; Cai et al., 2010). However,
Cloudina tubes are widely believed to be biomineralized or at least
weakly biomineralized (Grant, 1990), whereas Conotubus tubes
are either non-biomineralized or only weakly biomineralized (Hua
et al., 2007). Furthermore, some Cloudina species show evidence of
branching, whereas no branching or splitting specimens of Cono-
tubus have been known so far.
Conotubus varius (Li et al., 1992) is a tubular fossil found in thin
sections of upper Gaojiashan Member carbonate and is 3–25 mm
in length and 0.6–2 mm in maximum diameter. Li et al.’s (1992)
comparative documentation suggested that C. varius would have
been a variant of C. mimicus, as they share a broadly similar
size and morphology but differ in host rock lithology. C. varius is
much smaller than C. hemiannulatus, but similar in size to Cloud-
ina riemkeae. Indeed, its very thin calcareous walls, as visible in
thin sections (Li et al., 1992), are identical to those of C. riemkeae.
It appears that C. varius has biomineralized tube walls. There-
fore, we suggest that C. varius should be excluded from Conotubus
 Y. Cai et al. / Precambrian Research 191 (2011) 46–57
and regarded as a species of Cloudina, perhaps conspecific with C.
riemkeae.
C. hemiannulatus Zhang and Lin in Lin et al. (1986)
C. hemiannulatus Zhang and Lin in Lin et al., 1986, p. 12, pl. I,
figs. 6, 10 (holotype not designated); Zhang, 1986, p. 74–76,
pl. I, figs. 10, 13; Li et al., 1992, p. 91, pl. XI, figs. 9, 16, 18; Zhang
and Hua, 2000, p. 329, pl. I, fig. 8; Cai and Hua, 2007, figs. 1–2.
C. gaojiashanensis Zhang and Lin 1986 in Lin et al., p. 12, pl. I,
figs 3–5, 7; Zhang, 1986, p. 76, pl. I, figs. 1, 3, 9; Li et al., 1992,
p. 91–92, pl. XI, fig. 13; pl. XII, figs. 1, 4.
C. impolitus Zhang and Lin in Lin et al., 1986, p. 12, pl. I, figs. 1, 11;
Zhang, 1986, p. 76, pl. I, fig. 11; Li et al., 1992, p. 92, pl. XI, fig. 14.
C. mimicus Zhang and Lin in Lin et al., 1986, p. 12, pl. I, figs. 2, 9a,
pl. II, fig. 9; Zhang, 1986, p. 76, pl. I, fig. 6; Li et al., 1992, p. 92,
pl. XI, fig. 2, pl. XII, fig. 2.
C. hamatus Zhang, 1986, p. 76–77, pl. I, figs. 2, 4; Li et al., 1992,
p. 92–93, pl. XI, fig. 15.
C. cephalotus Zhang, 1986, p. 77, pl. I, fig. 7; Li et al., 1992, p. 93, pl.
XII, fig. 10.
C. adpressus Zhang, 1986, p. 77, pl. I, figs. 8, 14, pl. II, fig. 7; Li et al.,
1992, p. 93, pl. XII, figs. 3, 5–6.
C. sp., Zhang, 1986, p. 77, pl. I, fig. 12.
Arthrophycus ichnosp., Lin et al., 1986, p. 14, pl. I, figs. 8, 12.
Intrites cf. punctatus, Lin et al., 1986, p. 13, pl. II, fig. 1.
Intrites cf. punctatus, Zhang, 1986, p. 83, pl. IV, fig. 6.
Neonereites ichnosp., Lin et al., 1986, p. 14, pl. II, fig. 16.
Neonereites icnosp., Zhang, 1986, p. 82, pl. III, fig. 11a.
Palaeopascichnus cf. delicatus, Lin et al., 1986, p. 14, pl. II, fig. 10.
Planolites annularis, Zhang, 1986, p. 80, pl. III, fig. 1.
Planolites beverleyensis (Zhang, 1986), p. 80, pl. III, fig. 7.
Planolites sp., Lin et al., 1986, p. 13–14, pl. II, fig. 17.
Planolites ichnosp., Zhang (1986), p. 80, pl. IV, fig. 1.
Skolithos isolatus, Zhang, 1986, p. 82–83, pl. IV, fig. 9.
Unnamed undescribed specimen, Li et al., 1992, pl. XII, fig. 9.
C. sp., Zhang and Hua, 2000, pl. I, fig. 16.
C. sp., Hua et al., 2000, pl. I, fig. 7a.
Skolithos isolatus, Hua et al., 2000, pl. II, fig. 16.
C. sp., Cai and Hua, 2007, fig. 3b–c.
Conotubus, Hua et al., 2007, figs. 5.1–5.3, 6.1–6.2, 7.2–7.3, 7.5, 8.2a.
C. sp., Cai and Hua, 2008, fig. 1G–H.
C. sp., Ma et al., 2008, fig. 3C–F.
Conotubus, Cai et al., 2010, figs. 4B–F, 8A–B, 8D–E, 9A, 9C–J, 10A–B,
10G–I.
Neotype: The specimen illustrated in Fig. 3A is here designated
as a neotype.
Original diagnosis: Compressed conical tube larger [than other
Conotubus species]. Tube diameter 4–10 mm, and anterior [apertu-
ral] end is typically wider [than apical end]. Tube length 40–80 mm.
Tube slightly curved and preserved parallel to bedding surface.
Irregular tube surface at anterior [apertural] end. Posterior [apical]
part of the tube characterized by parabolic annulations or ridges
whose convex side points posteriorly [adapically]; this feature is
prominent on the dorsal surface. Ventral surface uneven. Trans-
verse cross section of tube oblate or elliptical. Tube wall pyritized
with double or multiple layers. [Translated from Lin et al. (1986),
with current authors’ annotations in brackets; the diagnosis pro-
vided in Zhang (1986) is similar.]
Emended diagnosis: Gently curved conotubular fossil, with an
open aperture (Fig. 3A) and a rounded apex (Fig. 3B) that often pen-
