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Origination and diversification of Devonian ambocoelioid brachiopods in South China

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Abstract

The superfamily Ambocoelioidea is an important group of the Brachiopoda in the Devonian, both globally and in South China. In the Devonian, this group is also most diverse compared with that in other periods. Based on features of cardinal process and cruralium, three types of cardinalia are recognised, including Ambocoelia-type, Emanuella-type, and Rhyncospirifer-type. Our study shows that the Rhynchospirifer-type cardinalia is a distinct character that distinguishes them from the rest of the Ambocoeliidae; therefore, the Rhynchospiriferinae is re-elevated to the family rank. Guangxiispirifer of the previous Ambocoeliinae is reassigned to the Rhynchospiriferidae while Ambothyris, Choperella, Crurispina, Diazoma, Emanuella, Ilmenia, Ilmospirifer, Ladjia, Moravilla, and Zhonghuacoelia of the previous Rhynchospiriferinae are transferred to the Ambocoeliidae (= previous Ambocoeliinae). Three earliest ambocoelioids in South China are Ambothyris, Amboglossa, and Prolazutkinia in the upper Emsian; their emergences were likely associated with the global Upper Zlichov Event. The upper Emsian occurrences of Amboglossa and Prolazutkinia are their FADs (first appearance datum) globally. These two genera probably gave rise to the Rhynchospiriferidae and the Lazutkiniidae, respectively. After an initial gradual increase in generic richness, the Ambocoelioidea reached the highest diversity (10 genera including four endemic genera) in the late Eifelian and Early Givetian in South China. From then on, the diversity gradually decreased till the end of the Devonian (two genera) and was least affected by the Frasnian–Famennian Event. The Ambocoelioidea is a strongly facies-controlled group. Their temporal and spatial distributions show a close relationship with sea level changes. Ambocoeliids and rhynchospiriferids were adapted to different environments. The former inhabited deeper and partly dysoxic environments, therefore showed greater capabilities to migrate and survive extinctions. On the other hand, the rhynchospiriferids dwelled in a reef-related, high energy, and oxic environments, which were probably more sensitive to environmental changes; they also showed high diversity and endemism and more easily experienced rapid and regional extinctions. Three species representing the three types of cardinalia, Ambocoelia yidadeensis sp. nov., Ambothyris panxiensis, and Rhynchospirifer sp., are described.

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Acknowledgements

A number of colleagues and students are thanked for their help in the fieldwork and during the preparation of this paper, including Zhang Yubo (Peking University), Zong Pu (Chinese Academy of Geological Sciences), and Lü Dan (Research Institute of Petroleum Exploration and Development). Andrzej Baliński (Instytut Paleobiologii PAN) was thanked for providing some reference papers. This manuscript benefited from comments by Andrzej Baliński and an anonymous reviewer.

Funding

This work was supported by the National Natural Science Foundation of China (Grant 41290260).

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    Meiqiong Zhang & Xueping Ma

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This article is a contribution to the special issue “Devonian palaeoecosystems and palaeoenvironments of South China”

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Appendix. Taxonomic data list

Appendix. Taxonomic data list

All ambocoelioid brachiopods from the Devonian of South China to our knowledge are listed below.

Superfamily Ambocoelioidea George, 1931

Family Ambocoeliidae George, 1931

Ambocoelia:

Ac. yidadeensis sp. nov.: Panxi (location no. 5 in Fig. 1), Yidade Fm., Lower Frasnian (Fig. 9)

Ac. elongata Zhang, 1983: Huama (Near Tewo, location no. 7), Lure Fm., upper Eifelian (Zhang et al. 1983)

Ac. pseudosinensis Zhang, 1983: Tewo (location no. 7), Lure Fm., upper Eifelian; Shaanxi, Lengshuihe Fm., Frasnian (Zhang et al. 1983)

Ac. sp. 1: Nanbiancun (location no. 8), Nanbiancun Fm., uppermost Famennian (Yu 1988)

Ac. sp. 2: Liujing (location no. 2), basal Mintang Fm., upper Eifelian (Sun 1992)

