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Pediculate Brachiopod Diandongia pista

from the Lower Cambrian of South China
W A N G Zhifei, HAN Jim, ZHANG Xingliang, LIU Jianni and SHU Degan
Early Life Institute and Key Laboratoryfor Continental Dynamics of the Ministry of Education,
Northwest University, Xi an, Shuanxi 710069;E-mail: zhangelle@sina.corn.cn
Abstract

The Chengjiang LagerstLitte has been celebrated for prolific soft-bodied fossils. Based on specimens recently
excavated in the Chengjiang Lagersatte by the Early Life Institute, Northwest University,Diundongiupistu Rong, 1974, is
directly revealed to be a pediculate brachiopod, assigned to the Family Botsfordiidae, as is further confirmed by the
exceptionally preserved vascular system including dorsal and ventral mantle canals. These specimens described herein
exhibit some peculiarities, notably the extremely thin and long pedicles, which suggest that Diundongiu is epifaunal rather
than burrowing brachiopod. A study of this group of animals indicates that they may be vital to understand the relationship
between the lingulids and the remaining brachiopods, and the character evolution of the early Cambrian brachiopods.

Key words Chengjiang Lagerstiitte, soft-bodied fossils, mantle canals. character evolution, Cambrian, China

1 Introduction
The Chengjiang Lagerstatte from the Lower Cambrian of
China is well-known for yielding delicate, disparate, softbodied fossils (Chen et al., 1995, 1996; Shu et al., 2001a),
particularly, the earliest-known true vertebrates (Shu et al.,
1999,2003) as well as the extinct primitive deuterostomes
(Shu et al., 2001b), providing us the best window into the
Cambrian explosion (Hou et al., 1999; Chen et al., 1996;
Luo et al., 1999). f h e soft-bodied fossils in this fauna throw
much light on the phylogenetic evolution and ecological
features of subsequent marine biota (Jin et al., 1991). The
brachiopods are one of the dominant groups of the
Chengjiang fauna, generally regarded as late Atdabanian in
age (Qian and Bengtson, 1989; Bengtson et al., 1990).
Extensive and intensive excavations of the Chengjiang
Lagerstiittehave made it possible to reveal new information
of the known species. Diandongia pista is initially
described by Rong in 1974, and is considered to be widely
distributed in the uppermost the Qiongzhusi (formerly
Chiungchussu) Formation, Lower Cambrian of Eastern
Yunnan (Rong, 1974). Recent field work by the Early Life
Institute (ELI), Northwest University, revealed that
Diandongia pista is widely distributed nearly in the whole
productive strata containing the Chengjing fauna in the
Shankoucun section, Anning; Ercaicun section and
Jianshan section in Haikou, Kunming (Luo et al.. 1997).
The faunal assemblages in these localities are dominated by
arthropods and worms (Zhang et al., 2001; Han et al.,
2003). Most specimens of Diandongia pista in these

localities are preserved as hard shells without soft bodies.

Fortunately, more than ten specimens in our collection
were preserved with slim pedicles and 12 are preserved
with some fine valve interiors. All the well-preserved
fossils will provide new insights into this group of animals.

2 Terminology
Terminology used in this paper is derived from that of
Rong (1974) and Sutton et al. (1999).

3 Systematic Paleontology
Family Botsfordiidae Schindewolf, 1955
Genus Diandongia Rong, 1974
Diandongia pista Rong, 1974

4 Materials
11 specimens, 12 individuals with tender pedicle; 12

specimens with impressions of vascula of valve interior;

more than 200 specimens with articulated shells and
disarticulated shells without soft body preserved. All these
specimens are reposited in the Early Life Institute (ELI),
Northwest University, Xian.

5 Stratigraphy and Locality

Yuanshan Member (Eoredlichia Zone), Qiongzhusi
Formation, Lower Cambrian. The majority of the

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289

generally stained with

reddish-brown pigment,
through
which
the
interiors of valves are
detectable. The pedicles
are partly or near
completely preserved. A
number of shells of
Diandongia pisra are
found as an attachment
for
other
animals,
illustrated here by a kind
of
gregarious
brachiopods (Fig. 1-1, 2)
with fleshy pedicle,
possible Longtancunella
chengjiangensis
Hou,
1999 (Hou et al., 1999).