etrates into sediments (Fig. 3A). The rounded apex (Fig. 3B) appears
to be anchored in sediments. Original tube wall probably not min-
eralized, and consists of a series of nested funnel-shaped cylinders
(Fig. 3C and D). The cylinders show some transverse annulations
(Fig. 3E). The cylinders are circular in cross section view (Fig. 3F and
49
G). Successive cylinders overlap one other and as many as three
cylinders (Fig. 3H) may be encountered at any given transverse
section of the tubes.
Morphological reconstruction is illustrated in Fig. 4.
Description: C. hemiannulatus is largely preserved as a pyritized
tube in the Gaojiashan Member, although examples of carbona-
ceous compression and glauconitic casting are known (see Cai et al.,
2010, their Fig. 4C and D). There is no strong evidence in support
of biomineralization. Complete tubes typically 3–80 mm in length,
0.1–4 mm in diameter at apex, and 0.3–12 mm in diameter at aper-
ture (see Fig. 5 for measurements of selected specimens). Individual
cylinders 5–8 mm in length, 3–5 mm in diameter at apertural end,
and 2–4 mm in diameter at apical end. Successive cylinders overlap
by 2–5 mm. Cylinders typically have a slightly thickened apertural
rim and an apical rim that can be seen as faint annulus. In cross-
section, tubes are circular in shape (Fig. 3G), although they can be
fractured or flattened due to compaction. Flattened tubes are often
preserved parallel to bedding surface and are semicircular in cross-
sectional views; their stratigraphically lower surface is typically
flatter than the upper surface. Rims and annulations are gener-
ally well developed in apertural part but absent in apical part. In
compacted specimens, rims and annulations are typically deformed
and tilted, so that they tend to imbricated adapically on the strati-
graphically upper surface, and adaperturally on lower surface (Cai
et al., 2010). Tubes devoid of septa, cross walls, or any other inter-
nal structures. Soft tissues are unknown, although Zhang (1986)
interpreted irregular pyrite mass at the apertural end (Fig. 3A) as
pyritized tentacles. The tubes of Conotubus are suggested to be non-
biomineralizing. This is supported by carbonaceous compression
taphonomic mode and flexible (but not brittle) deformation.
Occurrence: The late Ediacaran Gaojiashan Member of the
Dengying Formation in the Ningqiang area of Shaanxi Province,
South China.
Discussion: Zhang and Lin in Lin et al., 1986, illustrated two spec-
imens of C. hemiannulatus, but they did not designate a holotype.
Unfortunately both illustrated specimens were lost, thus we des-
ignate the specimen illustrated in Fig. 3A as a neotype. The species
diagnosis is also emended to eliminate taphonomic features (e.g.,
pyritization, flattening, and imbricated annulations) and unwar-
ranted interpretations (e.g., anterior-posterior and dorsal–ventral
differentiation).
A total of eight Conotubus species have been described in the lit-
erature. Considering taphonomy and ontogeny, we regard seven
as conspecific, and the type species C. hemiannulatus takes pri-
ority. C. hemiannulatus, C. gaojiashanensis, and C. mimicus were
differentiated on the basis of size variation, which are respectively
40–80 mm, 30–40 mm, and 9–15 mm in length, and are respec-
tively 4–10 mm, 3–6 mm, and 1.4–2 mm in apertural diameter. Our
observation suggests that there is a continuous range of length from
13 mm to 80 mm (Fig. 5). Thus, these three species likely represent
different ontogenetic stages of the same taxon. This interpretation
is also supported by the observation that C. mimicus, the smallest
of the three species, tends to be devoid of annulations, a feature
that also characterizes the apical end of the two large species, C.
hemiannulatus and C. gaojiashanensis.
Conotubus impolitus, C. hamatus, C. cephalotus, and C. adpressus
represent taphonomic variants of C. hemiannulatus. C. impolitus,
similar in size to the type species, is characterized by the lack of
annulae and a rough body surface; this difference is most likely due
to pyrite overgrowth on tube wall, leading to coarse pyrite crystals
and the appearance of a rough exterior texture. This may addi-
tionally explain the lack of annulations, as coarser-grained pyrite
may obscure finer surficial details. Only two specimens have been
assigned to C. hamatus, which was established on the basis of a
strongly curved apical part of the tube. However, this curvature
is most likely due to post-mortem deformation or rejuvenation
50 Y. Cai et al. / Precambrian Research 191 (2011) 46–57

