Palaeobiodiversity and Palaeoenvironments

, Volume 99, Issue 1, pp 63–90 | Cite as

Origination and diversification of Devonian ambocoelioid brachiopods in South China

  • Meiqiong Zhang
  • Xueping MaEmail author
Original Paper


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.


Devonian Brachiopods Ambocoelioidea Rhynchospiriferinae Classification Diversity South China 


Brachiopods were very abundant in the Devonian of South China, which play an important role in stratigraphic divisions, correlations, and palaeoenvironmental reconstruction. Because of this, brachiopod studies may be traced back to one and a half centuries ago, from morphological description (e.g. Davidson 1853) to modern comprehensive studies of morphology, stratigraphy, palaeogeography, ancient environments, evolution, and diversity change (e.g. Hou and Xian 1964; Jin 1988; Chen 1984; Chen and Yao 1999; Ma 1995, 2009; Ma and Day 2003, 2007; Ma et al. 2006) over.

The Ambocoelioidea is a major group in the Upper Palaeozoic, of which the family Ambocoeliidae accounts for 80% of the total 45 genera (data based on Johnson et al. 2006 and Gourvennec and Carter 2007). The Devonian saw the radiation of the superfamily, with the four families (Ambocoeliidae George, 1931, Lazutkiniidae Johnson and Hou, 1994, Eudoxinidae Nalivkin, 1979, and Verneuiliidae Schuchert, 1929) all present at various times. The Devonian is also the period during which this group is most diverse, with 34 out of 45 genera in the history of the Ambocoelioidea; the Rhynchospiriferinae of the Ambocoeliidae are even exclusively restricted to the Devonian. In South China, the Ambocoeliinae are also abundant. This paper focuses on the origin and diversification of the Ambocoeliidae in South China, with discussions on a new classification scheme of the family.

In the following text, formal Givetian and Frasnian substage divisions are consistent with the scheme approved by the Subcommission on Devonian Stratigraphy (SDS) (Becker 2007; Becker et al. 2012), with base of Po. rhenanus/Po. varcus Zone, base of (Lower) Schmidtognathus hermanni Zone, base of the Pa. punctata Zone, and approx. base of the Lower Pa. rhenana Zone as the base of the Middle Givetian, Upper Givetian, Middle Frasnian, and Upper Frasnian, respectively. The Emsian substages follow the suggestion of SDS, with the base of the Po. inversus Zone as the lower–upper Emsian substage boundary. For convenience, the Eifelian is temporarily divided into lower and upper Eifelian at the start of the Haikou Uplifting, approximately at the upper Tortodus kock. australis to lower Tortodus kock. kockelianus zones (Ma et al. 2017); the four informal Famennian substages are only broadly correlated with the conodont zonation, with base of the Lower Pa. rhomboidea Zone, base of the Lower Pa. ru. trachytera Zone, and base of the Middle Pa. gr. expansa Zone as the base of the middle, upper, and uppermost Famennian, respectively. Abbreviations used in this paper are: Pa. = Palmatolepis, Po. = Polygnathus, Fm. = Formation; Mbr = Member; for abbreviation of brachiopod generic names, see Appendix.

Geological setting

The South China plate was located in the low latitude areas in the eastern part of the Paleotethys during the Devonian (Fig. 1(a)). During the Early Devonian, marine transgression invaded the South China region from the southwest. Approximately at the start of the Middle Givetian, the palaeogeographic framework of the South China plate (Fig. 1(c)) became a network of intercepted carbonate platforms (shallow water) and grabens (deeper water) through at least two major rifting activities during the Emsian and Givetian (Ma et al. 2017). This platform-rift system continued to the end of the Frasnian in northern areas and of the Famennian in southern areas.
Fig. 1

Middle Devonian (Givetian) to Late Devonian (Frasnian) lithofacies map of South China ((c) is modified from Ma et al. 2009b, (a) from Scotese 2014b). Note location No. 7 is marked in (b) (to the north of the frame of (c))

Under the abovementioned palaeogeographic framework, four major basic facies types may be recognised in South China (Xian et al. in Hou and Wang 1988). The Xiangzhou type (subtidal benthic facies) is characterised by shallow, well-oxygenated, and agitated water environments with abundant various benthic organisms; the Nandan type (pelagic facies) occupied the platform basin areas (Fig. 1) and is characterised by predominant pelagic fossils and minor thin-shelled brachiopods; the Transitional type (mixed benthic and pelagic facies) represents a platform marginal facies between platforms and platform basins with alternating deeper and shallow water sediments; and the Qujing type (littoral facies) is characterised by intertidal clastic deposits with fish, vascular plants, and spores. The ambocoelioid brachiopods can be found in a variety of environments, but they mostly inhabited deeper water areas, including deeper subtidal and platform basin (rift-basin) environments; however, the Rhynchospiriferinae seem to prefer a platform setting.

Temporal and spatial distributions of the Ambocoelioidea

Global distribution

The Silurian saw the first ambocoelioid brachiopods in western North America (Nevada and northwestern Canada) and Canadian Arctic Islands: Eoplicoplasia and Dicoelospirifer in the upper Wenlock (Figs. 2 and 3; Zhang 1989a, b). During the Ludlow and Pridoli, this group began to expand its geographic distribution into the Baltica (e.g. Kraft and Kvaček 2017) and questionably into Western Australia (Johnson and Lenz 1992). The Ludlow occurrence of Eoplicoplasia? and Lochkovian occurrence of Ambocoelia in Australia are somewhat enigmatic on the other end of the distant either Paleotethys or Panthalassic Ocean (Fig. 3).
Fig. 2

Stratigraphic distribution of ambocoelioids (data mainly based on Johnson et al. 2006; see Appendix for additional sources). a Dorsal view of Dicoelospirifer dicoelospirifer (from Zhang 1989b). b Dorsal view of Eoplicoplasia cf. acutiplicata (from Zhang 1989b). c Dorsal view of Metaplasia hemicona (from Havlíček 1990). dMetaplasia paucicostata (from Johnson 1970). Note: genera of Baranov (2007, 2009) and Baranov and Alkhovik (2006) are excluded because their family assignment is doubtful (see Fig. 3: genus nos. 31 and 32 for generic names)