7 Description
7. 1 Shell
The description of hard
part mainly refers to that
of Rong (1974) and Luo
et al (1994). Biconvex
valves are approximately
subcircular in dorsoventral views with the
highest point at the umbo
(Fig. 2-1). Valves are
Fig. 1. Diandongia pista Rong attached by gregarious Longtancunella chengjiangensis Hou, 1999,
mainly ornamented with
from the Yuanshan Member of the Qiongzhusi (Chiungchussu) Formation, Lower Cambrian.
pronounced
concentric
Scale intervals are in millimeter.
lines, together with some
feeble costellae in the
specimens with soft bodies were collected from Haikou,
anterior surface of the shell (Fig. 2-1). There are some
Kunming, where the earliest-known agnathans vertebrate
small definite pustules arranging radially in the circular
Haikouichrhys and Myllokunmingia were found (Shu et al.,
protegulum, the posteromedial section of the shell.
1999, 2003), and the others from the Shankoucun section,
Oppositely, some pits in the internal surface of the
Anning, Yunnan Province.
protegulum radiates from the umbo, concentric mound the
umbo. Four neighboring pits form a rhombus (Fig. 2-12).
6 Preservation
The pseudointerarea of ventral valve is crescent-shaped,
extending anteriorly more than one-third length of the
These specimens were collected from the grayish-green
lateral margin (Fig. 2-2). But dorsal pseudointerarea is
and grayish-yellow mudstone in the Mafang section and
absent or comparatively short. Ventral pseudointerarea is
Jianshan section, Haikou, Kunming, and the Shankoucun
divided by deep pedicle groove (Fig. 2-2, 4), forming
section in Anning, Yunnan. Most of specimens of
triangular pseudodeltidium (Fig. 2-3, 4). The
Diandongia pista are preserved as articulated or
pseudodeltidium is about 0.8 nun long in a shell 10.0 mm
disarticulated shells, which were more or less modified by
wide, and is ornamented with transverse lines (Fig. 2-2, 3),
some factors of preservation. The valves of the specimens
as compared with that of Lingulepis nzafongensisRong (Jin
with soft anatomy preserved are commonly crushed or
et al, 1993). From the umbo radiate two pairs of ridges (Fig.
2-5, 6), reminiscent of lobed visceral platform on the floor
deformed. The casts and moulds of these fossils are

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Zhang et al.

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of the valve interior.

7.2 Pedicle
The pedicle is one of the important soft parts in fossil
brachiopods. The pedicle of Diundongiu extends from the
pedicle groove (Fig. 2-2, 4). In the specimens of our
collection, the pedicles of Diundongia pista are up to 16
mm in length (Fig. 2-8, 9), and measure about 0.7 mm in
diameter. Most of them are preserved as slimly linear
impressions (Fig. 2-7,8,9). Clearly they are different from
the pedicles of Cambrian lingulids (Jin et al., 1993) in that
no wrinkles are present and no coelom of the pedicle is
detectable. Nonetheless, there exist some distinct adhered
grains in the distal part of the pedicle of Diundongiu (Fig.
2-9).

7.3 Vascular system

Mantle canals are tubular branching extensions of the body
cavity (coelom). Within them the coelomic fluid circulates,
used mainly for respiration. The mantle canals of
Diundongia are commonly preserved as reddish brown
impressions in the internal moulds of dorsal and ventral
valves, relatively easy to be detected.
In the internal mould of the dorsal valve, a pair of vascula
lateralia arise from the body cavity at a lateral position, and
a pair of vascula media (Fig, 2-13, 14, 15) project from the
body cavity near the midline. As the vascula lateralia and
the vascula media extend anteriorly, each member of them
diverges into 5-6 finger-like branches, the secondary
vascular branches (Fig. 2-13, 14, 15). The arrangement of
vascula is simple in the shell less than 10 mm in width, with
paucity of branches in the peripheral belt of the mantle (Fig.
2-14,15). But the configuration of vascula is comparatively
complex in the shell more than 10 mm in width, showing
not less than four clusters of broom-shaped branches in
peripheral band of the mantle (Fig. 2-13). There arise
distinctly the third branches or distributaries at the end of
the secondary vascular markings (Fig. 2-13). They
terminate blindly near the peripheral belt (Fig. 2-13).
During the growth of a brachiopod its mantle edges become
longer, and the mantle canals would correspondingly tend
to become more widely spaced. Thus, the branches of
vascula accordingly become more and more.