Fig. 3. Reflected light photographs showing key morphological features of Conotubus hemiannulatus. (A) Three-dimensionally pyritized specimen preserved parallel to
bedding plane. Note that the apex (black arrow) penetrates into sediments, and a mass of pyrite (white arrow) is located just outside the aperture (ELI-GJSH2010-220;
neotype). (B) A polished slab cut longitudinally through a C. hemiannulatus tube, showing a rounded apex (white arrow). (C) Close-up view of a three-dimensionally pyritized
specimen, showing exterior view of nested cylinders. Note cylinder rim at the apertural end (black arrow) (ELI-GJSH2010-230). (D) A well preserved specimen showing
nested cylinders, each sloping into the underlying cylinder(s). Apical end of cylinders have vaguely double-annulated rim (black arrows). Additional transverse annulae
(white arrows) are visible (ELI-950251). (E) A flattened C. hemiannulatus replicated by clay minerals (with very little pyritization) showing rims of the cylinders (black
arrows) and transverse annulations (white arrows). (F) Fractured surface perpendicular to bedding plane, showing transverse cross section of a pyritized C. hemiannulatus
tube. (G) Transverse cross section view of a slightly flattened tube. Note the thin lamina (arrows) probably representing trace of cylinder wall on which pyrite was precipitated.
(H) Longitudinally fractured specimen, showing traces of multiple thin walls (arrows) of nested cylinders. Gold color is pyrite and dark color is calcite.
 Y. Cai et al. / Precambrian Research 191 (2011) 46–57 51