Ambothyris:

At. longmenshanensis Yang, 1988: Longmenshan (location no. 6), Jinbaoshi Fm., upper Eifelian (Hou et al. 1988)

At. panxiensis (Chu, 1974) (= Ambocoelia panxiensis Chu in Fang and Zhu 1974; Ambocoelia yunnanensis Wang et al. 1974; Emanuella aff. transversa in Bai et al. 1982, pp. 12–13, 1994, p. 141): Xiangzhou (location no. 1), Yingtang Fm. (Figs. 11(a–c) and 12); Panxi (location no. 5), Qujing Fm., Eifelian–Middle Givetian

At. yunnanensis (Chu, 1974): Panxi (location no. 5), Qujing Fm., Middle Givetian (Wang et al. 1974) (Fig. 11(e))

At. parvissima (Yang, 1988) (= Emanuella parvissima in Hou et al. 1988): Longmenshan (location no. 6), Jinbaoshi–Guangwushan Fm., upper Eifelian–Upper Givetian. It is very similar to Ambothyris yunnanensis, therefore be assigned to Ambothyris.

At.? curvimarginalis (Zhang, 1983) (= Emanuella curvimarginalis in Zhang et al. 1983): Tewo (location no. 7), Dangduo Fm., probably upper Emsian–lower Eifelian. It has a moderately high interarea, 2–3 weak costae on each flank and a shallow furrow and therefore is included in Ambothyris questionably.

At. sp.: Dushan (Location no. 3), Hejiazhai Mbr, Lower Frasnian (Fig. 11(d)) (data of this study)

Aviformia grandia Xian, 1988: Longmenshan (location no. 6), Shiliangzi Mbr, lower Eifelian (Hou et al. 1988)

Crurithyris:

Cr.? minuta Zhang, 1983 (our question mark): Tewo (location no. 7), Cakuohe Fm., Upper Frasnian (Zhang, in XIGMR and NIGP 1987). It is a biconvex elongated–rounded shell distinct from the transversely rounded shell of true Crurithyris.

Cr. aff. lunievensis Nalivkin, 1979: Nanbiancun (location no.8), Nanbiancun Fm., uppermost Famennian (Yu 1988)

Cr. urei (Fleming, 1828): Xikuangshan (location no. 10), Menggong’ao Fm., uppermost Famennian (Lü 2017)

Cyrtinoides guangsiensis (Sun, 1992): Liujing (location no. 2), Mintang Fm., upper Eifelian (Baliński and Sun 2016)

Diazoma:

Di. sp.: Dushan (location no. 3), Jiwozhai Mbr, Middle Givetian (Fig. 5(g))

Di. wenchuanensis Chen, 1978: Wenchuan (location no. 13), Guanwushan Fm., Givetian (Xu et al. 1978)

Emanuella:

Em. takwanensis (Kayser, 1883): Xiangzhou (location no. 1), Jide Mbr; Shetianqiao (location no. 4), “Yijiawan” Fm.; Panxi (location no. 5), Qujing Fm.; Longmenshan (location no. 6), Guangwushan Fm., Givetian (Grabau 1931; Bai et al. 1982; Hou et al. 1988, own data)

Em. insinosa Zhang, 1983: Fengxian (Shaanxi Provice), Gudaoling Fm., Givetian (Zhang et al. 1983)

Em. guangxiensis (Yang, 1977) (= Ilmenispina guangxiensis Yang, 1977): Xiangzhou (location no. 1), Donggangling Fm. (= Jide Mbr), Lower Givetian. It should be included in Emanuella beacause of its discrete crural plates and strongly incurved interarea.

