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A fresh look at Nisusia Walcott, 1905 from the Cambrian Kaili Formation in Guizhou
Accepted Manuscript Title: A fresh look at Nisusia Walcott, 1905 from the Cambrian Kaili Formation in Guizhou Author: Yong-Qin Mao Yuan-Long Zhao Cheng-Wen Wang Timothy Topper PII: DOI: Reference:
S1871-174X(16)00021-4 http://dx.doi.org/doi:10.1016/j.palwor.2016.03.001 PALWOR 354
To appear in:
Palaeoworld
Received date: Revised date: Accepted date:
23-6-2015 4-1-2016 8-3-2016
Please cite this article as: Mao, Y.-Q., Zhao, Y.-L., Wang, C.-W., Topper, T.,A fresh look at Nisusia Walcott, 1905 from the Cambrian Kaili Formation in Guizhou, Palaeoworld (2016), http://dx.doi.org/10.1016/j.palwor.2016.03.001 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
A fresh look at Nisusia Walcott, 1905 from the Cambrian Kaili Formation in Guizhou
a
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Yong-Qin Mao a, b, Yuan-Long Zhao c *, Cheng-Wen Wang a, Timothy Topper d
College of Earth Sciences, Jilin University, 130061 Changchun, Jilin Province, P.R.
b
cr
China
Institute of Mountain Resources, Guizhou Academy of Sciences, 550001 Guiyang,
College of Resources and Environmental Engineering, Guizhou University, 550025
Guiyang, Guizhou Province, P.R. China d
an
c
us
Guizhou Province, P.R. China
Department of Palaeobiology, Swedish Museum of Natural History, P.O. Box 50007,
M
SE-104 05, Stockholm, Sweden
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Abstract
d
* Corresponding author. E-mail address: [email protected]
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Nisusia Walcott, 1905 is an early genus of kutorginid brachiopod of the Subphylum Rhynchonelliformea. Originating in the Cambrian Series 2 and becoming extinct in the late Cambrian Series 3, Nisusia existed for a relatively short period of geologic
time, and is a brachiopod genus that exhibited a slow rate of morphological evolution. This paper details the brachiopod Nisusia from the middle Cambrian Kaili
Konservat-Lagerstätte. Nisusia, in terms of abundance, is one of the most important
brachiopods from the Kaili fauna. Two Nisusia species from the Kaili Formation of
the Cambrian Series 2–3 in the eastern part of Guizhou Province are described, Nisusia granosa and Nisusia guizhouensis n. sp. This study will give valuable insights into the morphological variability of Nisusia species, and provides new details on the composition of the brachiopod community in the Kaili Biota.
Keywords: Kaili fauna; Taxonomy; China; Brachiopoda; Lagerstätte
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1. Introduction Named by Walcott in 1905, Nisusia is a calcareous-shelled brachiopod genus found in the Cambrian Series 2–3 (traditionally called the lower and middle
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Cambrian). Nisusia is a relatively short-lived genus, first appearing in the Cambrian
Series 2 and disappearing from the fossil record in the late Cambrian Series 3 (Popov
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and Holmer, 1996; Popov and Williams, 2000). Despite its geologically brief
existence, the genus had a near cosmopolitan distribution in the Cambrian, and has
us
been documented from numerous localities across the Americas, Asia, Europe, and Oceania (Popov and Williams, 2000). Although frequently documented during the 20th
an
century (Walcott, 1912; Størmer, 1925; Schuchert and Cooper, 1931, 1932; Rowell, 1962, 1977) the lack of morphological details regarding the cardinalia area resulted in
M
systematic uncertainties. For a prolonged period of time, many authors considered Nisusia a representative of the Orthida (Williams and Wright, 1965), closely allied
d
with the billingselloids (Walcott, 1912; Schuchert and Cooper, 1931, 1932). It was transferred to the Kutorginida after a study by Rowell and Caruso (1985) that
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questioned the affinities of the genus and later work by Popov and Tikhonov (1990)
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that unveiled the unusual mode of articulation in Nisusia. The peculiar combination of calcareous valves and a distinct, simple type of articulation has led to the suggestion that the group most likely represent a distinct, short-lived, early representative of Rhynchonelliformean brachiopods (Williams et al., 1996; Popov and Williams, 2000). In China, most Cambrian brachiopods belong to the Subphylum Linguliformea
(Inarticulata in the traditional classification), whereas relatively few documented Cambrian brachiopods belong to the Subphylum Rhynchonelliformea (Articulata). Fossils of the Class Obolellata, Class Chileata, and Class Kutorginata of the Subphylum Rhynchonelliformea have been reported from the Chengjiang Lagerstätte of the early Cambrian Series 2. Among them, only fossils of the Family Kutorginidae of the Class Kutorginata, rather than those of the Nisusiidae, were found (Zhang, 2006; Zhang et al., 2007, 2008, 2011a, 2011b; Zhang and Holmer, 2013). Nevertheless, Nisusia specimens (unnamed species, referred to here as Nisusia sp. A) have been 2
Page 2 of 32
reported from the Guanshan Biota in the middle part of the Cambrian Series 2 (Luo et al., 1999, 2008; Hu et al., 2014). In terms of species diversity, brachiopods are the second most prominent phylum in the Kaili Formation (Cambrian Series 3), second only to arthropods (Zhao, 2011). There are 11 genera, including 8 genera of the
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Subphylum Linguliformea and 3 genera of the Subphylum Rhynchonelliformea. The Subphylum Rhynchonelliformea is represented in the Kaili Biota by the families
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Kutorginidae and Nisusiidae of the Class Kutorginata (Huang et al., 1994; Zhao, 2011; Mao et al., 2014). There are two species of Nisusia present. One is Nisusia granosa,
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originally described as Nisusia alaica and recently revised by Mao et al. (2014) based on specimens from the Kaili Formation at the Miaobanpo Section in Balang Village of
an
Jianhe County, Guizhou Province. The two species are morphologically very similar but differ in the thickness of costellae and the size of nodes on the costellae (Mao et
M
al., 2014). The other species was originally described as Nisusia sulcata (Zhao, 2011), with more than 300 specimens collected from sections near Balang Village, but also
d
from those near Zhuping Village (Fujiachong Section locates at southwest of Zhuping Village, about 2 km far away from it), Zhenyuan County (Yang et al., 2011) and near
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Nangao Village, Danzhai County, Guizhou Province (Zhao et al., 2012). The dorsal
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valves of those specimens, described as N. sulcata, do not display a sulcus and the ventral valves bear the maximum height at the apex, characters that are different from Nisusia sulcata and other documented Nisusia species. Thus, it is herein regarded as a
new species and is named Nisusia guizhouensis n. sp. Based on newly discovered
material, this study enhances the diversity of the genus Nisusia and provides new
details on the combination of morphological characteristics exhibited by the genus.
Abbreviations for specimen number: GTBM – Miaobanpo Section; (M) – Miaobanpo Section; (B) – Wuliu-Zengjiaya Section; (S) – Sanwan Section; (F) – Fujiachong Section.
2. Geological background The Kaili Formation, a lithostratigraphic unit of the Cambrian Series 2–3 3
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(traditionally the Lower and Middle Cambrian) (Zhao et al., 1993, 1994; Zhao, 2011; Yuan et al., 1999), is situated in the transitional belt between the Yangtze Platform Region and the Jiangnan Region in eastern Guizhou Province, South China (Fig. 1B) and has been interpreted as a neritic shelf environment in the Cambrian (Lu et al.,
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1982; Yuan et al., 1999; Zhao at al., 2001; Zhao, 2011). The Kaili Biota has been
predominantly documented from the middle–upper Kaili Formation at the Miaobanpo
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Section and the Wuliu-Zengjiaya Section near Balang Village of Jianhe County,
Guizhou Province. The Kaili Formation yielding the Kaili Biota is located primarily
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across Jianhe County, Danzhai County, Zhenyuan County and Tongren City in eastern Guizhou Province (Fig. 1A). The specimens studied in this paper were collected at the
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Miaobanpo Section and the Wuliu-Zengjiaya Section near Balang Village of Jianhe County, the Sanwan Section in Danzhai County and the Fujiachong Section in
M
Zhenyuan County (Fig. 2). The lower Miaobanpo Section is missing sections of the formation due to a fault contact with the Qingxudong Formation and the Fujiachong
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Section of the Kaili Formation has no top or bottom parts of the formations exposed; however, the other two sections have basically complete sequences, which are in
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conformable contact with the overlying and underlying strata. The studied sections
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from the Kaili Formation consist mainly of mudstones, with subordinate medium-bedded limestone, that vary in thickness at the top and base of the formation. (Zhao et al., 1993, 1994, 2012; Yuan et al., 1999, 2002; Yang et al., 2011).
Fig. 1 here
There are three trilobite zones present in the Kaili Formation. In ascending order,
these are the Bathynotus kueichouensis-Ovatoryctocara cf. granulata Zone, the Oryctocephalus indicus Zone, and the Peronopsis taijiangensis Assemblage Zone (Zhao, 2011; Zhao et al., 2012). Most of the Nisusia specimens examined in this study were collected of between 36.74 m and 111.69 m at the Miaobanpo Section in Jianhe County; however, a few specimens were collected from 53.3-164.13 m at the Wuliu-Zengjiaya Section, and 160-280 m at the Sanwan Section in Danzhai County, 4
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in addition to 11.9-106 m at the Fujiachong Section in Zhenyuan County. All specimens are from the Oryctocephalus indicus Zone (Fig. 2).
3. Geographic and stratigraphic distribution of Nisusia
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Fig. 2 here
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Thirty-nine documented species (including the new species described herein) of Nisusia are currently considered valid (Table 1) (Verneuil and Barrande, 1860;
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Billings, 1861; Tate, 1892; Walcott, 1886, 1889, 1905, 1908, 1912, 1924; Sun, 1924; Cooper, 1936; Endo and Resser, 1937; Lermontova, 1940; Bell, 1941; Shaw, 1954,
an
1957; Nikitin, 1956; Aksarina, 1960; Andreeva, 1962; Mount, 1981; Rowell and Caruso, 1985; Zeng, 1987; Popov and Tikhonov, 1990; Roberts and Jell, 1990; Brock,
M
1998; Wotte and Mergl, 2007; Benedetto and Foglia, 2012; Mao et al., 2014; Topper et al., 2014, 2015). The genus had a cosmopolitan distribution in the Cambrian and
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has been documented from the USA, Canada, Mexico, Argentina, Uzbekistan, Kazakhstan, Kyrghyzstan, Bohemia, Siberia, Australia, Spain, and China (Table 1).
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North America tops the list of the Nisusia species distribution, with 17 species found
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in USA and 6 species in Canada, followed by China with 6 species (including the new species documented herein). There are three Nisusia species documented from
Oceania.