A Aperture B C
Aperture Aperture

Rims

Rim

Rim

Rims

Apex Apex
Apex

D 1 cm Margins re
r tu
Ape

ex
Ap

Embyonic Conjectured
cone Second section Third section
tentacles

Fig. 4. Morphological reconstruction of Conotubus hemiannulatus. (A) Single cylinder. (B) Nested cylinders. (C) Cut-away view of nested cylinders. (D) Morphological
reconstruction of C. hemiannulatus.

from non-lethal burial. C. cephalotus is characterized by a large
and expanded tadpole-like apertural part, which possibly origi-
nated from secondary pyrite overgrowth or recrystallization giving
the appearance of an inflated apertural part. C. adpressus is char-
acterized by preservation as glauconitic casts, but it is otherwise
very similar to the type species; thus, we regard C. adpressus as a
taphonomic variant of C. hemiannulatus.
8 punctatus, Neonereites ichnosp., Palaeopascichnus cf. delicatus, and
Aperture diameter (mm)
Other Conotubus specimens published under open nomencla-
tures (Conotubus sp. of Zhang, 1986; Conotubus sp. of Zhang and
Hua, 2000; Conotubus sp. of Hua et al., 2000; Conotubus sp. of Cai
and Hua, 2007; Conotubus sp. of Cai and Hua, 2008; Conotubus
sp. of Ma et al., 2008; Conotubus of Hua et al., 2007; Conotubus
of Cai et al., 2010) are herein considered as synonymous to the
type species. Their size ranges and morphologies are similar to
C. hemiannulatus.
Some ichnofossil taxa—including Arthrophycus sp., Intrites cf.
Planolites ichnosp. (Lin et al., 1986); Intrites cf. punctatus, Neonere-

7
6 ites ichnosp., Planolites annularis, Planolites beverleyensis, Planolites
5
sp., and Skolithos isolatus (Zhang, 1986)—are reconsidered as syn-
onymous to C. hemiannulatus. They not only have similar size ranges
4
but also share diagnostic features with C. hemiannulatus, including
3
sparse cross annulations (Palaeopascichnus cf. delicatus, Planolites
2
ichnosp., Lin et al., 1986; Planolites annularis, Planolites beverleyen-
1 sis, Planolites ichnosp., Zhang, 1986) or corrugations (Arthrophycus
0 sp., Lin et al., 1986; Neonereites ichnosp., Zhang, 1986), conical
0 10 20 30 40 50 60 70 80
apical end (previously interpreted as vertical burrows; Intrites cf.
Tube length (mm) punctatus, Lin et al., 1986; Skolithos isolatus, Intrites cf. punctatus,
Zhang, 1986), and indications of incomplete pyritization of Cono-
Fig. 5. Size distribution of 25 relatively complete specimens of Conotubus hemian-
tubus tubes (Neonereites ichnosp., Lin et al., 1986).
nulatus.
52 Y. Cai et al. / Precambrian Research 191 (2011) 46–57

Fig. 6. Reflected light photographs of pyritized Conotubus hemiannulatus specimens showing ontogenetic features. (A) The upper right specimen shows disarticulation
(black arrow), possibly due to weaker connection between two tube sections. The lower left specimen consists of a smooth apical section (a possible embryonic cone), a
second section with sparse annulations, and a third section with nested cylinders. Boundaries between sections are marked by arrows, with white arrow marking possible
rejuvenation after obrution (ELI-GJSH2010-163). (B) Specimen with a smooth and thin apical section succeeded by a thick section with nested cylinders. Boundary between
sections is marked by an arrow (ELI-GJSH2010-195). (C and D) Specimens showing possibly three ontogenetic sections. Boundaries between sections are indicated by arrows,
with white arrows representing paleoecological phenomenon of rejuvenation or periodical growth and black arrows representing probable taphonomic disarticulation. (C)
ELI-950292 and (D) ELI-950002.