Fig. 3

Palaeobiogeography and generic distribution of amboceolioids during the Wenlock (middle Silurian) to Famennian (Late Devonian). aMetaplasia hemicona (from Havlíček 1990, Ludlow, Czech Republic). bAmbothyris? inopis (sic. question mark, from Mawson and Talent 1999, Lochkovian, southeastern Australia; our note: because of having a bilobed cardinal process, it possibly belongs to Ambocoelia or Metaplasia). cMetaplasia cf. lenzi (from Lenz and Johnson 1985, Lochkovian, southeastern Australia). dAmbocoelia bubovica (from Mergl 2003, Lochkovian, Bohemia). eAmbocoelia sp. (from Lenz and Johnson 1985, Lochkovian, southeastern Australia). fAmbocoelia aff. praecox (from Lenz 1977, Pragian, Yukon). gAmbothyris runnegari (Chatterton 1973, Emsian, southeastern Australia). h Spreading of generic distribution of amboceolioids with time; palaeogeographic maps after Scotese 2014a, b. Taxonomic data are from various sources (Bai et al. 1982, 1994; Baliński and Sun 2016; Baliński et al. 2016; Baranov 2007, 2009; Baranov and Alkhovik 2006; Beus 1965; Blodgett and Johnson 1994; Boucot 1959; Boucot and Lawson 1999; Boucot et al. 1970, 1986; Brice et al. 1973; Caldwell 1967; Chatterton 1973; Chatterton and Perry 1978; Chen 1984; Chen and Yao 1999; Cherkesova 1988; Chlupáč 1982; Chlupáč et al. 1998; Cooper and Williams 1935; Day 1988, 1996; De Keyser 1977; Dürkoop 1970; Fagerstrom 1961; Farrell 1992; Ficner and Havlíček 1978; Frost and Langenheim 1966; Goldman and Mitchell 1990; Havlíček 1953, 1990, 1998; Havlíček and Kukal 1990; Heidelberger and Amler 2002; Herrera 1995; Hou et al. 1988; Isaacson 1977; Johnson 1970, 1971, 1974, 1986; Johnson and Klapper 1990; Johnson and Lenz 1992; Johnson and Blodgett 1993; Johnson et al. 1969, 1996; Kelus 1939; Kesling and Chilman 1975; Klovan 1964; Koch 1981; Lenz 1970, 1972, 1973, 1977; Lenz and Johnson 1985; Lesperance and Sheehan 1975; Liashenko 1969; Mawson and Talent 1999; McGhee 1976; McGhee and Sutton 1985; Mergl 2003; Mergl and Massa 1992; Michels 1986; Morales 1965; Mottequin 2008; Norris and Uyeno 1998; Norris et al. 1992; Paulus 1957; Perry 1978, 1979, 1984; Racki 1992; Robinson 1963; Rode and Lieberman 2004; Rohr and Smith 1978; Rzhonsnitskaia 1955; Sanchez and Benedetto 1983; Sandford and Norris 1975; Sapelnikov et al. 1995; Savage 1974; Telford 1988; Tiazheva 1960; Veevers 1959a; Vogel et al. 1989; Vopni and Lerbekmo 1972; Wang and Zhu 1979; Wang et al. 1974, 1987; Xian 1983, 1998; Xian and Jiang 1978; XIGMR and NIGP 1987; Xu 1979; Xu et al. 1978; Yang et al. 1977; Yolkin et al. 1988, 2011; Yu 1988; Yudina and Rzhonsnitskaya 1985; Zambito and Schemm-Gregory 2013; Zhang 1989a, 1989b, 1991)

No ambocoelioids have been found in the Silurian in South China yet although there were favourable sedimentary rocks that yield various other brachiopod taxa. The upper Ludlow Guandi (= Kuanti) Fm. and Miaogao (= Miaokao) Fm. (especially the latter) yield various brachiopods including orthides, rhynchonellides, atrypides, athyridides, and spiriferides (without ambocoelioids) in eastern Yunnan (Qujing) (Zhang et al. 1996); the Pridoli Yanglugou Fm. (= upper Bailongjiang Fm.) of western Qinling bears relatively abundant and diverse brachiopods, including the spiriferide cyrtioids (Eospirifer and Hedeina of the Cyrtiidae), adolfioids (Nikiforovaena of the Adolfiidae), delthyridoids, and reticularioids as well as other brachiopods (Rong et al. in XIGMR and NIGP 1987); the Wenlock Zhoushan Group (including the Xiaolianggou and Miaogou formations; limestones and metamorphosed clastics) yields various brachiopods including rhynchonellides and athyridides, and the Pridoli(?) Bailongjiang Group (metamorphosed limestones and clastics) yields corals and brachiopods (atrypides, orthides, spiriferides, and others and conodonts, trilobites) (GBDMR 1989). Zhang (in XIGMR and NIGP 1987, vol. II, p. 133–134) mentioned that Amboglossa mosolensis ranges from the Pridoli Yanglugou Fm. to the Upper Givetian–Lower Frasnian Pulai Fm., but only the upper Emsian–lower Eifelian Dangduo Fm. and upper Eifelian Lure Fm. and basal Xiawuna Fm. can be confirmed that yield the species (with an Amboglossa sp. in the lower Givetian Xiawuna Fm.).

In the Devonian, the Ambocoelioidea is very abundant. According to Johnson et al. (2006) and Gourvennec and Carter (2007), there are 45 genera in total in the superfamily (Fig. 2), of which the Ambocoeliidae include 36 genera, accounting for 80%; the Lazutkiniidae, 2 genera; the Eudoxinidae, 4 genera, and the Verneuiliidae, 3 genera. Thirty-four genera (76% of the total ambocoelioid genera) lived in the Devonian, including 29 genera of the Ambocoeliidae (accounting for 85% of the total Devonian genera), 2 genera of the Lazutkiniidae, 1 genus of the Eudoxinidae, and 2 genera of the Verneuiliidae. Of the Devonian genera, 60% (27 genera) were present in the Middle Devonian, reaching the highest diversity in generic richness and widest geographic distributions (Figs. 2 and 3). The Rhynchospiriferinae and the Lazutkiniidae are restricted to the Devonian, with numerous endemic taxa in South China (Fig. 3: genus Nos. 27–29), making the South China region an important area for the study of this superfamily.

During the Early Devonian Lochkovian through Pragian, the Ambocoelioidea were most diverse in the Euramerica continent. They did not occur in South China until the late Early Devonian (Emsian) although brachiopods are relatively diverse from the Lochkovian through Pragian in the Yulin area of Guangxi, a facies characterised by deeper water clastics with alternating brachiopod and graptolite faunas (Wang and Yang 1998) including various spiriferides: Hedeina (Eospiriferinae, Cyrtioidea), Nikiforovaena of the Adolfiidae (Adolfioidea), Orientospirifer of the Delthyrididae (Delthyridoidea), Quadrithyrina and Quadrithyris of the Xenomartiniidae, and Reticulariopsis of the Reticulariidae (both Reticularioidea) as well as other brachiopods. In the Middle Devonian, the Ambocoelioidea were most diverse in the whole world, especially in the Euramerica continent and South China. During the Late Devonian, this group started to decrease in diversity, with a limited number of genera recorded in the Lower Frasnian (Veevers 1959a; Beus 1965; Mottequin 2008; Ma 2009; Baliński et al. 2016). The Frasnian–Famennian Mass Extinction Event does not seem to significantly affect the group (Fig. 2) as they were already very rare in the Frasnian. In the Carboniferous and Permian, the diversity kept on a low level until finally becoming extinct in the Early Triassic (Fig. 2).

Ambocoelioid distribution in South China

Stratigraphy and divisions

The Ambocoelioidea-bearing strata are mostly deposits of platform origin, with rare pelagic index fossils. Nevertheless, their underlying or overlying strata may bear relatively common to abundant pelagic fossils. The Liujing section (Fig. 4) is the best representative for both pelagic and benthic fossils, where the conodont zonation can be established, e.g. the Yujiang Fm. (Lu et al. 2017), the Moding Fm. (Kuang et al. 1989; Bai et al. 1994), the Najiao Fm. (Kuang et al. 1989), the Mintang Fm. (Kuang et al. 1989; Bai et al. 1994; Jiang et al. 2000), the Gubi Fm. (Kuang et al. 1989; Bai et al. 1994; Jiang et al. 2000), and the Rongxian Fm. (Kuang et al. 1989). The Longmenshan and Dale sections are dominated by benthic fossils but index conodonts are also common at some horizons and intervals (Hou et al. 1988; Bai et al. 1982, 1994).
Fig. 4

Correlation chart of Devonian stratigraphic units in South China. Dotted lines indicate the uncertainty in correlation with international stages. Vertical lines signify a hiatus

In the absence of pelagic index fossils, benthic coral and brachiopod faunas may also play an important role in the correlation (e.g. Ma et al. 2009a). The regional Yujiangian (lower Emsian) is characterised by the Dicoelostrophia–Rostrospirifer tonkinensis Assemblage in the Tonggeng, Yujiang, and Ganxi formations of various sections. During the Ertangian (mid-Emsian, approx. the conodont Eocostapolygnathus nothoperbonus Zone; Hou et al. 2017), the Moding deepening/rifting event happened, which probably resulted in the wide spread of the ammonoid Erbenoceras fauna in South China, e.g. in the Dale and Nandan sections (Ma et al. 2009a). The Sipaian (upper Emsian) is characterised by the Trigonospirifer trigonata–Otospirifer daleensis–Euryspirifer paradoxus Assemblage which is widely distributed.