Table 1 here
Of the 39 documented species of Nisusia (Table 1), 17 species have been
documented from the Cambrian Series 2 and 22 species from the Cambrian Series 3. Originating in the Cambrian Series 2 and becoming extinct in the Cambrian Series 3, Nisusia existed for a relatively short span of time compared to other brachiopod genera, such as Lingula (Williams et al., 1996). Nisusia sp. A, found in the Guanshan Biota, Wulongjing Formation in the Yunnan Province of China, which is equivalent to the middle Cambrian Series 2 and the Qiandongian Series Early Duyunian Stage, is 5
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considered herein to represent the earliest occurrence of Nisusia in China; the Zhangxia Formation in Luan County of Hebei Province (Sun, 1924) yields the youngest occurrence of Nisusia in China. Nisusia species in the Cambrian of China are distributed mainly in the North
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China Region and the Yangtze Region. A number of Nisusia species were originally documented from the North China Region, comprising N. hayasakai from Luan
cr
County of Hebei Province (Sun, 1924), N. mantaouensis, N. salebrosa, N. orientalis and N. concentrica of Liaoyang County of Liaoning Province (Endo and Resser,
us
1937). Nisusia mantaouensis and N. salebrosa have since been synonymized with N. concentrica (Wang et al., 1966), leaving three valid species of Nisusia in the North
an
China Region. Four Nisusia species (including the unnamed Nisusia sp. A) have been found in the Yangtze Region, including Nisusia liantuoensis in Yichang City of Hubei
M
Province (Zeng, 1987), Nisusia sp. A in Kunming City of Yunnan Province (Luo et al., 2008; Hu et al., 2014), N. granosa (Mao et al., 2014) and N. guizhouensis n. sp. from
d
the middle–upper part of the Kaili Formation of Balang Village in Jianhe County of Guizhou Province. Nisusia granosa and N. guizhouensis n. sp. are the focus of this
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paper. Some of the N. guizhouensis n. sp. specimens were also found in the
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Qingxudong Formation beneath the Kaili Formation (Zhao et al., 2012).
4. Morphological characteristics and morphological variation of Nisusia Except for N. granosa (Mao et al., 2014), N. ancauchensis (Benedetto and Foglia,
2012) and N. guizhouensis n. sp. (documented here) all of the 39 Nisusia species were reported and described during the 20th century. More than half of those descriptions were published prior to 1950. Owing to the oversimplified descriptions of some of the species and the limited number of fossil images illustrated, some specimens could not be assigned to the genus with great certainty and consequently the statistical work undertaken herein includes only 17 species in this paper (Table 2). During the Cambrian Series 2–3, Nisusia exhibited only minor morphological changes and can be regarded as a comparatively stable group. The stability and variability of Nisusia are
described as follows: 6
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Stability: Overall many morphological features of Nisusia remained relatively stable in the Cambrian. During the Cambrian Series 2–3, little morphological change is evident in Nisusia, and features such as the biconvex shells, anacline dorsal interarea, ornamentation of costellae, straight hinge line and the sulcus-bearing ventral
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valve remained the same for the majority of species. However, subtle morphological
variations within the genus have allowed Nisusia to be differentiated into two species
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groups. Bell (1941) recognized morphological disparate groups of Nisusia on the
basis of North American species and his ideas have been expanded upon here. Group
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A, consists of N. alaica (Cambrian Series 2), N. ancauchensis and N. guizhouensis n. sp. (Cambrian Series 2–3), and N. metula and N. grandis grandis (Cambrian Series 3).
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All members of Group A are characterized by apsacline to catacline interareas and an apex occupied at the maximum height of the ventral valve. Group B, includes N.
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festinata, N. kotujensis and N. fulleri (Cambrian Series 2), N. borealis, N. lickensis, N. brugessensis and N. hayasakai (Cambrian Series 3) and are all characterized by an
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Fig. 3 here
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apsacline interarea and an apex occupying the maximum height of the ventral valve.
Variability: In the Cambrian Series 2, three types of ventral valve interarea are
found: apsacline, catacline, and procline, whereas in the Cambrian Series 3, only two types of ventral valve interareas were present: apsacline and catacline, indicating a decreasing trend in the variation of interarea type. The shell convexity varied from unequally biconvex in the Cambrian Series 2 to unequally biconvex and equally biconvex in the Cambrian Series 3. The maximum height of the ventral valve was always at the apex in the Cambrian Series 2, but it appeared sometimes at the umbo between the apex and the middle of the valve in the Cambrian Series 3 (see type B1 in Fig. 3). In addition to this evolutionary trend, the variability of Nisusia lies in certain
evolutionary relationships between various species groups. For example, a variation trend type A1 (Fig. 3), which exhibits a procline interarea with the maximum height of the ventral valve at the apex, only existed in the Cambrian Series 2, consisting of 7
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species such as in N. liantuoensis. Type B1, another variation trend, displays apsacline interareas, with the maximum height of the ventral valve at the near posterior margin instead of the apex, and only existed in the Cambrian Series 3, including species such
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as N. deissi, N. fulleri and N. montanaensis.
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Table 2 here
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5. Systematic palaeontology
Subphylum Rhynchonelliformea Williams, Carlson, Brunton, Holmer and Popov,
an
1996
Class Kutorginata Williams, Carlson, Brunton, Holmer and Popov, 1996
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Order Kutorginida Kuhn, 1949
Superfamily Nisusioidea Walcott and Schuchert
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Genus Nisusia Walcott, 1905
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Family Nisusiidae Walcott and Schuchert
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Type species: Orthisina festinata Billings, 1861. Diagnosis: Shell subequally biconvex or unequally biconvex (ventribiconvex or
dorsibiconvex); ventral valve elevated and frequently more convex than dorsal; transverse subrectangular or semicircular in outline; anterior commissure rectimarginate or slightly sulcate; ventral interarea raised and triangular in shape, characterized as either apsacline, catacline or procline; basically anacline dorsal interarea, with the apex small and curled towards the posterior margin; dorsal interior with transverse, elevated notothyrial platform; dorsal adductor muscle scars may be discretely impressed on two pairs by prominent ridges; shell radially ornamented by costellae, with nodes or spines on some costae. Remarks: Andreeva (1987) described a new genus of Nisusiid, Narynella Andreeva,
1987 (type species Nisusia ferganensis Andreeva, 1962) from the Cambrian (Lenan) of Kyrgyzstan. The new genus was erected to distinguish brachiopod specimens that 8
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lack spinose costellae and exhibit a unisulcate commissure from the spinose, rectimarginate Nisusia (Andreeva, 1987; Popov and Williams, 2000). Not all Nisusia species definitively possess spines (Benedetto and Foglia, 2012) and few species of Narynella have been documented since for comparison (Geyer et al., 2014). Geyer et
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al. (2014) considered that the nature of the commissure may be the only
distinguishing feature between the two genera, combined with the confusion about the
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age and origin of the type material (see Geyer et al., 2014) and the fact that selected
Narynella is in need of clarification and revision.
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species of Nisusia are sulcate (Popov and Williams, 2000), and implied that the genus
Bell (1941) considered that the majority of Nisusia species could be divided into
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two groups: one group consisting of taxa that possess a hemiconical ventral valve with the highest point at the apex, such as N. festinata (Billings, 1861), whereas the other
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group is characterized by a convex ventral valve with an incurved beak and maximum height near the middle of the valve, such as N. sulcata (Rowell and Caruso, 1985) and
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N. deissi (Bell, 1941). The specimens under consideration here exhibit the highest point of their ventral valves at the apex and belong to the former group.
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Occurrence: Cambrian Series 2–3; North America, Asia, Oceania, and Europe.
Nisusia guizhouensis n. sp.
(Figs. 4, 5)
2011
Nisusia sulcata – Zhao, p. 107, figs. 102-106.
Etymology: guizhouensis, derived from the locality of the new species in Guizhou
Province. Materials: 300 specimens in total, including specimens consisting of external and internal molds of both the ventral and dorsal valves. All specimens are reposited in the Institute of Palaeontology and Biological Mineralization, Guizhou University (GUK). Types: Holotype, GUK(M)-201 (Fig. 4a), ventral valve, valve length 11.06 mm, valve width 14.89 mm. Paratypes, GUK(M)-202 (Fig. 4b), ventral valve, valve length 6.50 9
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mm, valve width 9.37 mm; GUK(M)-203 (Fig. 4c), dorsal valve, valve length 7.21 mm, valve width 11.64 mm; GUK(M)-204 (Fig. 4d), dorsal valve, valve length 8.90 mm, valve width 14.61 mm. Diagnosis: Shell large, relatively for the genus, transverse rectangular in outline,
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unequally biconvex in lateral view; surface radially ornamented by costellae,
increasing in number by bifurcations in the middle and front of shell surface;
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prominent concentric lines visible between the middle part and the anterior margin of each valve; ventral interarea apsacline to catacline, maximum height at the apex of
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ventral which is fine and convex, with a sulcus; dorsal interarea anacline.
Description: Shell large, relative for the genus, valve length shorter than valve width;
an
holotype shell length 11.06 mm, shell width 14.89 mm; outline transversely subrectangular; hinge line straight, equal to the greatest shell width, cardinal
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extremities near right-angle; unequally biconvex in lateral profile, convexity of the dorsal valve slightly larger than that of the ventral valve; maximum height of the
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ventral valve at apex, shallow and wide sulcus originating from umbo to the anterior part of valve; hinge area high and triangular; anterior commissure mostly
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rectimarginate or rarely slightly sulcate; thick costellae ornamented on shell surface,
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increasing in number by bifurcations in the middle and front of shell surface, in the adult phase about 7-9 costellae of 3 mm along the anterior margin; concentric lines prominent from the middle to the anterior margin of each valve in the adult phase. Ventral valve (Figs. 4a, b, f, j-o, 5j-r): moderately convex, maximum height at the
apex which is pointed, small and raised; outline transversely subrectangular; Interarea apsacline to catacline; in the central valve surface, a wide and shallow sulcus originating from umbo to anterior margin; muscle scars obscure; interior structure not clear. Dorsal valve (Figs. 4c-e, g-i, 5a-i): slightly more convex than ventral valve, apex incurved, umbo swollen, without fold, maximum height at about one-third of the valve length in front of umbo, interarea anacline; muscle scars obscure; interior structure not clear. Young stage (Fig. 5a-c, j-l): valve length less than 4 mm, width less than 6 mm; 10
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costae distinct, without bifucated costellae or concentric lines; apex pointed and projecting perpendicular to shell surface; sulcus obscure. Adolescent stage (Fig. 5d-f, m-o): valve length 4-6 mm, width 6-9 mm; clear costae on valve, with bifurcated costellae and few concentric lines appearing in the
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anterior margin of each valve; apex pointed, small and straight; sulcus distinct.
Adult stage (Fig. 5g-i, p-r): more than 6 mm in length of valve, more than 9 mm
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in width; clear costae on valve, with bifurcated costellae and strong concentric lines appearing in the anterior margin of each valve; apex pointed, small and straight; a
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wide and shallow sulcus originating from umbo to the anterior margin.