4. Morphological reconstruction and possible ontogeny
In spite of previous morphological descriptions of Conotubus (Lin
et al., 1986; Zhang, 1986; Li et al., 1992; Zhang et al., 1992; Hua
et al., 2007), which have focused specifically on the documentation
of morphological outlines and morphometric tube measurements,
the tube wall structure remains inadequately described. Our obser-
vation clearly shows that C. hemiannulatus tube wall consists of
nested cylinders with open ends and a slightly flaring aperture
(Fig. 4). Apertural rims of the cylinders are clearly visible (Fig. 3C
and D). Furthermore, cylinders can be ornamented by transverse
annulations (Fig. 3D and E), which, due to their commonly poor
preservation, can only be found on well-preserved fossils. Because
there are no transverse cross walls or longitudinal septa in the
tubes, we infer that the entire tube enclosed a single living space
rather than a series of separate chambers.
Completely preserved specimens allow us to speculate on the
ontogeny of C. hemiannulatus. Some complete specimens have a
non-annulated apical cone that can be interpreted as a possible
embryonic cone, which is frequently succeeded by one or more
tube sections characterized by nested cylinders with much greater
diameters (Figs. 4D and 6). These sections may represent onto-
genetic stages. The boundary between the embryonic and adult
sections is marked by a constriction preceded by a prominent aper-
tural rim (labeled “margins” in Fig. 4D; Fig. 6B), possibly reflecting
periodic growth. Adult growth of the tube appears to be incremen-
tal and episodic, with successively larger cylinders added to the
apertural end to accommodate the growth of the organism inside
the tube.
Another important feature of C. hemiannulatus is that the apical
part of the tube often shows a stronger degree of deformation than
the apertural part, as evidenced by strongly deformed specimens
(Fig. 7C, D and G). This observation indicates that the apical part
may have been made up of more labile and/or thinner cylinders,
and would have thus been more degradable and/or deformable.
5. Paleoecology
C. hemiannulatus likely inhabited muddy (and possibly
microbially-bound) substrates. The sedimentological and biostrati-
nomic analysis of the Gaojiashan Lagerstätte by Cai et al. (2010)
suggested that the fossiliferous interval is dominated by fine-
grained event deposits (Fig. 8A). Typically, the event beds in the
middle Gaojiashan Member consist of normally-graded siltstone
overlain by a thin mudstone. With the settling of fine-grained
event sediments, the mudstone layer would serve as a soft sub-
strate for colonization of post-event communities. In the absence of
pervasive grazing or bioturbating activities, Ediacaran siliciclastic
substrates are believed to be bound by phototrophic and het-
erotrophic microbes (Seilacher, 1999; Bottjer et al., 2000), although
 Y. Cai et al. / Precambrian Research 191 (2011) 46–57 53

Fig. 7. Reflected light photographs of pyritized Conotubus hemiannulatus specimens, showing important taphonomic and paleoecological features. (A and B) Specimens
showing longitudinal furrows (black arrows) resulting from compaction. (A) ELI-950050 and (B) ELI-950232. (C and D) Part and counterpart showing bending and folding
(black arrows) and shallow longitudinal furrows. (C) ELI-950003a and (D) ELI-950003b. (E) Apertural view (top bedding surface view) of a vertically oriented specimen,
probably preserved in life position (ELI-950018). (F) Side view of an obliquely oriented specimen (stratigraphic up direction on top), probably preserved in life position but
subsequently tilted by compaction. Shown here is the external mold of the specimen. (G) A densely annulated specimen showing post-mortem deformation (black arrows)
and sharp margins (white arrows) between different sections that are interpreted as episodes of growth (ELI-GJSH2010-234). (H) Specimen with dense annulations and
constriction at abrupt bending (white arrow), which is interpreted as rejuvenation from non-lethal burial or periodic growth (ELI-950017).

