The First Discovery of ​Marrella​ (Arthropoda, Marrellomorpha) from the

Balang Formation (Cambrian Series 2) in Hunan, China

Author(s): Qing Liu

Source: Journal of Paleontology, 87(3):391-394. 2013.

Published By: The Paleontological Society

DOI: ​http://dx.doi.org/10.1666/12-118.1

URL: ​http://www.bioone.org/doi/full/10.1666/12-118.1

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Journal of Paleontology, 87(3), 2013, p. 391–394
Copyright 2013, The Paleontological Society

0022-3360/13/0087-391$03.00

THE FIRST DISCOVERY OF MARRELLA (ARTHROPODA,

MARRELLOMORPHA) FROM THE BALANG FORMATION
(CAMBRIAN SERIES 2) IN HUNAN, CHINA
QING LIU

State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of
Sciences, Nanjing, 210008, China, ​,​gaius_julius_caesar@yahoo.cn​.

A​BSTRACT​—Marrella, a well-known and important component of the Burgess Shale, was only previously found in the
Burgess Shale and in the Kaili Biota. An exoskeleton attributed to Marrella is described herein from the lower part of the
Balang Formation (Cambrian Series 2) in Paiwu, northwestern Hunan Province, China. It represents not only the third
locality where the genus is found, but also the oldest known occurrence of this taxon.

stratigraphically, and provides new and significant information

regarding the origin of this genus.

INTRODUCTION GEOLOGICAL SETTING AND MATERIAL

The Balang Formation established by Yang Jingzhi and Qian

Yiyuan in 1959 is mainly limited to a Cambrian outcrop belt in

O​
NE OF ​the earliest fossils found by Walcott from the Burgess
Shale (Cambrian Series 3), Marrella first appeared as a ‘‘lace crab’’
sketch in Walcott’s notebook in 1909 before its formal description
as Marrella splendens (Walcott, 1912). Whittington (1971)
provided a major and detailed study of Marrella splendens. A
specimen showing Marrella splendens in the act of molting was
described by Garc´ıa-Bellido and Collins (2004), who also
undertook a restudy and provided new information on the
anatomy, functional morphology, and the behavior of this taxon
(Garc´ıa-Bellido and Collins, 2006). Meanwhile, Zhao et al. (2003)
reported the appearance of Marrella sp. in the lower-middle part of
the Kaili Formation (Cambrian Series 3), representing the only
occurrence of Marrella outside the Burgess Shale. Until the present
study, the stratigraphic range of Marrella was within the Cambrian
Series 3, and the geographic distribution of this genus was only in
the Burgess Shale in British Columbia, Canada and in the
eastern Guizhou Province and western Hunan Province (Zhou et
al., 1979, 1980; Yin, 1987, 1996). The Balang Formation varies
greatly in thickness from 100 m to 700 m (Yin, 1996); it is
commonly 300 m to 600 m thick in Guizhou Province and
generally about 200 m thick in Hunan Province.
Characteristi-cally, trilobites are abundant throughout the Balang
Formation, including genera such as Arthricocephalus, Balangia,
Duyunas-pis, Changaspis and Redlichia (Qian, 1961; Yin, 1987).
The Balang Biota, a Burgess Shale-type fossil assemblage, was
first described from the upper part of the Balang Formation at
Wenglingtang, western suburb of Kaili City, Guizhou Province,
China (Peng et al., 2005). It is characterized by abundant
eocrinoids and covers various phyla including algae,

Kaili Biota in Guizhou Province, China.