The lower Eifelian is characterised by the brachiopod Athyrisina–Yingtangella–Xenospirifer Assemblage Zone and the coral Utaratuia Assemblage. Some major elements of the assemblage such as Eospiriferina houershanensis subplanus and Athyrisina squamosa can also be found in the equivalent strata in the West Qinling region. In South China, the late Eifelian saw a general sea level fall (Haikou Movement of Hou 1978) as represented by the Changcun, Dahekou, and Tunshang members, and lower Jinbaoshi Fm. (Fig. 4).

The Stringocephalus Abundance Zone bears various terebratulide, spiriferide, and atrypide brachiopods, which spans the Lower and Middle Givetian times (Ma and Zong 2010). The Jide, Dushan, Yijiawan, Qiziqiao, Poxi, Qujing, upper Jinbaoshi, lower and middle Guanwushan, and Xiawuna formations yield various brachiopods of this zone. Within this zone, the coral Endophyllum is also wide spread, representing the Endophyllum transgression in South China (Liao and Ma 2011) approx. at the end of the Early Givetian. In the Upper Givetian, Stringocephalus was very rare; instead, ambocoeliids and leiorhynchids became relatively abundant. The base of the Upper Givetian in South China is characterised by a sharp lithological change (deposits resulting from a sea level rise) in a number of sections such as the Panxi section, Longmenshan, and Tewo sections. There are a few sections where the Upper Givetian deposits may be missing probably due to erosion during an assumed subsequent regression (Ma et al. 2017) such as the Dushan, Qiziqiao, and Xikuangshan sections (Liao et al. 1979; Wang et al. 1986).

Brachiopods of the Lower Frasnian in South China are similar in composition to those of the Upper Givetian; both intervals belong to the ambocoeliid-leiorhynchid Assemblage Zone of Ma and Zong (2010). However, the ambocoeliids Diazoma and Emanuella, and rare Stringocephalus may be found in the Upper Givetian, whereas the leiorhynchids Leiorhynchus, Calvinaria, and Yocrarhynchus are only found in the Frasnian (Zhang et al. 2015). In the Middle Frasnian, the emergence of the Cyrtospiriferidae and the Conispiriferidae in South China was approx. in the Pa. hassi Zone (Ma et al. 2006). The lower part of the Upper Frasnian is characterised by the flourishing of reefs and reefal carbonates at the Qilijiang highstand of Ma et al. (2017) and subsequent drowning of reefs due to a sea level rise (Lower Kellwasser Event) (Ma et al. 2016). All the above faunal and sea level change events may serve to make stratigraphic correlations.

The lower and middle Famennian (= local Xikuangshanian stage) are characterised by the Yunnanella (previously Yunnanellina)–Sinospirifer and Nayunnella (previously Yunnanella)–Hunanospirifer Assemblage Zones, respectively, which are readily recognised and differentiated by their taxonomic differences (Ma and Zong 2010). Probably due to an interpreted sustained sea level fall/lowstand in the early Famennian, deposits of the Yunnanella–Sinospirifer Assemblage Zone may be missing in many then coastal areas in central Hunan (Ma and Zong 2010). The Yangshuoan is represented by deposits (coarse clastics and dolostone) under a regressive phase in the isolated platform and nearshore areas. Benthic fossils are very rare in this interval in various pelagic and neritic sections so that no coral and brachiopod assemblages can be established (Ma et al. 2009a). The Shaodongian represents a renewed marine transgression (e.g. Ma et al. 2017). Subsequently, the corals Cystophrentis and/or Beichuanophyllum became widely distributed in South China in the interval approx. equivalent to the Etroeungt of Europe, commonly associated with the last cyrtospiriferids.

Stratigraphic and geographic distribution of the ambocoelioids

Ambocoelioid brachiopods mostly occur in Xiangzhou, Liujing, Nandan, and Nanbiancun of Guangxi, Dushan of Guizhou, Panxi and Qujing of Yunnan, Xikuangshan, Shetianqiao, and Ningxiang of central Hunan, Longmenshan of Sichuan, and Tewo of Gansu (see Fig. 1 for locations and Fig. 4 for stratigraphic correlation) (see Appendix for a full taxonomic list of the Ambocoelioidea in South China and their revised names).

The Xiangzhou area of Guangxi was characterised by deposits of a typical shallow water platform facies during the Devonian with abundant benthic organisms (Bai et al. 1982, 1994; Jin 1988; Wang and Zhu 1979; Yang et al. 1977; Chen and Yao 1999). The ambocoelioids occur in the upper Emsian Dale Fm. (Amboglossa and Prolazutkinia) and in the Yingtang Fm. (Amboglossa in the lower Eifelian Guche Mbr, Ambothyris in the lower Eifelian Gupa Mbr to upper Eifelian lower–middle parts of the Changcun Mbr, and Rhynchospirifer in the lower Eifelian Guche Mbr and Lower Givetian Jide Mbr). The Lower Givetian Donggangling Fm. (= Jide Mbr of Bai et al. 1994) yields Emanuella (including Ilmenispina guangxiensis Yang, 1977 and Emanuella sp., Fig. 5(f)) and Kosirium.
Fig. 5

adLadjia jiwozhaiensis (Wang et al. 1974) from the Middle Givetian Jiwozhai Mbr, Dushan section. (a1–a3) Dorsal, ventral views, and enlarged microornament on primary layer (worn off during later ultrasonic cleaning), showing spine bases (black dots) of specimen PUM16215; (b1–b3) dorsal, ventral views, enlarged microornament after slight abrasion (capillae and concentric growth lines) of specimen PUM16214; (c1, c2) ventral view and enlarged ventral margin showing faint radial striae on exfoliated valve of specimen PUM16213; (d1, d2) interior of ventral valve and interior of dorsal valve with one whorl of spire left on each side of specimen PUM16125. eLa. jiwozhaiensis (Wang et al. 1974) from the middle Yidade Formation (Lower Frasnian), Panxi section. (e1–e4) Dorsal, ventral, anterior, and lateral views of specimen PUM16201. fEmanuella sp. from the Lower Givetian Jide Mbr, Baqi, Xiangzhou. (f1–f7) Dorsal, ventral, anterior, posterior, lateral views, and enlarged exfoliated shell of specimen PUM16211, showing spine bases (pits on primary layer, worn off during later ultrasonic cleaning). gDiazoma sp. from the Middle Givetian Jiwozhai Mbr of the Dushan section. (g1–g7) Dorsal, ventral, anterior, and posterior views of specimen PUM16203. h, iEmanuella plicata Grabau, 1931 from the Middle Givetian Qujing Formation, Panxi section. (h1–h5) Dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16142; (i1–i5) dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16117. (j) Stringocephalus nevadensis Frost and Langenheim, 1966 from the lower Yidade Formation of the Panxi section, Upper Givetian. (j1–j3) Dorsal, ventral, and lateral views of specimen PUM16207

The Dushan section of Guizhou is also of a typical shallow water platform facies, with ambocoelioid brachiopods in various horizons (Liao et al. 1978; Xian and Jiang 1978; Wang and Zhu 1979; Ma et al. 2009a; this study). The lower Eifelian Longdongshui Fm. yields Amboglossa and Guangxiispirifer. The upper Eifelian Tunshang Mbr of the Bangzhai Fm. yields Levibiseptum (the Dahekou Mbr lacks fossils according to Liao et al. 1978). The Lower Givetian Jipao Mbr of the Dushan Fm. bears Rhynchospirifer. The basal Jiwozhai Mbr of the Dushan Fm. yields the coral Endophyllum, which indicates that the level still lies in the Lower Givetian (Hou et al. 1988; Liao and Ma 2011) and that the rest of the Jiwozhai Mbr is of Middle Givetian age with Ladjia and Diazoma. The Lower Frasnian basal Hejiazhai Mbr of the Wangchengpo Fm. yields Ambothyris and Ladjia (including Emanuella torrida and Em. takwanensis in Ma 2009, Crurithyris jiwozhaiensis in Wang et al. 1974).