the presence of microbial mat systems on the substrate has not been
directly observed in the Conotubus-bearing sequence. The muddy
substrate could have been coated with exopolymeric substances
(EPS) secreted by benthic microbes, which provided a cohesive sur-
face for the attachment of C. hemiannulatus and potentially other
organisms with similar life modes.
Some vertically oriented specimens of C. hemiannulatus (Fig. 7E
and F) are interpreted to have been preserved in original life posi-
tion (Fig. 8C), therefore providing direct evidence for a benthic life
mode: at least in juvenile stage, C. hemiannulatus shared a life mode
similar to epibenthic “mat stickers” (Seilacher, 1999). However,
adult individuals of C. hemiannulatus may have employed special-
ized life strategies, with only the most apertural part elevated into
the water column and the bulk of the tube body lying prone on the
muddy substrate. This inference is based on the observation that
different sections of the tubes often have different orientations. We
speculate that, although the non-annulated embryonic cone of C.
hemiannulatus was inserted in the muddy substrate, the apertural
part of the tube may have extended uprightly or obliquely into the
water column. As the tube grew larger and longer during ontogeny
54 Y. Cai et al. / Precambrian Research 191 (2011) 46–57

Fig. 8. Interpretative diagrams illustrating the taphonomy and paleoecology of Conotubus hemiannulatus, emphasizing the role of event deposition. Event beds are numbered
and typically consist of normally graded siltstone overlain by a thin mudstone layer. Tentacles of C. hemiannulatus are conjectural. Three scenarios are considered. (A) C.
hemiannulatus was killed, sometimes fragmented, and buried by obrution of event deposit. Pyritization occurred after burial. (B) C. hemiannulatus was partially buried by event
sedimentation but re-emerged and rejuvenated from partial burial. Rejuvenation was possible because millimetric event beds were not able to kill and bury C. hemiannulatus.
(C) C. hemiannulatus was not affected by the very thin layers of event sediments. It survived the obrution and a section of the tube can be preserved vertically in the sediments.

(Fig. 9A–E), it became increasingly difficult for C. hemiannulatus to
maintain a vertical orientation of a straight tube. Thus, one or more
ontogenetically older sections of the tube may become procumbent
to increase the stability of the tube and to maintain an upright posi-
tion of the ontogenetically youngest section of the tube in order to
elevate the aperture for feeding purposes (Fig. 9F–I).
Alternatively, the different tube sections may represent growth
interruption (by obrution) and post-event rejuvenation: procum-
bent sections may reflect partial burial by obrution deposits and
upright sections represent rejuvenation after partial but non-lethal
burial (Fig. 8B). Cai et al. (2010) noted that the fine-grained event
beds are often only millimetric in thickness and therefore not likely
thick enough to completely bury C. hemiannulatus, which allowed
the possibility of rejuvenation after partial burial. Not surprisingly,
large-sized tubes were more prone to rejuvenation from sediment
obrution than small-sized individuals, which were more easily
smothered by obrution.
Regardless, C. hemiannulatus represents another example of
epibenthic sessile organisms in the Ediacaran Period. Like other
coeval fossils, including the lightly calcified tubular fossil Cloud-
ina identified from multiple continents (Cortijo et al., 2010 and
references therein), the probably calcified goblet-shaped fossil
Namacalathus from Namibia (Grotzinger et al., 2000), Canada
(Hofmann and Mountjoy, 2001) and Oman (Amthor et al., 2003),
and the probably soft-bodied tubular fossil Corumbella from Brazil
(Hahn et al., 1982; Babcock et al., 2005), Conotubus employed a
epibenthic sessile life mode which is characteristic of organisms
occupying tiers above the substrate in the marine realm during the
Ediacaran Period.
6. Phylogenetic position and potential relationship to
Cloudina
The phylogenetic position of C. hemiannulatus is uncertain, and
it is difficult to speculate about its phylogenetic affinity as no
soft parts have been observed. Like other Ediacaran fossils, the
lack of similar post-Ediacaran analogues also presents significant
challenges for phylogenetic interpretation of C. hemiannulatus.
However, the conotubular morphology, apical and apertural dif-
ferentiation, cylinder-in-cylinder tube wall structure, and the
reconstructed paleoecological strategies collectively support a
metazoan affinity. While C. hemiannulatus was originally described
as a bilaterian (Lin et al., 1986; Zhang, 1986), the dorsal–ventral
differentiation, as discussed above, is a taphonomic artifact due
to compaction, and the anterior and posterior ends represent the
aperture and apex of the tubular fossils. Some modern organisms,
such as the primarily deep sea polychaete family Siboglinidae (for-
merly pogonophorans and vestimentiferans; see Brusca and Brusca,
2003), have flanged tubes that are broadly similar to the nested
cylinders of C. hemiannulatus. However, the decidedly more com-
plex and variable tube structures of siboglinids hinder apt biological
comparisons between these two taxa.
The closest relative of C. hemiannulatus is the late Ediacaran
conotubular fossil Cloudina hartmannae. Morphologically, Cono-
tubus differs from Cloudina by its varying growth orientations
between tube sections. In addition, although Cloudina shows
dichotomously branching or split tubes (Hua et al., 2005; Cortijo
et al., 2010), it cannot be confirmed that such bifurcation (and
inferred asexual reproductive strategies) existed in Conotubus. Also,
 Y. Cai et al. / Precambrian Research 191 (2011) 46–57 55