Remarks: Specimens of Nisusia guizhouensis n. sp. from the Cambrian of China
an
were originally described as Nisusia sulcata (Zhao, 2011). The absence of a sulcus on the dorsal valve and the maximum height of the ventral valve occurring at the apex
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rather than the center of the valve clearly distinguish N. guizhouensis n. sp. from N. sulcata. Nisusia guizhouensis n. sp. also has a similar shell shape to Nisusia granosa
d
from the Kaili Formation at the Miaobanpo Section of Guizhou Province (Mao et al., 2014, p. 232, pl. 1). However, N. guizhouensis n. sp. possesses thick and convex
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costae lacking nodes, whereas Nisusia granosa has fine and rounded costae bearing
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many small nodes. In addition, Nisusia guizhouensis n. sp. has stronger concentric lines than N. granosa.
N. grandis glabra from the Cambrian Series 2–3 of Australia (Roberts and Jell,
1990, p. 266, pl. 7, figs. A-Y) is similar to the species from Guizhou in both shell shape and the presence of well defined concentric growth lines. But N. grandis glabra tends to be generally alate in outline, has thicker costellae, and lacks the pointed apex (Fig. 4n-o) when compared with the species from Guizhou. As for the shell shape and costellae ornamentation, Nisusia guizhouensis n. sp.
closely resembles N. festinata from the Cambrian Series 2 of North America (Walcott, 1912; Cooper, 1936, p. 212, pl. 26, figs. 33, 39-42, 47) and N. borealis from the Middle Cambrian in Alaska (Cooper, 1936, p. 212, pl. 26, figs. 22, 23, 25-26, 28, 30-32, 34-35, 37-38, 43-44, 46, 48-50). Although these three species all have a sulcus on the ventral valve, Nisusia guizhouensis n. sp. possesses a sulcus on the ventral 11
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valve that extends from the umbo to the anterior margin, whereas the other two species have a sulcus extending from umbo to the middle part of the valve. Additionally, both Nisusia festinata and Nisusia borealis have nodes on their costellae, which are absent in the new species.
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Occurrence: The middle–upper part of Kaili Formation of the Cambrian Series 2–3 of Balang Village in Jianhe County, Sanwan Village in Danzhai County, Zhuping
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Village in Zhenyuan County, Guizhou Province; the middle–upper part of the
Nisusia granosa Mao, Wang and Zhao, 2014
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(Fig. 6)
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Balang Village, Jianhe County, Guizhou Province.
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Qingxudong Formation of the Cambrian Series 2 Stage 4 at the Songshan Section near
Nisusia alaica Popov and Tikhonov – Zhao, p. 107, figs. 107, 108.
2014
Nisusia granosa – Mao et al., p. 223, pl. 1, figs. 1-15.
d
2011
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Materials: 15 specimens in total (including 8 complete specimens).
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GTBM-9-4-21597 is the holotype (Fig. 6a; also Mao et al., 2014, pl. 1, fig. 1), ventral valve, valve length 10.12 mm, valve width 14.30 mm. Diagnosis: Shell relatively large for the genus, outline transversely rectangular, lateral
view unequally biconvex; shell radially ornamented with fine, rounded and low costae, increasing in number by intercalations or bifurcations in the middle and front of shell surface; many nodes exist on the costellae; ventral valve with a fine and pointed apex at its maximum height, bearing a sulcus. Description: Shell relatively large for the genus, valve length shorter than valve width; holotype shell length 10.12 mm, shell width 14.30 mm; outline transversely rectangular, hinge line straight, equal to the greatest shell width, cardinal extremities near right-angle or acute angle; lateral view unequally biconvex, convexity of the dorsal valve slightly larger than that of the ventral valve; ventral valve with maximum height at the apex, possessing a shallow and wide sulcus originating from umbo to 12
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anterior margin; hinge area high and triangular; anterior commissure mostly rectimarginate or slightly sulcate; fine ornamented costae on shell surface, increasing in number by intercalations or bifurcations in the middle and front of shell surface in the adult phase about 6-7 costellae of 3 mm along the anterior margin; many tiny
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nodes appearing on costellae, concentric growth lines visible in anterior margin.
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Table 3 here
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Ventral valve (Fig. 6a, d-f, h-i): moderately convex, maximum height at the apex which is pointed, small and raised; outline transversely rectangular; interarea
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apsacline to catacline; in the central valve surface, a wide and shallow sulcus originating from umbo to anterior margin; valve surface radially ornamented by fine
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and close growth lines, increasing in number by intercalations or bifurcations towards anterior margin, interarea apsacline to catacline; interior structure not clear.
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Dorsal valve (Fig. 6b, c, g): slightly more convex than ventral valve, umbo swelling, without fold, maximum height at about one-third of the valve length in front
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not clear.
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of umbo, apex incurved, interarea anacline; muscle scars obscure; interior structure
Comparison: Nisusia granosa Mao, Wang and Zhao, 2014 closely resembles Nisusia
alaica from the Cambrian Series 2, found in south Kirgizia (Popov and Tikhonov, 1990, p. 22, pl. 3, figs. 23, 24) in shell shape, costae furcations and ornamentation of nodes and spines. Nisusia granosa Mao, Wang and Zhao, 2014 however tends to be
larger (holotype shell width 14.30 mm and length 10.12 mm) than Nisusia alaica (holotype shell width 9.0 mm and length 6.2 mm), and has finer costellae with a higher density on shell surface. The ornamentation of Nisusia granosa Mao, Wang
and Zhao, 2014 is dominated by faint concentric lines, morphological features absent in N. alaica. Nisusia granosa Mao, Wang and Zhao, 2014 appears to bear a close resemblance to N. lickensis from the Cambrian Series 3 of Montana (Bell, 1941, pl. 32, figs. 9-12) in shell shape and the presence of costellae spines. Nisusia granosa differs from N. 13
Page 13 of 32
lickensis in being larger and in possessing a median sulcus on the ventral valve from the umbo to anterior margin. Nisusia granosa Mao, Wang and Zhao, 2014 is similar to N. festinata from the Cambrian Series 2, of North America (Walcott, 1912, pl. 100, fig. le) in features such
ip t
as shell shape, costellae and nodes on the costellae. Though they both have a sulcus
on the ventral valve, the sulcus on the ventral valve of N. festinata extends only from
cr
the umbo to the central part of the valve. The biggest difference between these two species is the absence of a sulcus on the dorsal valve of Nisusia granosa.
us
Nisusia granosa is also similar to N. kotujensis from the Cambrian Series 2–3 found from Siberia (Andreeva, 1962, p. 88) in shell shape, but it is significantly larger
an
and the lack of a detailed description makes comparison difficult.
Occurrence: Middle–upper part of the Kaili Formation of the Cambrian Series 2–3
M
(Wuling Series) Stage 5 (Taijiangian Stage), at the Miaobanpo Section in Balang
Acknowledgments
d
Village of Jianhe County, Guizhou Province.
te
Special thanks are given to Prof. Lars Holmer from Uppsala University (Sweden) and
Ac ce p
Prof. Zhi-Fei Zhang from Northwest University (China) for their valuable instructions and suggestions for this paper, Academician Xu Chen and Prof. Mao-Yan Zhu from Nanjing Institute of Geology and Palaeontology, China Academy of Sciences for their help and financial support of the fossil specimen photographs in this paper, Prof. Jin Peng from Guizhou University for her instructions and guidance, and Dr. Hai-Zhou Wang from Uppsala University (Sweden) for helping take photographs of some fossils in this paper. This paper benefited greatly from constructive reviews by Prof. Zhi-Fei Zhang, Christian Skovsted, and an anonymous reviewer. This study is supported by grants from the Major State Basic Research Development Program of China (973 program (N): 2013CB835002), the National Natural Science Foundation of China (41372019,40762018, 41362002, 41172005, 41330101) to Wang, Zhao, Peng, Yang; The foundation of Chinese Committee of Stratigraphy, Foundation of Guizhou Science Cooperation foreign (Gscf.G2010-70001) to Zhao; Guizhou Engineering and 14
Page 14 of 32
Technology Research Center for Development and Utilization of Karst Cave (Tourism) Resources (G [2014] 4004).
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brachiopods: A perspective from South China. Gondwana Research 14, 241-254. Zhang, Z.F., Holmer, L.E., Popov, L.E., Shu, D.G., 2011a. An obolellate brachiopod with soft-part preservation from the early Cambrian Chengjiang Fauna of China. Journal of Paleontology 85 (3), 460-463.
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Zhang, Z.F., Holmer, L.E., Ou, Q., Han, J., Shu, D.G., 2011b. The exceptionally
preserved Early Cambrian stem rhynchonelliform brachiopod Longtancunella
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and its implications. Lethaia 44, 490-495.
Zhao, Y.L., 2011. The Kaili Biota — Marine Organisms from 508 Million Years Ago.
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Guizhou Science and Technology Publishing House, Guiyang, 251 pp. (in Chinese).
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Zhao, Y.L., Yuan, J.L., Zhang, Z.H., Mao, J.R., Huang, Y.Z., Gong, X.Y., Wang, K., 1993. A preliminary study of the Kaili Formation in the transitional area in south
Chinese, with English abstract).
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China and its synchronous strata. Journal of Stratigraphy 17 (3), 170-178 (in
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Zhao, Y.L., Yuan, J.L., Huang, Y.Z., Mao, J.R., Qian, Y., Zhang, Z.H., Gong, X.Y., 1994. Middle Cambrian Kaili Fauna in Taijiang, Guizhou. Acta Palaeontologica
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Zhao, Y.L., Yu, Y.Y., Yuan, J.L., Yang, X.L., Guo, Q.J., 2001. Cambrian stratigraphy at Huanglian, Guizhou Province, China: Reference section for bases of the Nangaoan and Duyunian Stages. Palaeoworld 13, 172-180.
Zhao, Y.L., Peng, J., Yuan, J.L, Babcock, L.E., Guo, Q.J., Yin, L.M., Yang, X.L., Tai, T.S., Wang, C.J., Lin, J.P., Gaines, R.R., Sun, H.J., Yang, Y.L., 2012. Discussion of candidate stratotypes for the GSSP defining the conterminous base of Cambrian provisional Series 3and Stage 5. Journal of Guizhou University 29 (Supp. 1), 35-48.
Figure captions
20
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Fig. 1. (A) Location map of the study area in Guizhou. (B) Cambrian outcrops in the transitional belt of east Guizhou, South China (the shaded part of the map is adapted from Yuan et al., 2002).
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Fig. 2. Stratigraphic column of the Kaili Formation showing the distribution of
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Nisusia species in east Guizhou Province.
us
Fig. 3. Morphological variation of Nisusia.