In central Hunan, the Middle and Upper Givetian “Yijiawa Fm.” of the Shetianqiao section yields Emanuella (Ma et al. 2004); the Middle Givetian Qiziqiao Fm. in the Ningxiang area bears Zhonghuacoelia sinensis (= Ambocoelia sinensis in Tien 1938); the uppermost Famennian Menggong’ao Fm. in the Xikuangshan section yields Crurithyris (own data), a brachiopod that is distinct in the cardinalia (see the next section for discussion).

The Liujing area of Guangxi, an important reference section for correlation of different facies, is characterised by deposits of a platform–“basin” transitional facies, with both benthic and pelagic fossils. Brachiopods of the Mintang Fm. have been studied in detail by Xian (1983, 1998), Sun (1992), and Baliński and Sun (2016), including Ambocoelia, “Moravilla” (our quotation marks), Changtangella, Guangxiispirifer, Ilmenispina, Cyrtinoides, and Rhynchospirifer in the Eifelian part (basal Mintang Fm.) and Changtangella, Guangxiispirifer, Rhynchospirifer, and Ilmeniopsis in the Lower Givetian part (lower Mintang Fm.).

In eastern Yunnan, the Panxi section is also represented by platform and platform marginal deposits. The benthic fossils (corals and brachiopods) have been studied by many workers (e.g. Hou and Xian 1964; Liao et al. 2006). Ambocoelioid brachiopods have been found in the probably upper Eifelian Nanpanjiang Fm. (Rhynchospirifer), in the Middle Givetian Qujing Fm. (Ambothyris and Emanuella), and in the Lower Frasnian Middle Member of the Yidade Fm. (Ambocoelia and Ladjia). The Lower Member of the Yidade Fm. has been demonstrated to be of the Upper Givetian by the occurrences of the brachiopod Stringocephalus nevadensis (Fig. 5(j1–j3)) and the conodont Po. cristatus, which is an Upper Givetian conodont (own data).

With the comprehensive study of Hou et al. (1988), the complete Devonian sequence of the Longmenshan area has been established (see Ma et al. 2009a for the column of litho- and biostratigraphy). Ambocoelioid brachiopods have been found in the lower Eifelian Shiliangzi Mbr of the upper Yangmaba Fm. (Amboglossa and Aviformia), in the upper Eifelian lower Jinbaoshi Fm. (Amboglossa and Ambothyris), in the Lower and Middle Givetian upper Jinbaoshi Fm. and the Guanwushan Fm. (Emanuella and Ambothyris) in the Upper Givetian upper Haijiaoshi Mbr of the Guanwushan Fm. (Emanuella and Ambothyris), and in the Lower and Middle Frasnian Tuqiaozi Fm. (Zhonghuacoelia). No specific age was provided for Diazoma wenchuanensis from the Givetian Guanwushan Fm. of Wenchuan (Xu et al. 1978).

According to Zhang et al. (1983) and Cao et al. (in XIGMR and NIGP 1987), ambocoelioid brachiopods in the Western Qinling include Amboglossa in the upper Emsian through upper Eifelian Dangduo to basal Xiawuna formations, Ambocoelia in the upper Eifelian part of the Lure Fm. and the Frasnian Lengshuihe Fm., Rhynchospirifer in the Lower Givetian part of the Xiawuna Fm., “Emanuella” (our quotation marks) in the Lower Frasnian upper Pulai Fm. (our note: true Emanuella is Givetian in South China; there is no description and illustrations for Emanuella sp. from the Pulai Fm., and thus, it is excluded from our discussion), and Crurithyris? (our question mark, Cr. minuta Zhang, 1983) in the Upper Frasnian lower part of the Cakuohe Fm. Ambothyris? (our quotation marks, including Emanuella curvimarginalis in Zhang et al. 1983) from the Dangduo Fm. without an exact stratigraphic position (upper Emsian to lower Eifelian). The Cakuohe Fm. yields Zhonghuacoelia, but the exact horizon is uncertain (Upper Frasnian to probably lower Famennian).

In addition, the uppermost Famennian part of the Nanbiancun Fm. in the Nanbiancun section of Guangxi yields Ambocoelia and Crurithyris (Yu 1988). The upper Emsian–lower Eifelian Tangxiang Fm. in the Nandan section of Guangxi bears Amboglossa.

Revised classification of the Ambocoeliidae George, 1931

Major morphological characteristics

According to Johnson et al. (2006), the superfamily Ambocoelioidea George, 1931 consist of four families of commonly small shells that are characterised by lacking well-developed fold and sulcus, but possessing a commonly simple, knoblike cardinal process, broad and well-developed outer hinge plates, and a variably developed cruralium. The two subfamily divisions of the Ambocoeliidae are based on the nature of crural plates, microornament, and dental plates: the Ambocoeliinae is characterised by vestigial or lacking crural plates, commonly fine concentric growth lamellae, and lacking dental plates, whereas the Rhynchospiriferinae is characterised by well-developed crural plates, commonly fine capillae, and variably developed dental plates. In fact, the development of crural plates and dental plates as well as the cardinal process is quite variable in the whole superfamily Ambocoelioidea. For example, in the Rhynchospiriferinae, Crurispina possesses a small fixed (sessile) cardinal process, short and discrete crural plates and lacking dental plates, which is similar to Ambocoelia (see transverse section of Goldman and Mitchell 1990, figs. 10, 11, 13, 14) of the Ambocoeliinae, whereas Crurithyris of the Ambocoeliinae completely lacks crural plates, with the dorsal hinge structure identical to that of many other spiriferides such as the superfamilies Martinioidea Waagen, 1883 and Reticularioidea Waagen, 1883 (see Lü and Ma 2017, fig. 16, Sun and Baliński 2011, fig. 23, for comparison of the two types: ambocoelioid type and non-ambocoelioid type dorsal hinge structure).

Many workers have questioned the use of dental plates as an important feature to divide the Ambocoeliidae (Nalivkin 1941, p. 217; Veevers 1959a, p. 124; Johnson and Trojan 1982). Our study shows that the development of dental plates is quite variable, from lack of dental plates (the so-called dental ridges) to well-developed dental plates. The micro-ornamentation is also variable, either in different species or in the same species, even in the same specimen at different shell areas and therefore cannot be solely used to divide the Ambocoeliidae. In addition, microornament may be seen at two different shell layers (see Xian 1983): surficial microornament on the primary layer (spines and pustules) and inner microornament after abrasion of the shell primary layer (capillae or pits), which is evidenced in Ladjia jiwozhaiensis (Fig. 5(a3, b3, c2)) and Ambothyris sp. (Fig. 11(d6)) of Guizhou and Emanuella sp. of Xiangzhou (Fig. 5(f6, f7)). Furthermore, the same species may display different types of microornament with different degrees of erosion (Baliński 1975), which is also evidenced in Ladjia jiwozhaiensis (Fig. 5(a3, b3, c2)). Therefore, the state of shell preservation is important for correct identification of the microornamentation.