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291

The pallid sinus trunks of the ventral valve of

Diundongiu are represented by a single pair of arcuate
impressions (Fig. 2-10, 16), and they curve forward from a
lateral position on the anterior body wall, recurved
anteriorly toward the symmetrical plane of the valve,
subparallel with the shell margin. Along either of the two
arcuate pallid sinus trunks project peripherally 6 7
baculate branches, at the distal of which exist several minor
digitate distributaries (Fig. 2-10, 16). It is apparent that
these vascular markings are arranged symmetrically about
the median plane in the internal mould of the dorsal valve,
which is indicative of the nature of symmetry of bilateral
metazoan in the Early Cambrian.

8 Comparison
The only brachiopod comparable to Diandongiu is
Heliomedusa orienta Sun and Hou, 1987 (Sun and Hou,
1987; Jin and Wang, 1992) from the Chengjiang fauna.
Diundongiu has been considered as a senior synonym of
Heliomedusu (Jin and Wang, 1992; Geyer, 1994), as
Diundongiu shares some characters with Heliomedusa (Jin
and Wang, 1992). Nevertheless, there are some distinctive
differences between them. The shell of Diundongiu is
smaller and calcification of both valves is much stronger
than that of Heliomedusu. Especially, Diundongiu has a
more complex vascular system (Fig. 2-10, 13, 14, 15, 16), a
pair of pinnately divided vascula lateralia and a pair of
digitately extending dorsal vascula media (Fig. 2-13, 14,
15). By contrast, Heliomedusa possesses a pair of dorsal
vascula lateralia projecting anteriorly from the body cavity
(Jin and Wang, 1992).
Although the length of the pedicle is of little significance
in the specific identification because the pedicle is very
extensible, the presence or absence of pedicles for a
brachiopod may be vital to its definition and identification.
Heliomedusa is one of the dominant groups of brachiopods
in the Chengjiang Lagerslltte (Sun and Hou, 1987; Jin and
Wang, 1992), and an extensive excavation has failed to
reveal the evidence for the pedicle of Heliomedusa,
considered as a late synonym of Diundongiu (Jin and
Wang, 1992). Occurrences of the pedicles of Diundongiu in
conjunction with the differences discussed above,

Fig. 2. Diandongia pisru Rong from the Chengjiang fauna.

1. A pair of articulated valves, ELI-D-0013A; 2. Pedicle valve with faint pedicle impression. showing ventral pseudointerarea and pseudodeltidium, ELI-D-0049 (1);
3. Detail of the pseudodeltidium of 2 with transverse lines; 4. Showing the detail of pedicle groove, ELI-D-0075; 5. Pedicle valve preserved on a cephalon of trilobite,
showing the lobed platform, ELI-D-0037; 6. Ventral exterior, showing the lobed platform, ELI-D-002; 7. Two ventral valves in the same specimen. the lower with
impressions of vascula and the proximal part of pedicle preserved, ELI-D-0001; 8. Ventral valve with elongated, thin pedicle, ELI-D-0072; 9. Pedicle valve with Ushaped recurved pedde, at the distal part of which some adhered grains are preserved, ELI-D-0072; 10. Displaying the mantle canals of ventral valve and the
proximal part of pedicle, ELI-D-0047; 11. Showing a rod-like pedicle jointed with the posterior part of the ventral valve, ELI-D-0022; 12. Showing arrangement of
pits in interior valve, ELI-Doo73; 13. 14 and 15, Dorsal interior showing the vascular system; 13. Fossible adult with vascular markings diverging ~teriorlyand
laterally more times, ELI-D-0003; 14 and 15, Possible young valve showing relatively simple vascular markings of the dorsal valve; 14. ELI-poo42; 15. ELI-DOM I; 16. Showing the mantle canals of ventral valve, ELI-D-0070.
Scale intervals are in millimeter.

292

Lower Cambrian Pediculate Brachiopod from South China

therefore, confirm that Heliomedusa and Diandongia are

two different kinds of brachiopods from the Chengjiang
fauna, both of them indicating the diversity and abundance
of brachiopods in the Lower Cambrian.

