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Palaeobiodiversity and Palaeoenvironments

, Volume 99, Issue 1, pp 91–99 | Cite as

Palaeobiogeographic dynamics of brachiopod faunas during the Frasnian-Famennian biotic crisis in South China

  • Li QiaoEmail author
  • Wenkun Qie
Original Paper
  • 90 Downloads

Abstract

The Late Devonian biotic crisis has been studied for a long time mainly on the basis of its taxonomic severity, but with little attention paid on the relationship between biogeography and evolutionary dynamics. In this study, brachiopod faunas across the biotic crisis are surveyed based on revised data compiled from 15 sections in South China. Data of brachiopod faunas are plotted using a network method to detect biogeographic variation and dynamics through the upper Frasnian, lower and middle Famennian. In addition, frequency analysis is performed on the occurrence of brachiopod faunas within the different localities and depositional settings through time. These data show significant taxonomic loss in the five depositional zones in South China, particularly in the littoral clastic and restricted-platform facies. Refuges during the survival interval were probably located in deeper water zones, especially in the inter-platform basin, but also in the peritidal and the open carbonate platform. A decrease of biogeographic connections within the brachiopod occurrence network, accompanied with an increase of low frequency taxa in the survival and recovery intervals, supports biogeographic divergence of ecological preferences of survival and recovery faunas in South China, as well as a turnover of brachiopod faunas through the biotic crisis. Reasons for variation of biogeographic distribution and connections are less known, besides the faunal colonisation that facilitated by sea-level fluctuation through the biotic crisis.

Keywords

Biogeography Network Late Devonian Mass extinction 

Introduction

The Frasnian-Famennian mass extinction (hereinafter “F-F biotic crisis”) has been originally ranked among the “big five” mass extinction events according to biodiversity loss by Raup and Sepkoski (1982). To date, discussions on the evolutionary dynamics of the F-F biotic crisis and associated various reasons are still being discussed. Recent studies focus on the ecological impacts which have degraded the taxonomic severity of the Late Devonian mass extinction, as emphasised by low species origination and speciation rates (Bambach et al. 2004; Stanley 2007; McGhee et al. 2012, 2013). On the other hand, knowledge on the biogeographic distribution during the biotic crisis is limited, which prevents understanding of the relationship between biogeography and evolutionary dynamics in the brachiopod fauna. In addition, the Late Devonian has long been recognised as interval of strong faunal cosmopolitanism during which marine faunas attained wide geographic coverage driven by wide faunal dispersal. The widespread distribution of taxa was consequently considered to be correlative to a decrease of endemism and the loss of the total diversity across the F-F biotic crisis based on regional data from North America (Johnson 1970; Stigall 2010, 2012).

To understand the biogeographic dynamics and its relationship with faunal turnover and recovery during the biotic crisis, we carry out a regional study on the brachiopod fauna from South China. This study focuses on the regional brachiopod distribution through the F-F biotic crisis, based on brachiopod occurrences from 15 sections representing various depositional environments in the late Frasnian, early Famennian, and middle Famennian. The patterns of brachiopod biogeography before and after the biotic crisis are detected by the network method, with interests in the changing connections and biogeographic distribution, which provide an opportunity to assess the relationship between the brachiopod diversity, distribution and their environmental background at both temporal and spatial scales.

Data and temporal framework

Devonian brachiopods have been investigated in various sedi-mentary settings of South China. The database compiled here include a large number of brachiopod genera and species that have been reported mostly in monographs and systematic papers, as well some lists from the reports on regional geology. Brachiopod genera have been revised herein following the Revised Brachiopod Treatise and more importantly by the “Phanerozoic brachiopod genera of China” published recently (Hou et al. 2017 in Rong et al. 2017) which proposed alternative systematic criteria to some old taxa, e.g. the most common genera of yunnanellides or cyrtospiriferides in the Frasnian and Famennian. The range and distribution of the revised brachiopod genera are given in Fig. 1 (also see Table 1 for detailed lists of genera).
Fig. 1

Biostratigraphic framework and zonation adopted in this study (brachiopod zones modified from Ma and Zong 2010; Hou et al. 2017; Qie et al., in press), with abundance of the revised brachiopod genera showing range and distribution of brachiopods through the F-F biotic crisis in South China

The Late Devonian sequences in South China have been studied and updated recently, allowing a credible biostratigraphic framework and taxonomic occurrence lists of brachiopods. Moreover, comprehensive studies involving systematic, biostratigraphic, litho-, and bio-facies have improved subdivision and correlation of conventional zonations of benthic faunas (Hou et al. 1985; Bai et al. 1994; Xian et al. 1995; Ma and Bai 2002; Liao and Ruan 2003; Hou and Ma 2005; Ma and Sun 2008; Ma and Zong 2010; Ma et al. 2016). We here have subdivided the brachiopod genera into three time intervals across the Frasnian and Famennian biotic crisis, mainly following the conodont or brachiopod zonation as shown in Fig. 1.