Fig. 9. Reconstructions of possible ontogenetic stages (A–E) and paleoecological strategies (F–I) of Conotubus hemiannulatus. (A–E) Possible ontogenetic stages, including an
embryo (A), smooth initial embryonic cone (B and C), and successive sections of the adult tube (D and E). (F–I) Changing paleoecological strategies at different ontogenetic
stages and after rejuvenation. It became increasingly difficult for C. hemiannulatus to maintain the vertical orientation of a straight tube as it grew larger during ontogeny.
Thus, one or more older sections of the tube may become procumbent to increase the stability of the tube and to maintain an upright position of the ontogenetically youngest
section of the tube. Alternatively, the procumbent sections were partially buried by obrution deposits and the upright section represents rejuvenation after partial but
non-lethal burial. White arrows indicate contacts between tube and substrate.

Cloudina likely had biomineralized tubes (but see Porter, 2007),
whereas Conotubus was probably not biomineralized. Notably,
Cloudina has been suggested to be a prey organism as evidenced
by micro-borings found on the shell (Bengtson and Yue, 1992; Hua
et al., 2003), but these predatory traces have not been observed on
the tube walls of Conotubus. The cosmopolitan distribution of Cloud-
ina also corresponds to a globally consistent latest Neoproterozoic
age assignment of the Cloudina-bearing strata, although whether C.
hemiannulatus shares a cosmopolitan nature is yet unknown as our
knowledge is currently restricted to specimens from Gaojiashan
Member siliciclastics. However, the similarity between Cloudina
and Conotubus in fundamental tube construction with nest funnels
means that they can mutually illuminate each other’s phylogenetic
and morphological obscurities.
Cloudina has been previously compared with both modern
annelids and cnidarians (Grant, 1990; Hua et al., 2005), and sim-
ilar comparisons can be made for Conotubus. Indeed, both Cloudina
and Conotubus occur in the Dengying Formation at the Gaojiashan
section, with the first occurrence of Conotubus precedes that of
Cloudina, leading Hua et al. (2007) to consider Conotubus as a non-
biomineralizing evolutionary precursor of Cloudina. Earlier studies
of Cloudina were primarily based on observation in randomly cut
thin sections (Germs, 1972; Grant, 1990), where Cloudina shells and
constituent cylinders show variable morphologies depending on
the orientation of the fossil within the thin section. On the basis
of these petrographic observations, Grant (1990) reconstructed
Cloudina as having a cone-in-cone test. However, examination of
phosphatized Cloudina fossils extracted from Ediacaran carbon-
ates in South China (Hua et al., 2005) has clearly demonstrated
that Cloudina tests had specifically a funnel-in-funnel construction,
i.e., its constituent cylinders, except the most basal one, are open
at both ends. The funnel-in-funnel construction has been recently
confirmed by secondarily silicified Cloudina material from central
Spain (Cortijo et al., 2010). More importantly, rims at the apertural
end of a given cylinder that is marked by projections and trans-
verse annulations/corrugations are also present in well-preserved
specimens in both Conotubus and Cloudina.
The successive appearance of C. hemiannulatus and Cloudina
hartmannae in the Ediacaran Dengying Formation provides an
opportunity to link paleoecological differences of these two taxa. At
the Gaojiashan section, Conotubus is primarily autochthonously or
para-autochthonously preserved in fine-grained siliciclastics that
are stratigraphically overlain by the Cloudina-bearing limestone
(Cai et al., 2010; Hua et al., 2007). The Cloudina-bearing limestone
sequence contains microbial structures, Cloudina debris, and rip-
up clasts, and is interpreted to have been deposited near or above