In this study, an exoskeleton attributed to Marrella was

discovered in the lower part of the Balang Formation (Cambrian
Series 2) in Paiwu, northwestern Hunan Province, China,
representing the first record of this taxon found in the Balang
Formation in China. This new discovery undoubtedly extends the
distribution of Marrella both paleogeographically and
 and Miao Autonomous Prefecture, Hunan Province, China (Fig.
1.1). This specimen was found in dark gray calcareous mudstone
chancelloriids, cnidarians, priapulids, molluscs, brachiopods, from the lower part of the Balang Formation, and about 61 meters
arthropods and echinoderms, as well as many trace fossils (Peng above the base of the Balang Formation (Fig. 1.2). The specimen
et al., 2005, 2007, 2012). Since then, the Balang Biota has been was wetted with anhydrous alcohol immediately before digital
reported in many other localities and from the middle to the photography to increase contrast.
upper part of the Balang Formation, but was limited to Guizhou
Province until the present study (Peng et al., 2010, 2012; Ma et The specimen described in this paper is housed in the Nanjing
al., 2011). Institute of Geology and Palaeontology, Chinese Academy of
Sciences (prefix NIGP).
The Balang Formation is equivalent to the lower Duyunian
SYSTEMATIC PALEONTOLOGY
Stage (Stage 4) of Qiandongian (Cambrian Series 2) (Peng and
Babcock, 2001). The upper part of the Balang Formation, which
contains abundant Redlichia (Pteroredlichia) chinensis Walcott, Class M​ARRELLOMORPHA​ Beurlen, 1934 Order
1905, is correlated with the lower part of the Ordian of the M​ARRELLIDA​ (Raymond, 1920) Family M​ARRELLIDAE
¨
Walcott, 1912 Genus M​ARRELLA​ Walcott, 1912
Cambrian of Australia (Opik, 1970; Peng et al., 2012).
Arthricocephalus chauveaui and Changaspis elongata are known
Type species.—Marrella splendens Walcott, 1912. From the
in the lower Cambrian Henson Gletscher Formation of Greenland
Burgess Shale, British Columbia, Canada.
(Blaker and Peel, 1997; Peng et al., 2012). The position of the
Diagnosis.—See Garc´ıa-Bellido and Collins (2006).
Balang Biota can be correlated with that of the Guanshan Fauna
and the Emu Bay Shale Lagersta¨tte (Nedin, 1995; Gehling et al.,
Remarks.—In addition to the type species, an unnamed species
2011; Luo et al., 2008; Peng et al., 2010, 2012).
(Marrella sp.) was reported from the Kaili Biota, Guizhou
Province, China (Zhao et al., 2003).
The only specimen of Marrella examined in this study was
collected from Paiwu Village, Huayuan County, Xiangxi Tujia

391
392 JOURNAL OF PALEONTOLOGY, V. 87, NO. 3, 2013

F​IGURE 1—1, map showing the locality where the only specimen of Marrella sp. indet. was collected in Paiwu, northwestern Hunan Province, China; 2,
stratigraphic column showing the position where the specimen was collected from the lower part of the Balang Formation; 3, stratigraphic distribution of the
genus Marrella, including its oldest known occurrence from the Balang Formation.

Occurrence.—North America and Asia, Cambrian Series 2 to

Cambrian Series 3.
 M​ARRELLA​ sp. indet
Figure 2
Material.—One incomplete exoskeleton (NIGP 156216).
Description.—The only specimen is an incomplete exoskele-
ton. The maximum width of the exoskeleton is 8 mm. Cephalic
shield wedge-shaped, bearing one pair of lateral spines, and one
pair of median spines. Left median spine with most parts absent
and the right median spine apically lost in this specimen. Lateral
spines and median spines smooth, with no auxiliary spines.
Lateral spines present medial groove (Fig. 2.1, 2.2).
Occurrence.—From the lower part of the Balang Formation,
Duyunian Stage (Stage 4), Qiandongian (Cambrian Series 2),
Paiwu Village, Huayuan County, Xiangxi Tujia and Miao
Autonomous Prefecture, Hunan Province, China.
Remarks.—Based on the single incomplete specimen, many
in-depth aspects are unclear. Thus, it has been classified as an
undefined species. Compared with the type species Marrella
splendens Walcott, 1912 from the Burgess Shale in British
Columbia, Canada, the most obvious distinction is that the
median cephalic spines have serrated edges in the Canadian
taxon whereas they are smooth in the specimen described herein.
Furthermore, all the specimens of Marrella sp. reported from the
Kaili Biota are represented by exoskeletons, except for one
specimen with an impression of soft tissues (Zhao et al., 2003).
The specimen from the Balang Formation described here is quite
similar to those found in the Kaili Biota in that both have smooth
median cephalic spines.
 LIU—DISCOVERY OF MARRELLA FROM HUNAN, CHINA 393

F​IGURE 2—1, Marrella sp. indet. from the lower part of the Balang Formation in Paiwu, northwestern Hunan Province, China, NIGP 156216; 2, camera lucida
drawing of NIGP 156216. Scale bar is 3 mm. Abbreviations: ms​¼​median cephalic spine; ls​¼​lateral cephalic spine; mg​¼​medial groove.

has been found, which further extends its paleogeographic

DISCUSSION distribution.