Fig. 4. Nisusia guizhouensis n. sp. Except where indicated, scale bars represent 5 mm.
an
(a-f, h-o) Specimens collected from the Kaili Formation in Balang Village of Jianhe County, in Guizhou Province, South China; (a) GUK(M)-201, holotype, external view
M
of ventral valve; (b) GUK(M)-202 external view of ventral valve; (c) GUK(M)-203, external view of dorsal valve; (d) GUK(M)-204, external view of dorsal valve; (e)
d
GUK(M)-204, magnifying posterior margin of dorsal valve; (f) GUK(M)-205, external view of ventral valve, exhibiting sulcus from umbo to anterior part of valve;
te
(h) GUK(M)-207, external view of dorsal valve; (i) GUK(B)-208, external view of
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dorsal valve; (j) GUK(M)-209, external view of ventral valve; (k) GUK(M)-209, external view of ventral valve (scanning electron microscope); (l) GUK(M)-209, external view of ventral valve (scanning electron microscope) with magnified apex of ventral valve; (m) GUK(M)-210, external view of ventral valve; (n) GUK(M)-210, anterior view of ventral valve exterior, displaying the pointed convex beak of the shell; (o) GUK(M)-210, lateral view of ventral valve exterior, displaying the pointed convex beak of the shell. (g) Specimen collected from the Kaili Formation in Zhuping Village of Zhenyuan County, in Guizhou Province, South China, GUK(F)-206, external view of dorsal valve.
Fig. 5. Nisusia guizhouensis n. sp. Scale bars represent 2 mm for (a-c, j-l) (young stage); 3 mm for (d-f, m-o) (adolescent stage); 5 mm for (j-i, p-r) (adult stage). (a, k, n) Specimens collected from the Kaili Formation in Zhuping Village of Zhenyuan 21
Page 21 of 32
County, in Guizhou Province, South China; (a) GUK(F)-211, external view of dorsal valve; (k) GUK(F)-221, external view of ventral valve; (n) GUK(F)-224, external view of ventral valve. (b-g, i, j, l, m, o-r) Specimens collected from the Kaili Formation in Balang Village of Jianhe County, in Guizhou Province, South China; (b)
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GUK(M)-212, external view of dorsal valve; (c) GUK(M)-213, external view of
dorsal valve; (d) GUK(M)-214, external view of dorsal valve; (e) GUK(M)-215,
cr
external view of dorsal valve; (f) GUK(M)-216, external view of dorsal valve; (g)
GUK(M)-217 external view of dorsal valve; (i) GUK(M)-219, external view of dorsal
us
valve; (j) GUK(M)-220, external view of ventral valve; (l) GUK(M)-222, external view of ventral valve; (m) GUK(M)-223, external view of ventral valve; (o)
an
GUK(M)-225, external view of ventral valve; (p) GUK(M)-226, external view of ventral valve; (q) GUK(M)-227, external view of ventral valve; (r) GUK(M)-228,
M
external view of ventral valve. (h) Specimen collected from the Kaili Formation in Sanwan Village of Danzai County, in Guizhou Province, South China, GUK(S)-218,
d
external view of dorsal valve.
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Fig. 6. Nisusia granosa. Except where indicated, scale bars represent 3 mm. All
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specimens were collected from the Kaili Formation in Balang Village of Jianhe County, in Guizhou Province, South China. (a) GTBM-9-4-21597, holotype, external view of ventral valve. (b) GTBM-9-5-6200, external view of dorsal valve. (c) GTBM-9-3-5362, external view of dorsal valve. (d-f) GTBM-9-3-3757, ventral valve; (d) external view; (e) shell showing posterior margin (scanning electron microscope); (f) magnified spines of surface valve (scanning electron microscope). (g) GTBM-9-5-2668, external view of dorsal valve. (h, i) GTBM-9-2-3521, ventral valve; (h) external view, showing pointed beak of shell; (i) lateral view, showing sulcus from umbo to anterior part of valve.
22
Page 22 of 32
Table 1. Geological era and occurrence of Nisusia species. Cambrian Series
Species name
Second
Location
Third
N. deissi (Bell, 1941)
×
USA
N. lickensis (Bell, 1941)
×
USA
N. montanaensis (Bell, 1941)
×
USA
N. borealis (Cooper, 1936)
×
USA
N. sulcata (Rowell and Caruso ,1985)
×
USA
N. fulleri (Mount, 1981)
USA
×
USA
N. erecta (Walcott, 1905)
× × ×
×
N. transversa (Walcott, 1886)
×
N. bivia (Walcott, 1912)
×
an
N. argenta (Walcott, 1905)
N. spencei (Walcott, 1905) N. nautes (Walcott, 1905) N. kanakensis (Walcott, 1908) N. orientalis (Endo and Resser, 1937) N. hayasakai (Sun, 1924)
te
N. liantuoensis (Zeng, 1987)
d
N. concentrica (Endo and Resser, 1937)
M
N. rara (Walcott, 1908)
Ac ce p
N. amii (Walcott, 1905)
USA
×
USA
×
USA
×
USA
×
USA
×
China
×
China
×
China China
× ×
N. alberta (Walcott, 1889) N. oriens (Walcott, 1924)
USA
×
N. guizhouensis n. sp.
USA
USA USA
×
N. granosa (Mao, Wang and Zhao, 2014)
USA
us
N. utahensis (Walcott, 1905) N. cloudi (Shaw, 1954)
ip t
×
cr
N. festinate (Billings, 1861)
China China Canada
× ×
Canada Canada
N. spinigera (Walcott, 1924)
×
Canada
N. burgessensis(Walcott, 1924)
×
Canada
N. lowi (Walcott, 1908)
×
Canada
N. grandis grandis (Roberts and Jell, 1990)
×
Australia
N. metula (Brock, 1998)
×
Australia
×
Australia
N. compta (Tate, 1892)
N. kotujensis (Andreeva, 1962)
×
N. nasuta (Nikitin, 1956)
×
N. alaica (Popov and Tikhonov, 1990)
×
Kyrgyzstan
N. pospelovi (Aksarina, 1960)
×
Siberia
N. minussensis (Lermontova, 1940)
×
N. vaticina (Verneuil and Barrande, 1860) N. ancauchensis (Benedetto and Fogli, 2012)
×
Siberia ×
Kazakhstan
Siberia ×
Spain
×
Argentina
23
Page 23 of 32
Table 2. Morphological analysis and comparison of Nisusia species.
Hinge line
Anterior commissure Sulcus
Maximum height
Foramen
Costellae
TR0
0
0
1
0(V) 0(D)
1
0
0
N. montanaensis
3(M)
0
1
2
0
1
0(V) 0(D)
1
0
0
N. sulcata
3(M)
0
T1
0
0
1
0(D)
0(V)
1
0
N. grandis grandis
3(M)
1(D)
0
2
1
0(V)
0
0
N. lickensis
3(M)
1
T0
0
0
N. borealis
3(M)
1(V)
?
0, 1
0
N. metula
3(M)
1
0, 3
N. burgessensis
3(M)
1(V)
N. hayasakai
3(M)
1
2
N. granosa
2~3
1(D)
T1
N. guizhouensis n. sp.
2~3
1
N. liantuoensis N. alaica N. festinata N. fulleri N. ancauchensis
1
17(V)
0
0
1
18.1(V)
0
0
0
1
0
0(S)
0
0 to 2
1
0
0(T)
1
0
0(V) 0(D)
1
0
1
1
0
0(T)
0
2
0(V)
0
0
1
0(V)
0
0
0(V)
0
0
1
0
d
1
1
2
2
1
2(L)
1(V)
2
1
1
1
2(L)
1
0
1
1
2(L)
1
1
2
1
0(V) 0(D)
0
2(L)
1
T0
1
1
0(V)
0
2(L)
1
0
2
1
0(V)
0
0
0
cr
us 0
0
2(L)
ip t 0
0(V)
te
1
1
0
0
an
0
0 1, 2
0(V)
M
0
Ac ce p
N. kotujensis
1
Dorsal
Cardinal angles
1(D)
Maximum width (mm)
Ventral
Outline
3(M)
Interarea
Growth line
Shell shape
N. deissi
Fold
Age
Ornamentation
0
11.8(D)
1
0(S)
5 23(D)
0(P)
0
0(P)
0
9(V) 0 to 2
1
0
1
14.3(V)
0(P)
0
0
1
10.5(D)
0
0
3
1
8~14
0 to 2
1
9.0(V)
1
0(S) 0
0
0
0
0
0
0
0
0
0
2
15
Note: morphological information used in this table: V = ventral valve; D = dorsal valve. Age: 2 = Cambrian Series 2, 3 = Cambrian Series 3, M = middle, L = lower. Common shell characters: Shell shape: 0 = almost equally biconvex, 1 = unequally biconvex, 1(V) = unequally biconvex, the 24
Page 24 of 32
ventral valve is more convex than the dorsal valve, 1(D) = unequally biconvex, the dorsal valve is more convex than the ventral valve. Outline: 0 = subrectangular, 1 = subquadrate, 2 = semielliptical, 3 = subcircular, T =
transversely subrectangular, TR0 = transversely rounded subrectangular.
line: 0 = hinge line less than greatest width of shell, 1 = hinge line equal
cr
Cardinal angles: 0 = obtusely angle, 1 = right angle, 2 = acute angle. Hinge
ip t
transversely, R = rounded, e.g., T1 = transversely subquadrate, T0 =
greatest width of shell. Anterior commissure: 0 = rectimarginate, 1 = sulcate.
us
Sulcus: 0 = present, 1 = absent, 0(V) = sulcus of ventral valve, 0(D) =
sulcus of dorsal valve. Fold: 0 = present, 1 = absent, 0(V) = fold of ventral
an
valve. Maximum ventral valve height: 0 = in the apex, 1 = in near the
middle of valve. Foramen: 0 = present. Costae: 0 = present, 1 = absent, 0(S)
M
= 0(Spines), 0(T) = 0(Tubercle), 0(P) = 0(Pustula). Growth line: 0 = present, 1 = absent. Interarea: 0 = apsacline, 1= anacline, 2 = catacline, 3 = procline.
Ac ce p
te
show no description in this paper.
d
Width: W(V) = width of ventral valve, W(V) = width of dorsal valve. Space
25
Page 25 of 32
Table 3. Measurements of Nisusia granosa.
Dorsal valve
W (mm)
L (mm)
N
Cw (mm)
Sw (mm)
GTBM9-1-1128
15.56
10.73
7
0.25
0.25
GTBM 9-2-3521
15.41
10.65
6
0.26
0.26
GTBM 9-4-21597
14.3
10.12
6
0.28
0.29
GTBM 9-3-1132
12.66
10.02
6
0.25
0.27
GTBM 9-5-5963
12.34
9.67
7
0.26
0.27
GTBM 9-3-5362
14.93
11.01
6
0.26
0.26
GTBM 9-5-6200
13.27
8.17
7
0.28
0.28
GTBM 9-3-4127
8.96
6.63
7
0.26
0.26
13.4288
9.625
6.5
0.2625
Average value
ip t
Ventral valve
Specimen number
cr
Shell type
0.2675
Note: W – width of valve, L – length of valve, N – number of valve costae per 3 mm, Cw – width of Costellae, Iw
Ac ce p
te
d
M
an
us
– interspaces width of between lines.