In South China, the Frasnian and Famennian brachiopods have been recorded in various depositional environments spanning a proximal to distal transect across the sedimentary basin. In this study, we have selected 15 sections (Table 1) representing different sedimentary settings that can be categorised into I to V (see Fig. 2), whereas, benthic faunas are absent in the deeper water Qinzhou-Fangcheng trough (category VI) which are characterised by siliceous rocks and argillaceous mudstone across the F-F transition. In the littoral clastic-dominated settings (I), brachiopod data have been selected from Yongxin and Huangjinpin in central Hunan, and in Zhenba at the north-western margin of South China (sections 1, 2, 15 in Fig. 2). The late Frasnian and Famennian brachiopods largely inhabited the open carbonate platform (II) and (semi-)restricted platform (III), as have been recorded in the Xikuangshan and Qiziqiao sections (3, 4), and in the Shujiapin, Mujintang, Tangjiawan, Dushan and Lu’nan sections (7, 8, 9, 11, 12), respectively. The brachiopod faunas from the Shetianqiao and Jiangjiaqiao sections (5, 6) have been associated with inter-platform basin (Yu et al. 1990; Ma and Zong 2010; Ma et al. 2016), whereas those from the Debao section (10) are related to the deep-water marginal slope or peri-platform basin facies (IV), and consist of nodule limestone, thin to middle-bedded marls and siliceous rocks. At the northwestern margin of the South China Block, the late Frasnian sequences, containing brachiopods of the Schizophoria-Gypidula-Atrypa and Cyrtospirifer assemblages in Ganxi and Jiangyou sections (13, 14) in Sichuan, are assigned to peritidal and lagoonal environments (V) (Chen 1983, 1984; Hou et al. 1985; Xian et al. 1995). Table 2 lists the number of genera in the five sedimentary settings.
Fig. 2

Palaeogeographical map of South China in the Late Devonian, showing distribution of the selected brachiopod faunas across the Frasnian-Famennian biotic crisis. Sections name as indicated in the text (see also Table 1) and depositional categories are compiled from Zhao (1985), Wu et al. (1997), Ma and Zong (2010) and Yang (2014)

Table 2

Number of brachiopod genera assigned to palaeoenvironmental categories I to V in South China through the Frasnian-Famennian biotic crisis intervals

Interval\sedimentary category

Littoral clastic facies (I)

Carbonate platform (II)

Restricted platform (III)

(Peri-) or inter-platform (IV)

Peritidal/lagoon (V)

Middle Famennian

11

7

21

12

0

Lower Famennian

0

3

0

13

5

Upper Frasnian

14

12

18

19

9

Table 1

Revised brachiopod genera recorded from different sections in South China. Section name: 1, Yongxin; 2, Huangjinpin; 3, Xikuangshan; 4, Qiziqiao; 5, Shetianqiao; 6, Jiangjiaqiao; 7, Shujiapin; 8, Mujintang; 9, Tangjiawan; 11, Dushan; 12, Lu’nan; 13, Ganxi; 14, Jiangyou; 15, Zhenba. A, upper Frasnian; B, lower Famennian; C, middle Famennian. Brachiopods from Debao section (10) were recorded but are not analysed here due to lack of systematic description

Genus

Section

1

2

3

4

5

6

7

8

9

11

12

13

14

15

Interval

A

C

A

B

C

A

A

B

A

B

C

C

C

A

A

A

C

A

B

A

C

Athyris

×

  

×

 

×

×

 

×

×

 

×

×

×

 

×

×

×

   

Atrypa

×

 

×

   

×

      

×

×

  

×

 

×

 

Camarotoechia

×

×

             

×

 

×

×

×

 

Chonetes

×

          

×

  

×

      

Costatrypa

      

×

              

Cyrtiopsis

       

×

 

×

×

×

×

     

×

  

Cyrtospirifer

×

×

×

 

×

×

×

  

×

×

×

×

×

×

  

×

 

×

 

Desquamatia

×

 

×

  

×

×

 

×

     

×

    

×

 

Dmitria

           

×

         

Gypidula

        

×

     

×

×

 

×

   

Hunanospirifer

 

×

        

×

×

      

×

  

Hunanotoechia

      

×

              

Hypothyridina

     

×

×

 

×

    

×

   

×

   

Iowatrypa

      

×

 

×

            

Lamarckispirifer

         

×

           

Leioproductus

         

×

×

          

Leptostrophoria

                