To summarise, the classification scheme of the Ambocoeliidae based on the abovementioned features (Johnson et al. 2006) has various problems and inconsistencies. Some taxa that are included in the same subfamily on the basis of similar short crural plates may be quite different in the other internal structures. For example, in the subfamily Ambocoeliinae, Ambocoelia has a pair of small crural plates and a bilobed cardinal process, whereas Guangxiispirifer has inner hinge plates united into a triangular platform and a prominent (single nodular) cardinal process. In addition, crural plates may be shown to be short or vestigial for shells with a low dorsal area and convex ventral umbo, in which crural plates are completely or mostly buried in the secondary shell thickening. This type of “lacking crural plates” is fundamentally different from “the lack of crural plates” in Crurithyris (non-ambocoelioid type dorsal hinge structure), which implies that Crurithyris may not be an ambocoelioid. Therefore, to achieve a uniform and consistent classification scheme, more characteristics need to be evaluated, e.g. nature of the cardinal process and cruralium.

Cardinal process

Cardinal process is one of the important criteria in the superfamily divisions of the Spiriferidina Waagen, 1883. The Ambocoelioidea George, 1931 was said to have a commonly simple, knoblike cardinal process (Johnson et al. 2006). Xian (1983) already noted that the Rhynchospiriferidae is characterised by a robust cardinal process, but did not give further description. Our study shows that this structure is quite varied and complicated. At least four types of cardinal process may be recognised in the Ambocoelioidea (Fig. 6).
Fig. 6

Comparison of four types of cardinal process in species that have been previously assigned to the Ambocoeliidae George, 1931. a Bilobed type (images and line drawings from Goldman and Mitchell 1990). b Triangular-knobbed type. c Ctenophoridium-like type. d Single nodular-knobbed type (images and line drawings from Paulus 1957 and Xian 1983)

  1. 1.

    Bilobed type (Fig. 6a): the cardinal process is cemented on the shell bottom, being bilobed or bi-knobbed in shape (in ventral view), e.g. Ambocoelia and Metaplasia.

  2. 2.

    Triangular-knobbed type (Fig. 6b): it is also directly cemented on the shell bottom, but triangle-shaped in ventral view, e.g. Ladjia.

  3. 3.

    Ctenophoridium-like type (Fig. 6c): it is similar to but less developed than a ctenophoridium, with an apical vacuity (terminology of Goldman and Mitchell 1990) beneath the cardinal process.

  4. 4.

    Single-knobbed type (Fig. 6d): a single-knobbed cardinal process is supported by a single plate from bottom (median septum), with or without a coronal-like structure (muscle attachment) on top of the cardinal process.


The cardinalia with the Bilobed type, Triangular-knobbed type, and Ctenophoridium-like type cardinal process is characterised by the sessile crural plates (i.e. attached to the shell bottom; Fig. 6a–c).


The cruralium is a V- or U-shaped structure formed by union of crural plates (= inner hinge plates), whereas the notothyrial cavity is the space between the two socket plates (= outer hinge plates). Based on relative position, interrelation, and specific shapes of socket plates, crural bases, and crural plates, three types may be recognised for the cruralium (Fig. 7).
Fig. 7

Types of the cruralium in dorsal transverse serial sections of the Ambocoeliidae and the relationships with types of cardinal process. a Broad-sessile type. b Narrow-sessile type. c Elevated type. Numbers are distances in millimeters from the ventral beak; pictures are in different scale; line drawings and photo images are from Goldman and Mitchell (1990), Frost and Langenheim (1966), Paulus (1957), Havlíček (1959), Johnson et al. (2006), Xian (1983), and this study

  1. 1.

    Broad-sessile type (Fig. 7a): the cruralium is low and broad, sitting on the shell bottom (Fig. 7a). The socket plates diverge laterally at an angle equal to or greater than 90°. The crural plates are short and rudimentary; the crural bases are subrounded in cross section. This type of cruralium is usually characteristic of the taxa with the Bilobed type cardinal process, e.g. Ambocoelia (here referred as the Ambocoelia-type cardinalia).

  2. 2.

    Narrow-sessile type (Fig. 7b): the cruralium is low and narrow, sitting on the shell bottom (Fig. 7b). The socket plates are from nearly parallel vertically to divergent laterally at an angle smaller than 90°, resulting in a narrow and deep cruralium and notothyrial cavity. The crural plates are relatively developed, discrete, or united with each other; the crural bases are also subrounded in cross section. Taxa with this type of cruralium are usually characterised by Triangular-knobbed type cardinal process or Ctenophoridium-like type cardinal process. The combination of the Narrow-sessile type cruralium and the Ctenophoridium-like type cardinal process is here referred as the Emanuella-type cardinalia.

  3. 3.

    Elevated type (Fig. 7c): distinct from the two previous types, this type of cruralium is characterised by poorly developed socket plates with blade-like crural bases on their inner sides and well-developed crural plates that are united to form a V- or U-shaped cruralium, supported or unsupported by a median septum. Taxa with this type of cruralium are usually characterised by single-knobbed type cardinal process, e.g. Rhynchospirifer (here referred as the Rhynchospirifer-type cardinalia).


Revised classification

In the classification of Johnson et al. (2006), taxa of the Rhynchospirifer-type cardinalia are placed in both subfamilies, e.g. Guangxiispirifer of the Ambocoeliinae and Rhynchospirifer of the Rhynchospiriferinae, whereas taxa with different types of cardinalia were placed in the same subfamily, e.g. Guangxiispirifer and Ambocoelia in the Ambocoeliinae and Rhynchospirifer and Emanuella in the Rhynchospiriferinae. Our study supports the notion of Xian (1983) who elevated Rhynchospiriferinae Paulus, 1957 to the family level since their Rhynchospirifer-type cardinalia is a distinct character that distinguishes them from the rest of the Ambocoeliidae, which is, as represented by Ambocoelia, Emanuella, and Diazoma (previously in the Ambocoeliinae and the Rhynchospiriferinae, respectively), characterised by the sessile crural plates.

Superfamily Ambocoelioidea George, 1931

Family Ambocoeliidae George, 1931

Crural plates variably present, vestigial or well developed, discrete or connected with each other on the valve floor; cruralium of the Broad- and Narrow-sessile types; cardinal process varied, Bilobed, Triangular-knobbed, and Ctenophoridium-like types; crural bases commonly rounded; dental plates commonly lacking, except for a few genera in which they are weakly developed.

Genera included (those present in the Devonian of South China):Ambocoelia Hall, 1860, Ambothyris* George, 1931, Aviformia Xian, 1988, Choperella* Liashenko, 1969, Crurispina* Goldman and Mitchell, 1990, Cyrtinoides Yudina and Rzhonsnitskaia, 1985, Diazoma* Dürkoop, 1970, Emanuella* Grabau, 1923, Ilmenia* Nalivkin, 1941, Ilmospirifer* Liashenko, 1969, Ladjia* Veevers, 1959, Moravilla* Havlíček, 1953, and Zhonghuacoelia* Chen, 1978 (genera with an asterisk were previously in the Rhynchospiriferinae of Johnson et al. (2006); Crurithyris George, 1931 is probably not a ambocoelioid because of its non-ambocoelioid type dorsal hinge structure).

Family Rhynchospiriferidae Paulus, 1957

[nom. transl. Xian in Xian and Jiang 1978, ex Rhynchospiriferinae Paulus, 1957]

Crural plates developed, commonly joined to form an Elevated type cruralium supported by median septum or suspended from valve floor anteriorly; cardinal process of the Single-knobbed type; crural bases commonly elongated and blade-like in transverse section; dental plates commonly well developed (except in Guangxiispirifer and Amboglossa: weak or vestigial).