26
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S1871-174X(16)00021-4 http://dx.doi.org/doi:10.1016/j.palwor.2016.03.001 PALWOR 354
To appear in:
Palaeoworld
Received date: Revised date: Accepted date:
23-6-2015 4-1-2016 8-3-2016
Please cite this article as: Mao, Y.-Q., Zhao, Y.-L., Wang, C.-W., Topper, T.,A fresh look at Nisusia Walcott, 1905 from the Cambrian Kaili Formation in Guizhou, Palaeoworld (2016), http://dx.doi.org/10.1016/j.palwor.2016.03.001 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
A fresh look at Nisusia Walcott, 1905 from the Cambrian Kaili Formation in Guizhou
a
ip t
Yong-Qin Mao a, b, Yuan-Long Zhao c *, Cheng-Wen Wang a, Timothy Topper d
College of Earth Sciences, Jilin University, 130061 Changchun, Jilin Province, P.R.
b
cr
China
Institute of Mountain Resources, Guizhou Academy of Sciences, 550001 Guiyang,
College of Resources and Environmental Engineering, Guizhou University, 550025
Guiyang, Guizhou Province, P.R. China d
an
c
us
Guizhou Province, P.R. China
Department of Palaeobiology, Swedish Museum of Natural History, P.O. Box 50007,
M
SE-104 05, Stockholm, Sweden
te
Abstract
d
* Corresponding author. E-mail address: [email protected]
Ac ce p
Nisusia Walcott, 1905 is an early genus of kutorginid brachiopod of the Subphylum Rhynchonelliformea. Originating in the Cambrian Series 2 and becoming extinct in the late Cambrian Series 3, Nisusia existed for a relatively short period of geologic
time, and is a brachiopod genus that exhibited a slow rate of morphological evolution. This paper details the brachiopod Nisusia from the middle Cambrian Kaili
Konservat-Lagerstätte. Nisusia, in terms of abundance, is one of the most important
brachiopods from the Kaili fauna. Two Nisusia species from the Kaili Formation of
the Cambrian Series 2–3 in the eastern part of Guizhou Province are described, Nisusia granosa and Nisusia guizhouensis n. sp. This study will give valuable insights into the morphological variability of Nisusia species, and provides new details on the composition of the brachiopod community in the Kaili Biota.
Keywords: Kaili fauna; Taxonomy; China; Brachiopoda; Lagerstätte
Page 1 of 32
1. Introduction Named by Walcott in 1905, Nisusia is a calcareous-shelled brachiopod genus found in the Cambrian Series 2–3 (traditionally called the lower and middle
ip t
Cambrian). Nisusia is a relatively short-lived genus, first appearing in the Cambrian
Series 2 and disappearing from the fossil record in the late Cambrian Series 3 (Popov
cr
and Holmer, 1996; Popov and Williams, 2000). Despite its geologically brief
existence, the genus had a near cosmopolitan distribution in the Cambrian, and has
us
been documented from numerous localities across the Americas, Asia, Europe, and Oceania (Popov and Williams, 2000). Although frequently documented during the 20th
an
century (Walcott, 1912; Størmer, 1925; Schuchert and Cooper, 1931, 1932; Rowell, 1962, 1977) the lack of morphological details regarding the cardinalia area resulted in
M
systematic uncertainties. For a prolonged period of time, many authors considered Nisusia a representative of the Orthida (Williams and Wright, 1965), closely allied
d
with the billingselloids (Walcott, 1912; Schuchert and Cooper, 1931, 1932). It was transferred to the Kutorginida after a study by Rowell and Caruso (1985) that
te
questioned the affinities of the genus and later work by Popov and Tikhonov (1990)
Ac ce p
that unveiled the unusual mode of articulation in Nisusia. The peculiar combination of calcareous valves and a distinct, simple type of articulation has led to the suggestion that the group most likely represent a distinct, short-lived, early representative of Rhynchonelliformean brachiopods (Williams et al., 1996; Popov and Williams, 2000). In China, most Cambrian brachiopods belong to the Subphylum Linguliformea
(Inarticulata in the traditional classification), whereas relatively few documented Cambrian brachiopods belong to the Subphylum Rhynchonelliformea (Articulata). Fossils of the Class Obolellata, Class Chileata, and Class Kutorginata of the Subphylum Rhynchonelliformea have been reported from the Chengjiang Lagerstätte of the early Cambrian Series 2. Among them, only fossils of the Family Kutorginidae of the Class Kutorginata, rather than those of the Nisusiidae, were found (Zhang, 2006; Zhang et al., 2007, 2008, 2011a, 2011b; Zhang and Holmer, 2013). Nevertheless, Nisusia specimens (unnamed species, referred to here as Nisusia sp. A) have been 2
Page 2 of 32
reported from the Guanshan Biota in the middle part of the Cambrian Series 2 (Luo et al., 1999, 2008; Hu et al., 2014). In terms of species diversity, brachiopods are the second most prominent phylum in the Kaili Formation (Cambrian Series 3), second only to arthropods (Zhao, 2011). There are 11 genera, including 8 genera of the
ip t
Subphylum Linguliformea and 3 genera of the Subphylum Rhynchonelliformea. The Subphylum Rhynchonelliformea is represented in the Kaili Biota by the families
cr
Kutorginidae and Nisusiidae of the Class Kutorginata (Huang et al., 1994; Zhao, 2011; Mao et al., 2014). There are two species of Nisusia present. One is Nisusia granosa,
us
originally described as Nisusia alaica and recently revised by Mao et al. (2014) based on specimens from the Kaili Formation at the Miaobanpo Section in Balang Village of
an
Jianhe County, Guizhou Province. The two species are morphologically very similar but differ in the thickness of costellae and the size of nodes on the costellae (Mao et
M
al., 2014). The other species was originally described as Nisusia sulcata (Zhao, 2011), with more than 300 specimens collected from sections near Balang Village, but also
d
from those near Zhuping Village (Fujiachong Section locates at southwest of Zhuping Village, about 2 km far away from it), Zhenyuan County (Yang et al., 2011) and near
te
Nangao Village, Danzhai County, Guizhou Province (Zhao et al., 2012). The dorsal
Ac ce p
valves of those specimens, described as N. sulcata, do not display a sulcus and the ventral valves bear the maximum height at the apex, characters that are different from Nisusia sulcata and other documented Nisusia species. Thus, it is herein regarded as a
new species and is named Nisusia guizhouensis n. sp. Based on newly discovered
material, this study enhances the diversity of the genus Nisusia and provides new
details on the combination of morphological characteristics exhibited by the genus.
Abbreviations for specimen number: GTBM – Miaobanpo Section; (M) – Miaobanpo Section; (B) – Wuliu-Zengjiaya Section; (S) – Sanwan Section; (F) – Fujiachong Section.
2. Geological background The Kaili Formation, a lithostratigraphic unit of the Cambrian Series 2–3 3
Page 3 of 32
(traditionally the Lower and Middle Cambrian) (Zhao et al., 1993, 1994; Zhao, 2011; Yuan et al., 1999), is situated in the transitional belt between the Yangtze Platform Region and the Jiangnan Region in eastern Guizhou Province, South China (Fig. 1B) and has been interpreted as a neritic shelf environment in the Cambrian (Lu et al.,
ip t
1982; Yuan et al., 1999; Zhao at al., 2001; Zhao, 2011). The Kaili Biota has been
predominantly documented from the middle–upper Kaili Formation at the Miaobanpo
cr
Section and the Wuliu-Zengjiaya Section near Balang Village of Jianhe County,
Guizhou Province. The Kaili Formation yielding the Kaili Biota is located primarily
us
across Jianhe County, Danzhai County, Zhenyuan County and Tongren City in eastern Guizhou Province (Fig. 1A). The specimens studied in this paper were collected at the
an
Miaobanpo Section and the Wuliu-Zengjiaya Section near Balang Village of Jianhe County, the Sanwan Section in Danzhai County and the Fujiachong Section in
M
Zhenyuan County (Fig. 2). The lower Miaobanpo Section is missing sections of the formation due to a fault contact with the Qingxudong Formation and the Fujiachong
d
Section of the Kaili Formation has no top or bottom parts of the formations exposed; however, the other two sections have basically complete sequences, which are in
te
conformable contact with the overlying and underlying strata. The studied sections
Ac ce p
from the Kaili Formation consist mainly of mudstones, with subordinate medium-bedded limestone, that vary in thickness at the top and base of the formation. (Zhao et al., 1993, 1994, 2012; Yuan et al., 1999, 2002; Yang et al., 2011).
Fig. 1 here
There are three trilobite zones present in the Kaili Formation. In ascending order,
these are the Bathynotus kueichouensis-Ovatoryctocara cf. granulata Zone, the Oryctocephalus indicus Zone, and the Peronopsis taijiangensis Assemblage Zone (Zhao, 2011; Zhao et al., 2012). Most of the Nisusia specimens examined in this study were collected of between 36.74 m and 111.69 m at the Miaobanpo Section in Jianhe County; however, a few specimens were collected from 53.3-164.13 m at the Wuliu-Zengjiaya Section, and 160-280 m at the Sanwan Section in Danzhai County, 4
Page 4 of 32
in addition to 11.9-106 m at the Fujiachong Section in Zhenyuan County. All specimens are from the Oryctocephalus indicus Zone (Fig. 2).
3. Geographic and stratigraphic distribution of Nisusia
ip t
Fig. 2 here
cr
Thirty-nine documented species (including the new species described herein) of Nisusia are currently considered valid (Table 1) (Verneuil and Barrande, 1860;
us
Billings, 1861; Tate, 1892; Walcott, 1886, 1889, 1905, 1908, 1912, 1924; Sun, 1924; Cooper, 1936; Endo and Resser, 1937; Lermontova, 1940; Bell, 1941; Shaw, 1954,
an
1957; Nikitin, 1956; Aksarina, 1960; Andreeva, 1962; Mount, 1981; Rowell and Caruso, 1985; Zeng, 1987; Popov and Tikhonov, 1990; Roberts and Jell, 1990; Brock,
M
1998; Wotte and Mergl, 2007; Benedetto and Foglia, 2012; Mao et al., 2014; Topper et al., 2014, 2015). The genus had a cosmopolitan distribution in the Cambrian and
d
has been documented from the USA, Canada, Mexico, Argentina, Uzbekistan, Kazakhstan, Kyrghyzstan, Bohemia, Siberia, Australia, Spain, and China (Table 1).
te
North America tops the list of the Nisusia species distribution, with 17 species found
Ac ce p
in USA and 6 species in Canada, followed by China with 6 species (including the new species documented herein). There are three Nisusia species documented from
Oceania.