storm wave base (Cai et al., 2010). Redeposited Cloudina fragments
were subjected to sorting and corrosion, and thus led to a tapho-
nomically biased record with similarly sized specimens/fragments
concentrated in particular beds. Sorting may cause the observed
size bias of Cloudina fossils, which is exemplified in Cloudina-
bearing units of the Gaojiashan section. In this locale, complete
cm-sized Cloudina individuals can be found within micritic lime-
stones of the lower part of the Cloudina-bearing interval, whereas
only mm-scale debris can be found in thin sections of the upper
part of the Cloudina-bearing interval. Unfortunately, the Cloudina-
bearing limestone sequence is currently inadequately investigated,
and whether it contains direct evidence of the life mode of Cloud-
ina is uncertain. However, C. hemiannulatus specimens preserved in
life position found in the fine-grained siliciclastic sequence provide
direct evidence for life mode reconstruction. Our study suggests
that at least the small-sized C. hemiannulatus occupied an erect life
56 Y. Cai et al. / Precambrian Research 191 (2011) 46–57
mode by anchoring the rounded apex to the possibly microbially-
bound muddy substrate. We argue that Conotubus and Cloudina are
related organisms, with their primary paleoecological difference
falling on environmental preference and a combined procumbent-
erect life mode in Conotubus. They share strong similarities in terms
of morphology, tube wall-building units, and similar life styles.
These similarities reflect their very close phylogenetic affinities.
7. Conclusions
C. hemiannulatus is a cm-sized conotubular fossil that consists of
nested cylinders, with an apex possibly representing an embryonic
cone. Vertically preserved tubes interpreted to be in life posi-
tion suggest that C. hemiannulatus likely had an epibenthic and
sessile mode of life. Integrated morphological, taphonomic, and
paleoecological data allow us to reconstruct C. hemiannulatus as
an epibenthic mat/mud sticker in early ontogeny but employed
other stabilization strategies such as procumbent bending to sta-
bilize larger tubes in late ontogeny. The apertural end, however,
was always elevated to the water column to support a suspension
feeding life mode. Our observations additionally suggest a solitary
rather than a colonial community structure of C. hemiannulatus.
Solitary growth is advantageous in many of the more extreme envi-
ronments, and tends to dominate modern restricted or deep-water
conditions. Their comparatively large size enhances the potential
for survival and rejuvenation after incomplete burial by obrution
sediments. Event-generated sediments may have served as a tapho-
nomic agent for quick burial, adaptive stimulus for rejuvenation,
and a muddy substrate for renewed colonization of C. hemiannula-
tus.
Although it remains phylogenetically problematic, C. hemian-
nulatus plays an important role in deciphering paleoenvironmental
conditions, and is particularly valuable for characterizing Ediacaran
marine ecosystems as both the biosphere and surface environment
of this timeframe are still poorly understood by Phanerozoic stan-
dards. The specialized biological and sedimentological interactions
recorded in the Gaojiashan Lagerstätte provide important insights
into our understanding of paleoecological dynamics and sedimen-
tological regimes of late Ediacaran Period.
Acknowledgments
We thank Hongfu Liu, Shixue Hu, Xingliang Zhang, and Maoyan
Zhu for discussion and Sören Jensen and Brian Pratt for critical
reviews. This work was supported by the National Natural Sci-
ence Foundation of China (Nos. 40872021, 41030209), Program for
New Century Excellent Talents in University, NASA Exobiology and
Evolutionary Biology Program, State Key Laboratory of Continen-
tal Dynamics Research Project, and Northwest University Doctoral
Dissertation Funds (No. 09YYB01).
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