Em. orbicularia Yang, 1977: Xiangzhou (location no.1), Donggangling Fm. (= Jide Mbr), Upper Givetian (Yang et al. 1977)

Em. plicata Grabau, 1931: Xiangzhou (location no. 1), Jide Mbr, Lower Givetian; Panxi (location no. 5), Qujing Fm., Middle Givetian (Bai et al. 1982; Fig. 5(e, f))

Em. plicata pentagona Grabau, 1931:Xiangzhou (location no. 1), Jide Mbr; Panxi (location no. 5), Qujing Fm.; Longmenshan (location no. 6), Jinbaoshi Fm., Lower–Middle Givetian (Bai et al. 1982, Hou et al. 1988, own data)

Em. transversa Grabau, 1931: Longmenshan (location no. 6), Jinbaoshi–Guangwushan Fm., Lower Givetian–Middle Givetian (Hou et al. 1988)

Em. sp.: Xiangzhou (location no. 1), Jide Mbr, Lower Givetian (Fig. 5(d))

Ladjia:

La. torrida (Veevers, 1959a) (= Emanuella torrida in Ma 2009): Dushan (location no. 3), Hejiazhai Mbr, Lower Frasnian

La. jiwozhaiensis (Wang et al., 1974) (= Crurithyris jiwozhaiensis in Wang et al. 1974): Dushan (location no. 3), Jiwozhai and Hejiazhai Mbrs; Panxi (location no. 5), Yidade Fm., Middle Givetian–Lower Frasnian (Wang et al. 1974) (Fig. 5(a, b))

La. sp. (= Emanuella takwanensis in Ma 2009) Dushan (location no. 3), Hejiazhai Mbr, Lower Frasnian. It is internally identical with a type specimen of Emanuella takwanensis sectioned by Veevers (1959b). However, there are more numerous Givetian specimens of Emanuella takwanensis that are internally identical with another type specimen Emanuella takwanensis sectioned by Dürkoop (1970). To keep the name stability of the Givetian Emanuella takwanensis, Zhang (2016) and Hou et al. (2017) suggested to choose Dürkoop’s figure as the representative of the type species of Emanuella takwanensis. Em. takwanensis in Ma 2009 has the Triangular-knobbed type cardinal process; it should be assigned to Ladjia.

Zhonghuacoelia:

Zh. bispina Chen, 1984: Longmenshan (location no. 6), Tuqiaozi Fm., Lower–Middle Frasnian (Chen 1984; Hou et al. 1988)

Zh. oranta Chen, 1984: Longmenshan (location no. 6), Tuqiaozi Fm., Lower–Middle Frasnian (Chen, 1984; Hou et al. 1988)

Zh. sinensis (Tien, 1938): Longmenshan (location no. 6), Tuqiaozi Fm.; Tewo (location no. 7), Cakuohe Fm.; Ningxiang (location no. 11), Qiziqiao Fm., Middle Givetian–Upper Frasnian (probably lower Famennian), (Chen 1984; Hou et al. 1988; Zhang in XIGMR and NIGP 1987; Tien 1938).

Family Rhynchospiriferidae Paulus, 1957

Rhynchospirifer:

Rh. abnormalis Yang, 1977: Xiangzhou (location no. 1), Donggangling Fm. (= Jide Mbr), probably Lower Givetian (Yang et al. 1977)

Rh. hengxianensis Xian, 1983: Liujing (location no. 2), Mintang Fm., Lower Givetian (Xian 1998)

Rh. liujingensis Fang, 1974: Liujing (location no. 2), Mintang Fm., Lower Givetian (Xian 1998)

Rh. otalata Zhang: Tewo (location no. 7), Xiawuna Fm., Lower Givetian (Cao et al. in XIGMR and NIGP 1987, without description and illustration)

Rh. pteratus Zhang, 1983: Tewo (location no. 7), Xiawuna Fm., Lower Givetian (Zhang in XIGMR and NIGP 1987)

Rh. rectangularis Wang et al., 1974: Dushan (location no. 3), Jipao Mbr, Lower Givetian (Wang et al. 1974)

Rh. regularis Yang, 1977: Liujing (location no. 2), Mintang Fm., Lower Givetian (Xian 1998)

Rh. trigonum Xian, 1983: Liujing (location no. 2), Mintang Fm., upper Eifelian–Lower Givetian (Xian 1998)

Rh. sp.: Xiangzhou (location no. 1), Guche Mbr, Jide Mbr, lower Eifelian–Lower Givetian (Bai et al. 1982)