Table 1 here
Of the 39 documented species of Nisusia (Table 1), 17 species have been
documented from the Cambrian Series 2 and 22 species from the Cambrian Series 3. Originating in the Cambrian Series 2 and becoming extinct in the Cambrian Series 3, Nisusia existed for a relatively short span of time compared to other brachiopod genera, such as Lingula (Williams et al., 1996). Nisusia sp. A, found in the Guanshan Biota, Wulongjing Formation in the Yunnan Province of China, which is equivalent to the middle Cambrian Series 2 and the Qiandongian Series Early Duyunian Stage, is 5
Page 5 of 32
considered herein to represent the earliest occurrence of Nisusia in China; the Zhangxia Formation in Luan County of Hebei Province (Sun, 1924) yields the youngest occurrence of Nisusia in China. Nisusia species in the Cambrian of China are distributed mainly in the North
ip t
China Region and the Yangtze Region. A number of Nisusia species were originally documented from the North China Region, comprising N. hayasakai from Luan
cr
County of Hebei Province (Sun, 1924), N. mantaouensis, N. salebrosa, N. orientalis and N. concentrica of Liaoyang County of Liaoning Province (Endo and Resser,
us
1937). Nisusia mantaouensis and N. salebrosa have since been synonymized with N. concentrica (Wang et al., 1966), leaving three valid species of Nisusia in the North
an
China Region. Four Nisusia species (including the unnamed Nisusia sp. A) have been found in the Yangtze Region, including Nisusia liantuoensis in Yichang City of Hubei
M
Province (Zeng, 1987), Nisusia sp. A in Kunming City of Yunnan Province (Luo et al., 2008; Hu et al., 2014), N. granosa (Mao et al., 2014) and N. guizhouensis n. sp. from
d
the middle–upper part of the Kaili Formation of Balang Village in Jianhe County of Guizhou Province. Nisusia granosa and N. guizhouensis n. sp. are the focus of this
te
paper. Some of the N. guizhouensis n. sp. specimens were also found in the
Ac ce p
Qingxudong Formation beneath the Kaili Formation (Zhao et al., 2012).
4. Morphological characteristics and morphological variation of Nisusia Except for N. granosa (Mao et al., 2014), N. ancauchensis (Benedetto and Foglia,
2012) and N. guizhouensis n. sp. (documented here) all of the 39 Nisusia species were reported and described during the 20th century. More than half of those descriptions were published prior to 1950. Owing to the oversimplified descriptions of some of the species and the limited number of fossil images illustrated, some specimens could not be assigned to the genus with great certainty and consequently the statistical work undertaken herein includes only 17 species in this paper (Table 2). During the Cambrian Series 2–3, Nisusia exhibited only minor morphological changes and can be regarded as a comparatively stable group. The stability and variability of Nisusia are
described as follows: 6
Page 6 of 32
Stability: Overall many morphological features of Nisusia remained relatively stable in the Cambrian. During the Cambrian Series 2–3, little morphological change is evident in Nisusia, and features such as the biconvex shells, anacline dorsal interarea, ornamentation of costellae, straight hinge line and the sulcus-bearing ventral
ip t
valve remained the same for the majority of species. However, subtle morphological
variations within the genus have allowed Nisusia to be differentiated into two species
cr
groups. Bell (1941) recognized morphological disparate groups of Nisusia on the
basis of North American species and his ideas have been expanded upon here. Group
us
A, consists of N. alaica (Cambrian Series 2), N. ancauchensis and N. guizhouensis n. sp. (Cambrian Series 2–3), and N. metula and N. grandis grandis (Cambrian Series 3).
an
All members of Group A are characterized by apsacline to catacline interareas and an apex occupied at the maximum height of the ventral valve. Group B, includes N.
M
festinata, N. kotujensis and N. fulleri (Cambrian Series 2), N. borealis, N. lickensis, N. brugessensis and N. hayasakai (Cambrian Series 3) and are all characterized by an
te
Ac ce p
Fig. 3 here
d
apsacline interarea and an apex occupying the maximum height of the ventral valve.
Variability: In the Cambrian Series 2, three types of ventral valve interarea are
found: apsacline, catacline, and procline, whereas in the Cambrian Series 3, only two types of ventral valve interareas were present: apsacline and catacline, indicating a decreasing trend in the variation of interarea type. The shell convexity varied from unequally biconvex in the Cambrian Series 2 to unequally biconvex and equally biconvex in the Cambrian Series 3. The maximum height of the ventral valve was always at the apex in the Cambrian Series 2, but it appeared sometimes at the umbo between the apex and the middle of the valve in the Cambrian Series 3 (see type B1 in Fig. 3). In addition to this evolutionary trend, the variability of Nisusia lies in certain
evolutionary relationships between various species groups. For example, a variation trend type A1 (Fig. 3), which exhibits a procline interarea with the maximum height of the ventral valve at the apex, only existed in the Cambrian Series 2, consisting of 7
Page 7 of 32
species such as in N. liantuoensis. Type B1, another variation trend, displays apsacline interareas, with the maximum height of the ventral valve at the near posterior margin instead of the apex, and only existed in the Cambrian Series 3, including species such
ip t
as N. deissi, N. fulleri and N. montanaensis.
cr
Table 2 here
us
5. Systematic palaeontology
Subphylum Rhynchonelliformea Williams, Carlson, Brunton, Holmer and Popov,
an
1996
Class Kutorginata Williams, Carlson, Brunton, Holmer and Popov, 1996
M
Order Kutorginida Kuhn, 1949
Superfamily Nisusioidea Walcott and Schuchert
te
Genus Nisusia Walcott, 1905
d
Family Nisusiidae Walcott and Schuchert
Ac ce p
Type species: Orthisina festinata Billings, 1861. Diagnosis: Shell subequally biconvex or unequally biconvex (ventribiconvex or
dorsibiconvex); ventral valve elevated and frequently more convex than dorsal; transverse subrectangular or semicircular in outline; anterior commissure rectimarginate or slightly sulcate; ventral interarea raised and triangular in shape, characterized as either apsacline, catacline or procline; basically anacline dorsal interarea, with the apex small and curled towards the posterior margin; dorsal interior with transverse, elevated notothyrial platform; dorsal adductor muscle scars may be discretely impressed on two pairs by prominent ridges; shell radially ornamented by costellae, with nodes or spines on some costae. Remarks: Andreeva (1987) described a new genus of Nisusiid, Narynella Andreeva,
1987 (type species Nisusia ferganensis Andreeva, 1962) from the Cambrian (Lenan) of Kyrgyzstan. The new genus was erected to distinguish brachiopod specimens that 8
Page 8 of 32
lack spinose costellae and exhibit a unisulcate commissure from the spinose, rectimarginate Nisusia (Andreeva, 1987; Popov and Williams, 2000). Not all Nisusia species definitively possess spines (Benedetto and Foglia, 2012) and few species of Narynella have been documented since for comparison (Geyer et al., 2014). Geyer et
ip t
al. (2014) considered that the nature of the commissure may be the only
distinguishing feature between the two genera, combined with the confusion about the
cr
age and origin of the type material (see Geyer et al., 2014) and the fact that selected
Narynella is in need of clarification and revision.
us
species of Nisusia are sulcate (Popov and Williams, 2000), and implied that the genus
Bell (1941) considered that the majority of Nisusia species could be divided into
an
two groups: one group consisting of taxa that possess a hemiconical ventral valve with the highest point at the apex, such as N. festinata (Billings, 1861), whereas the other
M
group is characterized by a convex ventral valve with an incurved beak and maximum height near the middle of the valve, such as N. sulcata (Rowell and Caruso, 1985) and
d
N. deissi (Bell, 1941). The specimens under consideration here exhibit the highest point of their ventral valves at the apex and belong to the former group.
Ac ce p
te
Occurrence: Cambrian Series 2–3; North America, Asia, Oceania, and Europe.
Nisusia guizhouensis n. sp.
(Figs. 4, 5)
2011
Nisusia sulcata – Zhao, p. 107, figs. 102-106.
Etymology: guizhouensis, derived from the locality of the new species in Guizhou
Province. Materials: 300 specimens in total, including specimens consisting of external and internal molds of both the ventral and dorsal valves. All specimens are reposited in the Institute of Palaeontology and Biological Mineralization, Guizhou University (GUK). Types: Holotype, GUK(M)-201 (Fig. 4a), ventral valve, valve length 11.06 mm, valve width 14.89 mm. Paratypes, GUK(M)-202 (Fig. 4b), ventral valve, valve length 6.50 9
Page 9 of 32
mm, valve width 9.37 mm; GUK(M)-203 (Fig. 4c), dorsal valve, valve length 7.21 mm, valve width 11.64 mm; GUK(M)-204 (Fig. 4d), dorsal valve, valve length 8.90 mm, valve width 14.61 mm. Diagnosis: Shell large, relatively for the genus, transverse rectangular in outline,
ip t
unequally biconvex in lateral view; surface radially ornamented by costellae,
increasing in number by bifurcations in the middle and front of shell surface;
cr
prominent concentric lines visible between the middle part and the anterior margin of each valve; ventral interarea apsacline to catacline, maximum height at the apex of
us
ventral which is fine and convex, with a sulcus; dorsal interarea anacline.
Description: Shell large, relative for the genus, valve length shorter than valve width;
an
holotype shell length 11.06 mm, shell width 14.89 mm; outline transversely subrectangular; hinge line straight, equal to the greatest shell width, cardinal
M
extremities near right-angle; unequally biconvex in lateral profile, convexity of the dorsal valve slightly larger than that of the ventral valve; maximum height of the
d
ventral valve at apex, shallow and wide sulcus originating from umbo to the anterior part of valve; hinge area high and triangular; anterior commissure mostly
te
rectimarginate or rarely slightly sulcate; thick costellae ornamented on shell surface,
Ac ce p
increasing in number by bifurcations in the middle and front of shell surface, in the adult phase about 7-9 costellae of 3 mm along the anterior margin; concentric lines prominent from the middle to the anterior margin of each valve in the adult phase. Ventral valve (Figs. 4a, b, f, j-o, 5j-r): moderately convex, maximum height at the
apex which is pointed, small and raised; outline transversely subrectangular; Interarea apsacline to catacline; in the central valve surface, a wide and shallow sulcus originating from umbo to anterior margin; muscle scars obscure; interior structure not clear. Dorsal valve (Figs. 4c-e, g-i, 5a-i): slightly more convex than ventral valve, apex incurved, umbo swollen, without fold, maximum height at about one-third of the valve length in front of umbo, interarea anacline; muscle scars obscure; interior structure not clear. Young stage (Fig. 5a-c, j-l): valve length less than 4 mm, width less than 6 mm; 10
Page 10 of 32
costae distinct, without bifucated costellae or concentric lines; apex pointed and projecting perpendicular to shell surface; sulcus obscure. Adolescent stage (Fig. 5d-f, m-o): valve length 4-6 mm, width 6-9 mm; clear costae on valve, with bifurcated costellae and few concentric lines appearing in the
ip t
anterior margin of each valve; apex pointed, small and straight; sulcus distinct.
Adult stage (Fig. 5g-i, p-r): more than 6 mm in length of valve, more than 9 mm
cr
in width; clear costae on valve, with bifurcated costellae and strong concentric lines appearing in the anterior margin of each valve; apex pointed, small and straight; a
us
wide and shallow sulcus originating from umbo to the anterior margin.