9 Significance and Conclusions

The lack of wrinkles in the pedicle may be associated with
the thickness of the chitinous cuticle and muscles layer of
the pedicle. The fact that Diandongia possesses elongated,
thin pedicle without wrinkles may suggest that the
chitinous cuticle and muscle layer of the pedicle is very
thin. The reduction of the chitinous part in Glottidiu is
probably associated with its ability to collect sand grains or
mud (Mackay and Hewitt, 1978). It is possible, even
probable, that the pedicle of Diandongia has no functions
of contraction. At the same time, the thinness of the long
pedicle suggests that it might not have been able to support
the animal. Besides, a number of shells of Diandongia have
been found to serve for attachments by other pediculate
brachiopods (Fig. 1-1, 2). In addition, attention should be
given to the adhered grains at the distal of the pedicle (Fig.
2-9), and the fact that Diandongia was attached by the
pedicle to a free cheek of Eoredlichia (Zhang et al., 2001),
an index fossil for the Lower Cambrian. It is, therefore,
presumed that Diandongia must have been epifaunal, with
the pedicle anchored to another benthic organism or
cemented to other substrates.
The brachiopods are traditionally divided into
inarticulate and articulate. In terms of shell composition,
the brachiopods. could be subdivided into phosphaticshelled and calcareous-shelled brachiopods. Gorjansky and
Popov (1985) and Goryanskij and Popov (1986) placed all
phosphatic brachiopods in a new Class Lingulata outside
the Phylum Brachiopods. Recently, the long-standing
classification of the brachiopod phylum is broken. Chiefly
based on the framework of Popov et al. (1993) and Holmer
et al. (1995), Williams et al. (1996) divided Brachiopoda
into three Subphyla: Linguliformea, Craniformea and
Rhynchonelliformea (Rong and Li, 1997; Clarkson, 1998).
Diandongia is initiately considered as a senior synonym of
Heliomedusa (Jin and Wang, 1992; Geyer, 1994) related
closely to the non-pediculate craniopsids (Jin and Wang,
1992). Later, Diandongia was assigned to Family Obolidae
in 1994 and in 1999 (Luo et al., 1994, 1999). Diandongia,
with the slim, prolonged pedicle protruded posteriorly
between valves, lacking teeth and sockets, should be
referred to recent lingulides. However, Diandongia is
different from lingulides in size, shape, convexity of the
valves, and in the vascular system with dorsal vascula
media developed. It is worthy to be noted that recent
lingulides have a single pair of bifurcate mantle canals in

zhang et al.

both valves, with only vascula lateralia developed (Holmer,

1990). Diandongia has lobed platform on the floor of the
valve interior. The lobe-shaped visceral platform is
characteristic of Botsfordiidae, shared with all members of
Botsfordiidae. Therefore, Diandongia has been included in
this family (Holmer and Popov, 2000). In establishing their
classification, account should be taken of their arrangement
of the main mantle canal impressions. The arrangement of
the pallial sinus trunks of dorsal and ventral valves of
Diandongia is similar to the very nice illustrations of that of
Botsfordia (Popov, 1992). Therefore, their inclusion in
Family Botsfordiidae is further confirmed by the
occurrence of the vascular system of Diandongia. Family
Botsfordiidae is placed within Order Lingulida, Class
Lingulata in the revised Treatise on Invertebrate
Paleontology, Part H (Brachiopoda) (2000).
Diandongia is actually the only strongly mineralized
brachiopods hitherto known in the Chengjiang fauna, with
pinnate dorsal mantle canals and baculiform ventral mantle
canals, as well as minor and degenerating pedicle. This
group of animals, accordingly, may provide a link between
phosphatic-shell and calcareous-shelled brachiopods. They
may be of key importance for understanding the evolution
of the characters of brachiopods in the Early Cambrian, as
well as the origin of Brachiopods (Rowell, 1982;
Goryanskij and Popov, 1986).

Acknowledgements
This work is supported by the National Natural Science
Foundation of China (Grant No. 32070207) and the Science
and Technology Committee of Shaanxi Province. We owe
thanks to Sean P. Robson for his helpful discussion and
suggestions. Special thanks are given to Dick Jefferies,
Fatricio Dominguez and K. Yasui for some important
literatures. Thanks are due to an anonymous referee for
critical and constructive comments on the manuscript. We
also thank Cheng Meirong and Zhai Juanping for
preparation of these fossils, and Cheng Xiuxian for
photographing.
Manucript received Sept. 6,2002
accepted Jan. 12,2003
edited by Wang Sien and Xie Guanglian

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