Genera included (those present in the Devonian of South China):Rhynchospirifer Paulus, 1957, Amboglossa Wang and Zhu, 1979, Changtangella Xian, 1983, Levibiseptum Xian in Hou and Xian, 1975, Ilmeniopsis Xian, 1983, Ilmenispina Havlíček, 1959, Kosirium Ficner and Havlíček, 1975, and Guangxiispirifer* Xian, 1983 (genus with an asterisk was previously in the Ambocoeliinae of Johnson et al. 2006).

Origination and diversification of the ambocoelioids in South China

Immigration, endemism, and originations

North America is apparently the centre of origin for the Ambocoelioidea (Fig. 3). We agree that the tiny, small, smooth Ambocoelioidea evolved directly from the cyrtioids in the middle Silurian by the paedomorphic loss of ribbing (Carter and Gourvennec 2006) and that specifically the Eospiriferinae may be the ancestor (Pitrat 1965, p. H667; Johnson and Lenz 1992). According to Johnson and Lenz (1992), Gourvennec (2000), and Johnson et al. (2006), the oldest ambocoelioid genus is Eoplicoplasia in the late Wenlock of the Silurian, which is assigned to the Ambocoeliinae on the basis of its lacking crural plates. However, the so-called lack of crural plates needs further clarification because no serial sections were presented. For example, Goldman and Mitchell (1990) demonstrated through serial sections that Ambocoelia has short crural plates, which had commonly been known or described to lack crural plates. Dicoelospirifer (late Wenlock of Canadian Arctic Islands) serves as another alternative ancestor of the smooth-shelled ambocoeliids (Zhang 1989b) since its type species possesses short crural plates (Zhang 1991), although the author gave the statement “lacks crural plates” in the genus diagnosis. Zhang (1991) mentioned that, “It shows that from the very beginning the Ambocoeliinae were divided into non-plicate and plicate forms. It is possible that Dicoelospirifer is an ancestor of non-plicate ambocoeliids while Plicoplasia (=Eoplicoplasia; our note: Plicoplasia cf. acutiplicata of Zhang 1989b is reassigned to Eoplicoplasia) is an ancestor of plicate ones.” Compared with the non-plicate Dicoelospirifer, plicate Eoplicoplasia is more likely to be the ancestor of the smooth ambocoelioids for the following reasons. Plications decreased in strength in early intermediate forms of the ambocoelioids, e.g. Metaplasia hemicona (Fig. 3a, Havlíček 1990, from Czechia), which is considered to be (upper) Ludlow (Manda and Kříž 2006; Manda and Frýda 2014; Kraft and Kvaček 2017) to earliest Pridoli in age, Pridoli Metaplasia sp. (Lenz 1970, from Yukon), Lochkovian Ambothyris? inopis (Fig. 3: b, Mawson and Talent 1999, from southeastern Australia; our note: it should probably be assigned to Ambocoelia or Metaplasia), Metaplasia cf. lenzi (Fig. 3c, Lenz and Johnson 1985, from southeastern Australia) and Ambocoelia bubovica (Fig. 3d, Mergl 2003, from Bohemia), Pragian Ambocoelia aff. praecox (Fig. 3f, Lenz 1977, from Yukon). Dicoelospirifer differs from other early ambocoelioids in its bilobed-emarginated outline and poorly developed ventral interarea. The third earliest smooth-shelled ambocoeliid is Metaplasia, which represented an intermediate stage from the plicate to smooth forms. In the Silurian species Me. hemicona, the plications are still present, but much weaker than those of Eoplicoplasia (Fig. 2(b)); therefore, Metaplasia of the Ludlow is probably an intermediate genus linking to the Devonian forms.

The earliest three ambocoelioids in South China are Amboglossa, Prolazutkinia, and Ambothyris in the upper Emsian. The first two genera both have the Rhynchospirifer-type cardinalia and originated in South China, representing respectively two different families (Fig. 8). Amboglossa (Fig. 3: genus no. 12) invaded the Russian Platform and Urals during the Eifelian. Amboglossa has a high ventral interarea and narrow furrow on both valves, which may be derived from an ambocoeliid of Australia (probably Ambocoelia) and Ambothyris may directly migrate to South China from Australia.
Fig. 8

Temporal and spatial distribution of ambocoelioids in South China. a, b Palaeogeographic distribution during the upper Emsian–Lower Givetian (legends the same as in Fig. 1; lithofacies map modified from Ma et al. 2009b). c Stratigraphic distribution, diversity change, and possible phylogenetic relationships of ambocoelioids against regional sea level changes of Ma et al. 2017

During the Devonian, there were not only diverse cosmopolitan genera such as Cyrtinoides (Fig. 3: genus no. 6), Emanuella (Fig. 3: genus no. 17), Ambothyris (Fig. 3: genus no. 13), and Ladjia (Fig. 3: genus no. 22), but also numerous endemic genera (Fig. 3: genus nos. 27–29). It should be pointed out that those cosmopolitan genera are generally ambocoeliids whereas the rhynchospiriferids are basically restricted and short lived and have not been found in Laurentian (North America) and Gondwana, including Rhynchospirifer (Fig. 3: genus no. 11), Kosirium (Fig. 3: genus no. 21), Amboglossa (Fig. 3: genus no. 12), and Guangxiispirifer (Fig. 3: genus no. 30). In the latest Eifelian and Early Givetian, the Rhynchospiriferidae became diversified with quite a few endemic genera (Changtangella, Ilmeniopsis, and Levibiseptum) and lasted shortly before its final disappearance in South China at the end of the Early Givetian (Fig. 8).

The few endemic ambocoeliid genera include Zhonghuacoelia and Aviformia (poorly preserved). Zhonghuacoelia probably originated from Ambocoelia in light of its Ambocoelia-type cardinalia, whereas those cosmopolitan ambocoeliid genera are immigrants at various marine transgression episodes. The only genus that crosses the Frasnian–Famennian boundary in South China seems to be Ambocoelia. True Crurithyris probably first appeared in South China in the latest Famennian with a different cardinalia structure, which, along with other post-Devonian forms such as Attenuatella, Biconvexiella, and Cruricella, may represent a different evolutionary lineage.

Ladjia and Emanuella are very similar externally; their major difference lies in that the former is characterised by the Triangular-knobbed cardinal process, which implies that Ladjia should be a direct descendant of Emanuella. Emanuella and Diazoma are characterised by the Ctenophoridium-like type cardinal process and may be derived from Ambothyris. The Emanuella-type cardinalia having a more complicated cardinal process seems more advanced than the Ambocoelia-type cardinalia, which suggests that taxa of the former type are derived from the Ambocoelia-type taxa. The earliest genus of the Emanuella-type taxa is Ambothyris (Chatterton 1973, from the Receptaculites and Warroo limestones at Taemas, Fig. 3g) in the early Emsian of Australia (Lindley 2002, p. 111). Therefore, the differentiation of the Emanuella-type from the Ambocoelia-type should take place no later than (during or before) the early Emsian. Morphologically, all rhynchospiriferids are characterised by the Rhynchospirifer-type cardinalia. As the earliest genus of the Rhynchospiriferidae, Amboglossa possesses a median furrow on both valves, which also can be observed in Ambothyris, implying the high possibility that Rhynchospirifer-type forms are derived from the Emanuella-type forms. Such characteristics were also present in the lower Eifelian Guangxiispirifer bisinuatus but disappeared in the Lower Givetian Guangxiispirifer (Xian 1983). Therefore, Guangxiispirifer should be derived from Amboglossa. Other non-furrow rhynchospiriferid genera suddenly emerged at the basal Mintang Fm., which should originate from Rhynchospirifer. Prolazutkinia and Lazutkinia together constitute an evolutionary lineage because of their strong plication. Rhynchospiriferids and lazutkiniids are both characterised by the Rhynchospirifer-type cardinalia; therefore, they are closely related and possibly derived from the Ambocoeliidae.