Remarks: Specimens of Nisusia guizhouensis n. sp. from the Cambrian of China
an
were originally described as Nisusia sulcata (Zhao, 2011). The absence of a sulcus on the dorsal valve and the maximum height of the ventral valve occurring at the apex
M
rather than the center of the valve clearly distinguish N. guizhouensis n. sp. from N. sulcata. Nisusia guizhouensis n. sp. also has a similar shell shape to Nisusia granosa
d
from the Kaili Formation at the Miaobanpo Section of Guizhou Province (Mao et al., 2014, p. 232, pl. 1). However, N. guizhouensis n. sp. possesses thick and convex
te
costae lacking nodes, whereas Nisusia granosa has fine and rounded costae bearing
Ac ce p
many small nodes. In addition, Nisusia guizhouensis n. sp. has stronger concentric lines than N. granosa.
N. grandis glabra from the Cambrian Series 2–3 of Australia (Roberts and Jell,
1990, p. 266, pl. 7, figs. A-Y) is similar to the species from Guizhou in both shell shape and the presence of well defined concentric growth lines. But N. grandis glabra tends to be generally alate in outline, has thicker costellae, and lacks the pointed apex (Fig. 4n-o) when compared with the species from Guizhou. As for the shell shape and costellae ornamentation, Nisusia guizhouensis n. sp.
closely resembles N. festinata from the Cambrian Series 2 of North America (Walcott, 1912; Cooper, 1936, p. 212, pl. 26, figs. 33, 39-42, 47) and N. borealis from the Middle Cambrian in Alaska (Cooper, 1936, p. 212, pl. 26, figs. 22, 23, 25-26, 28, 30-32, 34-35, 37-38, 43-44, 46, 48-50). Although these three species all have a sulcus on the ventral valve, Nisusia guizhouensis n. sp. possesses a sulcus on the ventral 11
Page 11 of 32
valve that extends from the umbo to the anterior margin, whereas the other two species have a sulcus extending from umbo to the middle part of the valve. Additionally, both Nisusia festinata and Nisusia borealis have nodes on their costellae, which are absent in the new species.
ip t
Occurrence: The middle–upper part of Kaili Formation of the Cambrian Series 2–3 of Balang Village in Jianhe County, Sanwan Village in Danzhai County, Zhuping
cr
Village in Zhenyuan County, Guizhou Province; the middle–upper part of the
Nisusia granosa Mao, Wang and Zhao, 2014
M
(Fig. 6)
an
Balang Village, Jianhe County, Guizhou Province.
us
Qingxudong Formation of the Cambrian Series 2 Stage 4 at the Songshan Section near
Nisusia alaica Popov and Tikhonov – Zhao, p. 107, figs. 107, 108.
2014
Nisusia granosa – Mao et al., p. 223, pl. 1, figs. 1-15.
d
2011
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Materials: 15 specimens in total (including 8 complete specimens).
Ac ce p
GTBM-9-4-21597 is the holotype (Fig. 6a; also Mao et al., 2014, pl. 1, fig. 1), ventral valve, valve length 10.12 mm, valve width 14.30 mm. Diagnosis: Shell relatively large for the genus, outline transversely rectangular, lateral
view unequally biconvex; shell radially ornamented with fine, rounded and low costae, increasing in number by intercalations or bifurcations in the middle and front of shell surface; many nodes exist on the costellae; ventral valve with a fine and pointed apex at its maximum height, bearing a sulcus. Description: Shell relatively large for the genus, valve length shorter than valve width; holotype shell length 10.12 mm, shell width 14.30 mm; outline transversely rectangular, hinge line straight, equal to the greatest shell width, cardinal extremities near right-angle or acute angle; lateral view unequally biconvex, convexity of the dorsal valve slightly larger than that of the ventral valve; ventral valve with maximum height at the apex, possessing a shallow and wide sulcus originating from umbo to 12
Page 12 of 32
anterior margin; hinge area high and triangular; anterior commissure mostly rectimarginate or slightly sulcate; fine ornamented costae on shell surface, increasing in number by intercalations or bifurcations in the middle and front of shell surface in the adult phase about 6-7 costellae of 3 mm along the anterior margin; many tiny
ip t
nodes appearing on costellae, concentric growth lines visible in anterior margin.
cr
Table 3 here
us
Ventral valve (Fig. 6a, d-f, h-i): moderately convex, maximum height at the apex which is pointed, small and raised; outline transversely rectangular; interarea
an
apsacline to catacline; in the central valve surface, a wide and shallow sulcus originating from umbo to anterior margin; valve surface radially ornamented by fine
M
and close growth lines, increasing in number by intercalations or bifurcations towards anterior margin, interarea apsacline to catacline; interior structure not clear.
d
Dorsal valve (Fig. 6b, c, g): slightly more convex than ventral valve, umbo swelling, without fold, maximum height at about one-third of the valve length in front
Ac ce p
not clear.
te
of umbo, apex incurved, interarea anacline; muscle scars obscure; interior structure
Comparison: Nisusia granosa Mao, Wang and Zhao, 2014 closely resembles Nisusia
alaica from the Cambrian Series 2, found in south Kirgizia (Popov and Tikhonov, 1990, p. 22, pl. 3, figs. 23, 24) in shell shape, costae furcations and ornamentation of nodes and spines. Nisusia granosa Mao, Wang and Zhao, 2014 however tends to be
larger (holotype shell width 14.30 mm and length 10.12 mm) than Nisusia alaica (holotype shell width 9.0 mm and length 6.2 mm), and has finer costellae with a higher density on shell surface. The ornamentation of Nisusia granosa Mao, Wang
and Zhao, 2014 is dominated by faint concentric lines, morphological features absent in N. alaica. Nisusia granosa Mao, Wang and Zhao, 2014 appears to bear a close resemblance to N. lickensis from the Cambrian Series 3 of Montana (Bell, 1941, pl. 32, figs. 9-12) in shell shape and the presence of costellae spines. Nisusia granosa differs from N. 13
Page 13 of 32
lickensis in being larger and in possessing a median sulcus on the ventral valve from the umbo to anterior margin. Nisusia granosa Mao, Wang and Zhao, 2014 is similar to N. festinata from the Cambrian Series 2, of North America (Walcott, 1912, pl. 100, fig. le) in features such
ip t
as shell shape, costellae and nodes on the costellae. Though they both have a sulcus
on the ventral valve, the sulcus on the ventral valve of N. festinata extends only from
cr
the umbo to the central part of the valve. The biggest difference between these two species is the absence of a sulcus on the dorsal valve of Nisusia granosa.
us
Nisusia granosa is also similar to N. kotujensis from the Cambrian Series 2–3 found from Siberia (Andreeva, 1962, p. 88) in shell shape, but it is significantly larger
an
and the lack of a detailed description makes comparison difficult.
Occurrence: Middle–upper part of the Kaili Formation of the Cambrian Series 2–3
M
(Wuling Series) Stage 5 (Taijiangian Stage), at the Miaobanpo Section in Balang
Acknowledgments
d
Village of Jianhe County, Guizhou Province.
te
Special thanks are given to Prof. Lars Holmer from Uppsala University (Sweden) and
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Prof. Zhi-Fei Zhang from Northwest University (China) for their valuable instructions and suggestions for this paper, Academician Xu Chen and Prof. Mao-Yan Zhu from Nanjing Institute of Geology and Palaeontology, China Academy of Sciences for their help and financial support of the fossil specimen photographs in this paper, Prof. Jin Peng from Guizhou University for her instructions and guidance, and Dr. Hai-Zhou Wang from Uppsala University (Sweden) for helping take photographs of some fossils in this paper. This paper benefited greatly from constructive reviews by Prof. Zhi-Fei Zhang, Christian Skovsted, and an anonymous reviewer. This study is supported by grants from the Major State Basic Research Development Program of China (973 program (N): 2013CB835002), the National Natural Science Foundation of China (41372019,40762018, 41362002, 41172005, 41330101) to Wang, Zhao, Peng, Yang; The foundation of Chinese Committee of Stratigraphy, Foundation of Guizhou Science Cooperation foreign (Gscf.G2010-70001) to Zhao; Guizhou Engineering and 14
Page 14 of 32
Technology Research Center for Development and Utilization of Karst Cave (Tourism) Resources (G [2014] 4004).
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ip t
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brachiopods: A perspective from South China. Gondwana Research 14, 241-254. Zhang, Z.F., Holmer, L.E., Popov, L.E., Shu, D.G., 2011a. An obolellate brachiopod with soft-part preservation from the early Cambrian Chengjiang Fauna of China. Journal of Paleontology 85 (3), 460-463.
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Zhang, Z.F., Holmer, L.E., Ou, Q., Han, J., Shu, D.G., 2011b. The exceptionally
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Zhao, Y.L., Yuan, J.L., Zhang, Z.H., Mao, J.R., Huang, Y.Z., Gong, X.Y., Wang, K., 1993. A preliminary study of the Kaili Formation in the transitional area in south
Chinese, with English abstract).
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China and its synchronous strata. Journal of Stratigraphy 17 (3), 170-178 (in
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Zhao, Y.L., Yuan, J.L., Huang, Y.Z., Mao, J.R., Qian, Y., Zhang, Z.H., Gong, X.Y., 1994. Middle Cambrian Kaili Fauna in Taijiang, Guizhou. Acta Palaeontologica
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Zhao, Y.L., Yu, Y.Y., Yuan, J.L., Yang, X.L., Guo, Q.J., 2001. Cambrian stratigraphy at Huanglian, Guizhou Province, China: Reference section for bases of the Nangaoan and Duyunian Stages. Palaeoworld 13, 172-180.
Zhao, Y.L., Peng, J., Yuan, J.L, Babcock, L.E., Guo, Q.J., Yin, L.M., Yang, X.L., Tai, T.S., Wang, C.J., Lin, J.P., Gaines, R.R., Sun, H.J., Yang, Y.L., 2012. Discussion of candidate stratotypes for the GSSP defining the conterminous base of Cambrian provisional Series 3and Stage 5. Journal of Guizhou University 29 (Supp. 1), 35-48.
Figure captions
20
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Fig. 1. (A) Location map of the study area in Guizhou. (B) Cambrian outcrops in the transitional belt of east Guizhou, South China (the shaded part of the map is adapted from Yuan et al., 2002).
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Fig. 2. Stratigraphic column of the Kaili Formation showing the distribution of
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Nisusia species in east Guizhou Province.
us
Fig. 3. Morphological variation of Nisusia.
Fig. 4. Nisusia guizhouensis n. sp. Except where indicated, scale bars represent 5 mm.
an
(a-f, h-o) Specimens collected from the Kaili Formation in Balang Village of Jianhe County, in Guizhou Province, South China; (a) GUK(M)-201, holotype, external view
M
of ventral valve; (b) GUK(M)-202 external view of ventral valve; (c) GUK(M)-203, external view of dorsal valve; (d) GUK(M)-204, external view of dorsal valve; (e)
d
GUK(M)-204, magnifying posterior margin of dorsal valve; (f) GUK(M)-205, external view of ventral valve, exhibiting sulcus from umbo to anterior part of valve;
te
(h) GUK(M)-207, external view of dorsal valve; (i) GUK(B)-208, external view of
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dorsal valve; (j) GUK(M)-209, external view of ventral valve; (k) GUK(M)-209, external view of ventral valve (scanning electron microscope); (l) GUK(M)-209, external view of ventral valve (scanning electron microscope) with magnified apex of ventral valve; (m) GUK(M)-210, external view of ventral valve; (n) GUK(M)-210, anterior view of ventral valve exterior, displaying the pointed convex beak of the shell; (o) GUK(M)-210, lateral view of ventral valve exterior, displaying the pointed convex beak of the shell. (g) Specimen collected from the Kaili Formation in Zhuping Village of Zhenyuan County, in Guizhou Province, South China, GUK(F)-206, external view of dorsal valve.