Rh. sp.: Panxi (location no. 5), Nanpanjiang Fm., upper Eifelian (Fig. 11(f))

Amboglossa:

Ag. dafengmenensis (Wang et al., 1974): Xiangzhou (location no. 1), Dale Fm., upper Emsian (Chen and Yao 1999)

Ag. eleganta Yang, 1988: Longmenshan (location no. 6), lower Jinbaoshi Fm., upper Eifelian (Hou et al. 1988)

Ag. mosolensis (Ljaschenko, 1958): Xiangzhou (location no. 1), Guche Mbr; Dushan (location no. 3), Longdongshui Mbr; Tewo (location no. 7), Dangduo–Xiawuna Fms.; upper Emsian–Lower Givetian (Wang and Zhu 1979; Cao et al. in XIGMR and NIGP 1987; Jin 1988)

Ag. transversa (Wang et al., 1974): Xiangzhou (location no. 1), Dale Fm.; Dushan (location no. 3), Longdongshui Mbr; Longmenshan (location no. 6), Shiliangzi Mbr; upper Emsian–lower Eifelian (Wang and Zhu 1979; Hou et al. 1988)

Ag. waganovae (Breivel, 1952): Xiangzhou (location 1), Dale Fm., upper Emsian (Wang and Zhu 1979)

Ag. sp. (= Ilmenia cf. subhians in Xu, 1979): Nandan (location no. 9), Tangxiang Fm., upper Emsian–lower Eifelian (Xu 1979). It is sulcate, with dental plates and a cruralium supported by a median septum; thus, it is included in Amboglossa.

Changtangella:

Ch. bisepta Xian, 1983: Liujing (location no. 2), Mintang Fm., Lower Givetian (Xian 1998)

Ch. obtorta Xian, 1983: Liujing (location no. 2), Mintang Fm., Lower Givetian (Xian 1998)

Ch. sp.: Liujing (location no. 2), Mintang Fm., upper Eifelian (Sun 1992)

Guangxiispirifer:

Gu. subaequatus Xian, 1983: Liujing (location no. 2), Mintang Fm., Lower Givetian (Xian 1998)

Gu. bisinuatus Xian, 1978: Dushan (location no. 3), Longdongshui Mbr, lower Eifelian (Xian and Jiang 1978)

Ilmeniopsis:

Io. changtangensis Xian, 1983: Liujing (location no. 2), Mintang Fm., Lower Givetian (Xian 1998)

Io. ovata Xian, 1983: Liujing (location no. 2), Mintang Fm., Lower Givetian (Xian 1998)

Ilmenispina sp.: Liujing (location no. 2), Mintang Fm., upper Eifelian (Sun 1992)

Kosirium magnum (Yang, 1977): Xiangzhou (location no. 1), Jide Mbr, Lower Givetian (Bai et al. 1994)

Levibiseptum dushanense (Xian, 1975): Dushan (location no. 3), Bangzhai Fm. (in Tunshang Mbr), upper Eifelian (Xian and Jiang 1978)

“Moravilla” sp. (our quotation marks): Liujing (location no. 2), Mintang Fm., upper Eifelian (Xian 1998). It is problematic because there were only two dorsal valves that are characterised by the Rhynchospirifer-type cardinalia and hence are assignable to the Rhynchospiriferidae, but true Moravilla is reassigned to the Ambocoeliidae in this study.

Family Lazutkiniidae Johnson and Hou, 1994

Lazutkinia huaningensis (Wang et al., 1974): Panxi (location no. 3), Qujing Fm., Middle Givetian (Wang et al. 1974)

Prolazutkinia lata (Hou and Xian, 1983): Xiangzhou (location no. 1), Dale Fm., upper Emsian (Chen and Yao 1999)

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Zhang, M., Ma, X. Origination and diversification of Devonian ambocoelioid brachiopods in South China. Palaeobio Palaeoenv 99, 63–90 (2019). https://doi.org/10.1007/s12549-018-0333-4

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