To sum up, the origin of the Ambocoelioidea took place in northern North America during the Wenlock. Within the superfamily, various taxa may have their own place and time of origin. When early immigrants entered a new habitat, they may rapidly evolve into new forms. These new taxa may further expand their distribution into other continents. The Rhynchospiriferidae is such an example, which originated in South China and migrated into the Baltica continent. In light of the cardinalia evolution, the Ambocoelia-type taxa are ancestral forms for all the Ambocoelioidea. The differentiation of the Emanuella-type from the Ambocoelia-type should take place no later than (during or before) the early Emsian. The origination of the Rhynchospirifer-type taxa took place in South China and their differentiation from the Emanuella-type taxa probably happened in the upper Emsian.

Diversification and sea level changes

In South China, the first ambocoelioids Amboglossa and Prolazutkinia occurred in the upper part of the Dale Formation (upper Emsian), which consists of deposits indicative of a sea level rise environment (Dale sea level rise of Ma et al. 2017). The first Ambothyris appeared in the lower Dangduo Fm. of the Tewa area (Fig. 8), which is mainly composed of biogenic limestone and was probably also related to the Dale sea level rise (Fig. 8; Ma et al. 2017). The primary differentiation of the rhynchospiriferids is the appearances of Guangxiispirifer and Rhynchospirifer in the early Eifelian, probably related to the Gupa transgression (Chotec Event) at the end of the Po. partitus Zone (Ma et al. 2017). These two rhynchospiriferid genera are important for the subsequent radiation of the Rhynchospiriferidae in South China towards the end of the Eifelian, which was probably related to the Mintang transgression (Kacak Event) in the Po. ensensis Zone (Ma et al. 2017). After this radiation, the Rhynchospiriferidae reaches its highest diversity at the end of the Eifelian through Lower Givetian (six to seven genera) and apparently died out immediately afterwards in the main part of South China. This regional extinction happened within the Po. hemiansatus Zone (conodont data of Bai et al. 1994 in the Liujing section), which might be related to the Songjiaqiao sea level fall (Fig. 8). In the Qinling region (e.g. the Tewo area), rare rhynchospiriferids may range slightly higher, but still within Lower Givetian. In contrast, the Ambocoeliidae, which has more cosmopolitan genera, had no radiation event; its diversity increased gradually and reached the highest diversity (six genera) in the Middle–Upper Givetian. Ambocoeliid diversity started to decline in the Early Frasnian and continued towards the end of the Famennian.

Ambocoelioids are a group of strongly facies-controlled brachiopods. Their emergence and demise show a close relationship with sea level changes. The ambocoeliids are generally preserved in marl and shale intervals suggestive of deeper dysoxic environments, whereas rhynchospiriferids-bearing strata are often associated with reefal limestones, representing a high-energy and well-oxygenated environment (Xian 1983). Their different adaptions probably resulted in their different responses to environmental changes. The rhynchospiriferids were probably more sensitive to sea level changes, therefore prone to rapid and regional originations and extinctions. The ambocoeliids, inhabiting deeper water environments and having more cosmopolitan genera, underwent gradual originations, extinctions, and diversity changes.

Systematic palaeontology

All specimens with the prefix PUM are stored in the Geological Museum of Peking University.

Family Ambocoeliidae George, 1931

Genus Ambocoelia Hall, 1860

Ambocoelia yidadeensis sp. nov.

(Figs. 9(a–j) and 10)
Fig. 9

Ambocoelia yidadeensis sp. nov. from the middle Yidade Formation (Lower Frasnian), Panxi section. The nine specimens are to show convexity variations in the ventral umbonal area in ventral and lateral views. (a1–a5) Dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16101. (b1–b5) Dorsal, ventral, anterior, posterior, and lateral views of the paratype (PUM16103). (c1–c5) Dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16102. (d1–d4) Dorsal, lateral, microornament enlargement of d1, and microornament enlargement of d2 of specimen PUM16104. (e1–e5) Dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16106. (f1–f5) Dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16107. (g1–g5) Dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16108. (h1–h5) Dorsal, ventral, anterior, posterior, and lateral views of the holotype (PUM16109). (i1–i5) Dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16110. (j1–j2) Dorsal and enlarged posterior views of specimen PUM16105

Fig. 10

Transverse serial sections of Ambocoelia yidadeensis sp. nov. from the middle Yidade Formation (Lower Frasnian) of the Panxi section. a PUM16110 (Fig. 9(i)). b PUM16102 (Fig. 9(c)). Numbers refer to distance in millimeters from the ventral apex

Material: Ten specimens from the Yidade Formation of the Panxi section, including two sectioned specimens (Figs. 9(c, i) and 10a, b).

Derivation of name: Name after Yidade, a village, which is the type locality of the Panxi section.

Holotype: PUM16109 (Fig. 9(h)).

Diagnosis: Shell material thick in ventral umbo, ventral interarea relatively high, moderately curved; fold and sulcus lacking; spine bases broken off (giving a pitted appearance), which are densely arranged in a quincunx shape; faint striae widely spaced on exfoliated dorsal shell.

Description: Small-sized, generally around 10 mm in width; ventral valve more convex than dorsal valve, especially in the umbonal area; ventral umbo weakly inflated, beak strongly incurved; ventral interarea high; apical plate closing delthyrium apex, remains of deltidial plates along delthyrial edges; dorsal interarea very low; fold and sulcus lacking, some specimens displaying a shallow median depression near anterior margin on both valves; commissure rectimarginate; faint concentric growth lamellae; pits (after spine bases broken off) on primary layer, uniform in size, densely arranged in a quincunx shape; widely spaced faint radial striae on exfoliated dorsal valve.

Internally only dental ridges developed; teeth strong, triangular; cardinal process of the Bilobed type; hinge sockets deep, outer socket ridges partly fused to valve floor posteriorly; crural bases rounded, crural plates short; cruralium of the Broad-sessile type; crura small, rounded; spiralia about six–seven whorls.

Discussion: the new species differs from the type species Ac. umbonata in the higher and flatter ventral interarea, more convex dorsal valve and more indistinct median furrow. Ac. elongata Zhang, 1983 from the upper Eifelian Lure Fm. is smaller in size and has an elongated outline. Ac. pseudosinensis Zhang, 1983 from the Frasnian Lengshuihe Fm. shares some similarities externally, but differs from our specimen in having a shorter hinge line, more convex dorsal valve and more elevated socket plates.

Occurrence: Middle Yidade Formation of the Panxi section (stratigraphic column see Zhang et al. 2015, Panxi-Y2, Bed 9), Lower Frasnian.