Marrella is one of the most common fossils found in the strata
exposed in the Walcott Quarry in the Burgess Shale, and more
than 25,000 specimens have been collected (Garc´ıa-Bellido and
Collins, 2006). Furthermore, it is also the second most common
arthropod in the Walcott Quarry, after the bradoriids, compris-ing
7.3% of the specimens counted (Caron and Jackson, 2008). On
the other hand, only a few dozen specimens were reported from
the Kaili Biota in Guizhou Province, China (Zhao et al., 2003)
and only one specimen was discovered in the Balang Formation
in Hunan Province, China.
The known stratigraphic range of Marrella has long been
limited to the Cambrian Series 3, with the oldest representative
found in the Kaili Biota (Zhao et al., 2003) and the youngest
occurrence in the Burgess Shale (Garc´ıa-Bellido and Collins,
2006). Thus, the new discovery of Marrella sp. indet. from the
lower part of the Balang Formation greatly extends the
stratigraphic distribution of this genus down to the Stage 4 of
Cambrian Series 2 (Fig. 1.3).
The Marrellomorpha, a group of enigmatic arthropods, are only
known from a few species from the early to middle Paleozoic.
Until the present study, the constituent members of this group
include Marrella splendens Walcott, 1912, from the Cambrian
Burgess Shale in Canada (Garc´ıa-Bellido and Collins, 2006);
Marrella sp. from the Cambrian Kaili Biota in China (Zhao et al.,
2003); Austromarrella klausmuelleri Haug et al., 2012, from the
Cambrian Monastery Creek Phosphorite Formation in Australia
(Haug et al., 2012); Furca bohemica Fritsch, 1908, from the
Ordovician Letna´ Formation in Czech Republic (Rak et al.,
2012); Furca sp. from the Ordovician Fezouata Biota in Morocco
(Van Roy et al., 2010); Xylokorys chledophilia Siveter et al.,
2007, from the Silurian Herefordshire Lagersta¨tte in England
(Siveter et al., 2007); Mimetaster

The geographic distribution of Marrella was only within the

Burgess Shale in British Columbia, Canada until the genus was
discovered in the Kaili Biota in Guizhou Province, China (Zhao
et al., 2003). The new discovery of Marrella sp. indet. in Hunan
Province represents a second locality in China where the genus
hexagonalis Gu¨rich, 1931, and Vachonisia rogeri Lehmann,
1955, from the Devonian Hunsru¨ck Slate in Germany (Ku¨hl et
al., 2008; Ku¨hl and Rust, 2010). Two clades were identified with
distinct morphologies: one with a dorsal cordate shield such as
Xylokorys and Vachonisia, and another with prominent cephalic
spines such as Marrella, Furca and Mimetaster (Ku¨ hl et al.,
2008; Ku¨hl and Rust, 2010). The presence of the anterolateral
cephalic spines and secondary spines in Furca and Mimetaster
distinguishes them obviously from Marrella (Rak et al., 2012).
The recent discovery of Marrella sp. indet. from the Cambrian
Balang Formation in Hunan Province, China, provides new
material of the Marrellomorpha, but is also valuable as the oldest
known representative of this group.

ACKNOWLEDGMENTS

I appreciate the comments offered by associate editor S.

Westrop and reviewers D. C. Garc´ıa-Bellido and C. Haug. I am
grateful to Q. P. Lei (Nanjing Institute of Geology and
Palaeontology, Chinese Academy of Sciences) for her generous
help during the research and in field work. I am deeply indebted
to H. C. Zhang (Nanjing Institute of Geology and Palaeontology,
Chinese Academy of Sciences) for providing constructive
suggestions. I particularly thank S. C. Peng (Nanjing Institute of
Geology and Palaeontology, Chinese Academy of Sciences) for
his great support. I thank E. Jarzembowski (Nanjing Institute of
Geology and Palaeontology, Chinese Academy of Sciences) for
improving the English of this manuscript. I thank B. Wang
(Nanjing Institute of Geology and Palaeontology, Chinese
Academy of Sciences) for his kind help. I express great gratitude
to my parents S. Y. Liu and H. Y. Liu for providing great support
in this research.

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