Fig. 5. Nisusia guizhouensis n. sp. Scale bars represent 2 mm for (a-c, j-l) (young stage); 3 mm for (d-f, m-o) (adolescent stage); 5 mm for (j-i, p-r) (adult stage). (a, k, n) Specimens collected from the Kaili Formation in Zhuping Village of Zhenyuan 21
Page 21 of 32
County, in Guizhou Province, South China; (a) GUK(F)-211, external view of dorsal valve; (k) GUK(F)-221, external view of ventral valve; (n) GUK(F)-224, external view of ventral valve. (b-g, i, j, l, m, o-r) Specimens collected from the Kaili Formation in Balang Village of Jianhe County, in Guizhou Province, South China; (b)
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GUK(M)-212, external view of dorsal valve; (c) GUK(M)-213, external view of
dorsal valve; (d) GUK(M)-214, external view of dorsal valve; (e) GUK(M)-215,
cr
external view of dorsal valve; (f) GUK(M)-216, external view of dorsal valve; (g)
GUK(M)-217 external view of dorsal valve; (i) GUK(M)-219, external view of dorsal
us
valve; (j) GUK(M)-220, external view of ventral valve; (l) GUK(M)-222, external view of ventral valve; (m) GUK(M)-223, external view of ventral valve; (o)
an
GUK(M)-225, external view of ventral valve; (p) GUK(M)-226, external view of ventral valve; (q) GUK(M)-227, external view of ventral valve; (r) GUK(M)-228,
M
external view of ventral valve. (h) Specimen collected from the Kaili Formation in Sanwan Village of Danzai County, in Guizhou Province, South China, GUK(S)-218,
d
external view of dorsal valve.
te
Fig. 6. Nisusia granosa. Except where indicated, scale bars represent 3 mm. All
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specimens were collected from the Kaili Formation in Balang Village of Jianhe County, in Guizhou Province, South China. (a) GTBM-9-4-21597, holotype, external view of ventral valve. (b) GTBM-9-5-6200, external view of dorsal valve. (c) GTBM-9-3-5362, external view of dorsal valve. (d-f) GTBM-9-3-3757, ventral valve; (d) external view; (e) shell showing posterior margin (scanning electron microscope); (f) magnified spines of surface valve (scanning electron microscope). (g) GTBM-9-5-2668, external view of dorsal valve. (h, i) GTBM-9-2-3521, ventral valve; (h) external view, showing pointed beak of shell; (i) lateral view, showing sulcus from umbo to anterior part of valve.
22
Page 22 of 32
Table 1. Geological era and occurrence of Nisusia species. Cambrian Series
Species name
Second
Location
Third
N. deissi (Bell, 1941)
×
USA
N. lickensis (Bell, 1941)
×
USA
N. montanaensis (Bell, 1941)
×
USA
N. borealis (Cooper, 1936)
×
USA
N. sulcata (Rowell and Caruso ,1985)
×
USA
N. fulleri (Mount, 1981)
USA
×
USA
N. erecta (Walcott, 1905)
× × ×
×
N. transversa (Walcott, 1886)
×
N. bivia (Walcott, 1912)
×
an
N. argenta (Walcott, 1905)
N. spencei (Walcott, 1905) N. nautes (Walcott, 1905) N. kanakensis (Walcott, 1908) N. orientalis (Endo and Resser, 1937) N. hayasakai (Sun, 1924)
te
N. liantuoensis (Zeng, 1987)
d
N. concentrica (Endo and Resser, 1937)
M
N. rara (Walcott, 1908)
Ac ce p
N. amii (Walcott, 1905)
USA
×
USA
×
USA
×
USA
×
USA
×
China
×
China
×
China China
× ×
N. alberta (Walcott, 1889) N. oriens (Walcott, 1924)
USA
×
N. guizhouensis n. sp.
USA
USA USA
×
N. granosa (Mao, Wang and Zhao, 2014)
USA
us
N. utahensis (Walcott, 1905) N. cloudi (Shaw, 1954)
ip t
×
cr
N. festinate (Billings, 1861)
China China Canada
× ×
Canada Canada
N. spinigera (Walcott, 1924)
×
Canada
N. burgessensis(Walcott, 1924)
×
Canada
N. lowi (Walcott, 1908)
×
Canada
N. grandis grandis (Roberts and Jell, 1990)
×
Australia
N. metula (Brock, 1998)
×
Australia
×
Australia
N. compta (Tate, 1892)
N. kotujensis (Andreeva, 1962)
×
N. nasuta (Nikitin, 1956)
×
N. alaica (Popov and Tikhonov, 1990)
×
Kyrgyzstan
N. pospelovi (Aksarina, 1960)
×
Siberia
N. minussensis (Lermontova, 1940)
×
N. vaticina (Verneuil and Barrande, 1860) N. ancauchensis (Benedetto and Fogli, 2012)
×
Siberia ×
Kazakhstan
Siberia ×
Spain
×
Argentina
23
Page 23 of 32
Table 2. Morphological analysis and comparison of Nisusia species.
Hinge line
Anterior commissure Sulcus
Maximum height
Foramen
Costellae
TR0
0
0
1
0(V) 0(D)
1
0
0
N. montanaensis
3(M)
0
1
2
0
1
0(V) 0(D)
1
0
0
N. sulcata
3(M)
0
T1
0
0
1
0(D)
0(V)
1
0
N. grandis grandis
3(M)
1(D)
0
2
1
0(V)
0
0
N. lickensis
3(M)
1
T0
0
0
N. borealis
3(M)
1(V)
?
0, 1
0
N. metula
3(M)
1
0, 3
N. burgessensis
3(M)
1(V)
N. hayasakai
3(M)
1
2
N. granosa
2~3
1(D)
T1
N. guizhouensis n. sp.
2~3
1
N. liantuoensis N. alaica N. festinata N. fulleri N. ancauchensis
1
17(V)
0
0
1
18.1(V)
0
0
0
1
0
0(S)
0
0 to 2
1
0
0(T)
1
0
0(V) 0(D)
1
0
1
1
0
0(T)
0
2
0(V)
0
0
1
0(V)
0
0
0(V)
0
0
1
0
d
1
1
2
2
1
2(L)
1(V)
2
1
1
1
2(L)
1
0
1
1
2(L)
1
1
2
1
0(V) 0(D)
0
2(L)
1
T0
1
1
0(V)
0
2(L)
1
0
2
1
0(V)
0
0
0
cr
us 0
0
2(L)
ip t 0
0(V)
te
1
1
0
0
an
0
0 1, 2
0(V)
M
0
Ac ce p
N. kotujensis
1
Dorsal
Cardinal angles
1(D)
Maximum width (mm)
Ventral
Outline
3(M)
Interarea
Growth line
Shell shape
N. deissi
Fold
Age
Ornamentation
0
11.8(D)
1
0(S)
5 23(D)
0(P)
0
0(P)
0
9(V) 0 to 2
1
0
1
14.3(V)
0(P)
0
0
1
10.5(D)
0
0
3
1
8~14
0 to 2
1
9.0(V)
1
0(S) 0
0
0
0
0
0
0
0
0
0
2
15
Note: morphological information used in this table: V = ventral valve; D = dorsal valve. Age: 2 = Cambrian Series 2, 3 = Cambrian Series 3, M = middle, L = lower. Common shell characters: Shell shape: 0 = almost equally biconvex, 1 = unequally biconvex, 1(V) = unequally biconvex, the 24
Page 24 of 32
ventral valve is more convex than the dorsal valve, 1(D) = unequally biconvex, the dorsal valve is more convex than the ventral valve. Outline: 0 = subrectangular, 1 = subquadrate, 2 = semielliptical, 3 = subcircular, T =
transversely subrectangular, TR0 = transversely rounded subrectangular.
line: 0 = hinge line less than greatest width of shell, 1 = hinge line equal
cr
Cardinal angles: 0 = obtusely angle, 1 = right angle, 2 = acute angle. Hinge
ip t
transversely, R = rounded, e.g., T1 = transversely subquadrate, T0 =
greatest width of shell. Anterior commissure: 0 = rectimarginate, 1 = sulcate.
us
Sulcus: 0 = present, 1 = absent, 0(V) = sulcus of ventral valve, 0(D) =
sulcus of dorsal valve. Fold: 0 = present, 1 = absent, 0(V) = fold of ventral
an
valve. Maximum ventral valve height: 0 = in the apex, 1 = in near the
middle of valve. Foramen: 0 = present. Costae: 0 = present, 1 = absent, 0(S)
M
= 0(Spines), 0(T) = 0(Tubercle), 0(P) = 0(Pustula). Growth line: 0 = present, 1 = absent. Interarea: 0 = apsacline, 1= anacline, 2 = catacline, 3 = procline.
Ac ce p
te
show no description in this paper.
d
Width: W(V) = width of ventral valve, W(V) = width of dorsal valve. Space
25
Page 25 of 32
Table 3. Measurements of Nisusia granosa.
Dorsal valve
W (mm)
L (mm)
N
Cw (mm)
Sw (mm)
GTBM9-1-1128
15.56
10.73
7
0.25
0.25
GTBM 9-2-3521
15.41
10.65
6
0.26
0.26
GTBM 9-4-21597
14.3
10.12
6
0.28
0.29
GTBM 9-3-1132
12.66
10.02
6
0.25
0.27
GTBM 9-5-5963
12.34
9.67
7
0.26
0.27
GTBM 9-3-5362
14.93
11.01
6
0.26
0.26
GTBM 9-5-6200
13.27
8.17
7
0.28
0.28
GTBM 9-3-4127
8.96
6.63
7
0.26
0.26
13.4288
9.625
6.5
0.2625
Average value
ip t
Ventral valve
Specimen number
cr
Shell type
0.2675
Note: W – width of valve, L – length of valve, N – number of valve costae per 3 mm, Cw – width of Costellae, Iw
Ac ce p
te
d
M
an
us
– interspaces width of between lines.
26
Page 26 of 32
Page 27 of 32
ed
pt
ce
Ac M
an
us
cr
i
Page 28 of 32
d
te
Ac ce p us
an
M
cr
ip t
Page 29 of 32
ed
pt
ce
Ac M
an
us
cr
i
Page 30 of 32
d
te
Ac ce p us
an
M
cr
ip t
Page 31 of 32
d
te
Ac ce p us
an
M
cr
ip t
Page 32 of 32
ed
pt
ce
Ac M
an
us
cr
i