Genus Ambothyris George, 1931

Ambothyris panxiensis (Chu, 1974)

(Figs. 11(a–c) and 12)
Fig. 11

acAmbothyris panxiensis (Chu, 1974) from the Eifelian Yingtang Formation, Baqi section, Xiangzhou. (a1–a5) Dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16219; (b1–b5) dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16220; (c1–c5) dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16221. dAmbothyris sp. from the Hejiazhai Mbr (Lower Frasnian) of the Dushan section. (d1–d6) dorsal, ventral, anterior, posterior, lateral views, and microornament (slit-like furrows arranged radially: white arrows) on the exfoliated shell PUM16245. eAmbothyris yunnanensis (Chu, 1974) from the Middle Givetian Qujing Formation, Panxi section; (e1–e5) dorsal, ventral, anterior, posterior, and lateral views of specimen PUM16135. fRhynchospirifer sp. from the Nanpanjiang Formation (probably upper Eifelian) of the Panxi section. (f1–f6) Dorsal, ventral, anterior, posterior, lateral views, and microornament of specimen PUM16132

Fig. 12

Transverse serial sections of Ambothyris panxiensis (Chu, 1974) from the Eifelian Yingtang Formation, Baqi section. PUM16220 (Fig. 11(b))

1974 Ambocoelia panxiensis Chu; Fang and Zhu, p. 414, pl. 135, fig. 8–9.

1974 Ambothyris yunnanensis Wang et al. p. 244, pl. 124, fig. 6–9.

1982 Emanuella aff. transversa, Bai et al. p. 12–13.

Material: Ten specimens from the Yingtang Formation of the Baqi section, including one sectioned specimen (Figs. 11(b) and 12).

Description: Small-sized, generally around 5–10 mm in width, transverse in outline; dorsal valve arch-shaped; sub-equally biconvex; ventral beak small, strongly incurved; ventral interarea high; dorsal interarea low; fold and sulcus lacking, except for a faint median furrow on both valves; commissure rectimarginate or slightly sulcate; faint concentric growth lamellae.

Internally dental plates lacking, dental ridges strongly developed; very small apical plate closing the delthyrium apex; cardinal process of the Ctenophoridium-like type, apical vacuity developed; hinge socket plates high, crural plates short, cruralium of the Narrow-sessile type, crural bases rounded; spiralia at least six whorls.

Discussion: This species differs from other species such as At. yunnanensis (Chu in Fang and Zhu 1974), At. parvissima (Yang, 1988) and At. sp. (Fig. 11(d)) in its transverse outline and the arch-shaped dorsal valve. At. longmenshanensis has a very high ventral interarea, which is easily distinguishable from At. panxiensis. Our examination of specimens of Emanuella aff. transversa listed in Bai et al. (1982, pp. 12–13) and Bai et al. (1994, p. 141) shows that they clearly differ from Emanuella in the higher and flatter ventral interarea and that they should be assigned to Ambothyris panxiensis.

Ambothyris panxiensis (Chu, 1974) and Ambothyris yunnanensis Wang et al., 1974 are synonyms because both of them are from the Qujing Fm. of the Panxi section and are very similar morphologically. The two publications appeared in the same time (all November of 1974). As the first revisers, we choose Ambothyris panxiensis (Chu, 1974) as the valid species name. This can avoid to change the name Ambocoelia yunnanensis Chu in Fang and Zhu 1974 (which is a valid species and should also be assigned to Ambothyris). Ambothyris yunnanensis (Fig. 11(e)) can be distinguished from At. panxiensis (Fig. 11(a–c)) by its subpentagonal outline in adult individuals.

Occurrence: Yingtang Formation, Baqi section, Xiangzhou, Eifelian.

Family Rhynchospiriferidae Paulus, 1957

Genus Rhynchospirifer Paulus, 1957

Rhynchospirifer sp.

(Figs. 11(f) and 13)
Fig. 13

Transverse serial sections of Rhynchospirifer sp. from the Nanpanjiang Formation (Panxi section), upper Eifelian, PUM16132 (Fig. 11(f))

Material: One specimen.

Diagnosis: Inflated; cardinal extremities rounded; ventral interarea moderately high and strongly incurved; microornament of concentric rows of dense spiny bases on primary layer.

Description: Small- to medium-sized, ventribiconvex, inflated; cardinal extremities rounded; maximum width near the posterior margin; ventral interarea moderately high and strongly incurved; fold and sulcus lacking; front commissure rectimarginate; faint concentric growth lamellae; microornament of concentric rows of dense spiny bases on primary layer.

Internally dental plates convergent ventrally; cardinal process of the Single-knobbed type (ridge-shaped anteriorly); outer socket ridges fused to valve floor posteriorly; socket plates short, crural bases blade-shaped, crural plates forming an Elevated type cruralium, which is V-shaped, supported by a median septum; muscle scars deep.

Discussion: The species has a rounded cardinal extremity and strongly incurved ventral interarea; thus, it differs from other Chinese species such as Rh. liujingensis, Rh. trigonus, and Rh. hengxianensis and may represent a new species. Rh. regularis is the most similar species, but differs from the species in the sub-rectangular outline and having a wide flat depression near the anterior margin on both valves.

The species differs from the type species Rh. halleri in having an inflated shell a wider hinge (thus, maximum width is near the posterior margin) and a V-shaped cruralium (U-shaped in the type species).

Occurrence: Nanpanjiang Fm. (probably upper Eifelian) of the Panxi section


  1. 1.

    The origination of the Ambocoelioidea took place in northern North America during the Wenlock. The earliest form is the plicated ambocoeliid Eoplicoplasia from Canadian Arctic Islands. The second earliest largely smooth shelled ambocoeliid Metaplasia of the Ludlow is probably an intermediate genus linking to the Devonian forms.

  2. 2.

    Our study shows that the cardinal process and cruralium are quite varied and complicated. At least four types of the cardinal process (Bilobed, Triangular-knobbed, Ctenophoridium-like, and Single-knobbed type) and three types of cruralium (Broad-sessile, Narrow-sessile, and Elevated types) are recognised in the Ambocoelioidea. Based on differences in the cardinal process and cruralium, three types of cardinalia, Ambocoelia-type, Emanuella-type, and Rhyncospirifer-type, can be distinguished. It is justified to elevate the Rhynchospiriferinae Paulus, 1957 to the family level since their Rhynchospirifer-type cardinalia (with the Single-knobbed cardinal process and the Elevated type cruralium) is a distinct character that distinguishes them from the rest of the Ambocoeliidae George, 1931.

  3. 3.

    The temporal and spatial distributions of the superfamily Ambocoelioidea are depicted globally and in South China. Three earliest ambocoelioids in South China are Ambothyris, Amboglossa, and Prolazutkinia in the upper Emsian; the latter two genera originated in South China, and Amboglossa is probably the ancestor of the whole Rhynchospiriferidae family. In light of the cardinalia evolution, Ambocoelia-type is the first of all ambocoelioids. The differentiation of the Emanuella-type from the Ambocoelia-type cardinalia should take place in the early Emsian or before. The Rhynchospiriferidae (characterised by the Rhynchospirifer-type cardinalia) originated in South China in the late Emsian. The Rhynchospiriferidae and the Lazutkiniidae are both characterised by the Rhynchospirifer-type cardinalia and therefore are closely related, but they were possibly independently derived from the Ambocoeliidae.

  4. 4.

    The Ambocoelioidea is a group of strongly facies-controlled brachiopods. Their emergence and demise show a close relationship with sea level changes. The first appearance of the ambocoelioids in South China was in the late Emsian, which was probably related to the Dale sea level rise (=Upper Zlichov Event). The radiation of the endemic rhynchospiriferids in the basal Mintang Fm. was apparently related to the Mintang transgression (=Kacak Event). Because of different environmental adaptations, ambocoeliids and rhynchospiriferids show different modes of diversity variation and extinction. The ambocoeliids inhabited deeper and dysoxic water environments and showed greater capabilities to migrate and survive extinctions, and therefore, they are largely cosmopolitan in distribution. By contrast, rhynchospiriferids prefer a high-energy and well-oxygenated water environment; thus, they are probably more sensitive to sea level changes, and as a result, they are mostly endemic in distribution, being prone to rapid and regional extinctions.




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 information

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

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.


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Authors and Affiliations

  1. 1.Key Laboratory of Orogenic Belts and Crustal Evolution, School of Earth and Space SciencesPeking UniversityBeijingChina

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