×

    

Mennespirifer

      

×

              

Monelasmina

                   

×

 

Nayunnella

 

×

  

×

     

×

 

×

       

×

Platyspirifer

         

×

×

×

         

Plectorhynchella

                

×

    

Plicapustula

         

×

           

Praewaagenoconcha

         

×

×

          

Productella

×

 

×

  

×

×

×

×

×

 

×

×

 

×

 

×

   

×

Productellana

 

×

  

×

     

×

    

×

    

×

Ptychomaletoechia

    

×

×

×

 

×

×

×

×

×

  

×

×

   

×

Pugnax

     

×

  

×

   

×

  

×

×

    

Pyramidaspirifer

     

×

               

Radiatrypa

              

×

      

Rhyssochonetes

      

×

              

Schizophoria

  

×

   

×

      

×

 

×

×

  

×

 

Schuchertella

×

×

       

×

×

×

×

        

Sinoproductella

          

×

          

Sinospirifer

×

×

  

×

×

 

×

×

×

  

×

    

×

 

×

 

Spinatrypa

×

 

×

  

×

×

 

×

     

×

  

×

   

Spinatrypina

     

×

  

×

    

×

 

×

     

Stenoscisma

                

×

    

Tenticospirifer

×

 

×

   

×

    

×

×

×

 

×

 

×

×

  

Theodossia

×

    

×

        

×

      

Uchtospirifer

           

×

         

Xinshaoella

    

×

           

×

   

×

Yunnanella

 

×

  

×

  

×

 

×

×

×

      

×

 

×

It should be mentioned that brachiopods from the carbonate platform margin and slope are insufficiently sampled compared with those from the carbonate platform settings, showing the overall lower diversity and abundance. Recorded genera from the Debao section are represented by rhynchonellides in both pre- and post-extinction intervals, with no systematic description or revision, thus are excluded from the network and frequency analysis.

Network method for assessing biogeographic structure

The network method has been applied in evaluating regional or global biogeographic structure and faunal connections and provincialism (e.g. Sidor et al. 2013; Vilhena and Antonelli 2015; Kiel 2017; Huang et al., 2018). It converts faunal distributional data as a bipartite network and builds connections between species-locality and locality-locality. The biogeographic links between localities (nodes) are represented as lines which run from the taxa they contain. Therefore, the network method incorporates both locality and taxa presence-absence information, and visualises directly the network diagrams similarities and differences between faunas.

In this study, the biogeographical data were compiled into a presence/absence matrix of each genus in each of the sections, and then were quantitatively assessed by the Gephi software package (www. Gephi.org). The bipartite density introduced by Sidor et al. (2013) as BC, and network graphic density (GD) are also calculated to quantify the biogeographic connectivity and endemicity before and after the biotic crisis. The bipartite density is dependent on the links in the network, with a low value close to zero suggesting a high proportion of endemic taxa (Sidor et al. 2013). The graphic density is positively related to the number of links between nodes, with high values being obtained when the faunas are intensively connected. Besides, the modularity (MD) is calculated to assess the network clustering (Blondel et al. 2008). When positive and increased modularity values are obtained, it means that the nodes are tightly connected or clustered and are more easily recognised as a distinct biogeographic biochore.

Biogeographic distribution and dynamics across the biotic crisis

The brachiopod biogeographic network integrates locality data and related generic occurrence data, providing a visualisation of both diversity and biogeographic connections. The late Frasnian brachiopods distributed from the nearshore area to offshore settings in South China show general connections, as complex network links constructed by a percentage of genera that are shared by at least two localities (Fig. 3a). It is noteworthy that even the section with the largest number of genera (section 5) has a lower proportion of “endemic” taxa, compared with other sections which contain lower generic diversity. Immediately after the biotic crisis, brachiopods are absent in most of localities in South China, with rare genera being recorded in four localities, which are not sufficient for network analysis. The post-extinction brachiopod faunas here referred to are those in the middle Famennian of the network analysis (Fig. 3b), showing remarkable changes in both geographic range and biogeographic connections of brachiopod faunas. Biogeographic links in the middle Famennian are reconstructed mainly among a few sections (6, 7, 8), and depict a trend of divergence of ecological preference of brachiopod faunas after the biotic crisis. Further investigation on the number of brachiopod genera from different depositional settings including littoral clastic, open platform, restricted platform, (peri- and) inter-platform bain, lagoon or peritidal zones reveals that the brachiopods survived largely in the deeper water regions, especially in the inter-platform facies which had the highest diversity (Table 2 and Fig. 4a, b).
Fig. 3

Network diagrams showing the biogeographic distribution and connection of brachiopod faunas in South China before the biotic crisis (a, late Frasnian) and after extinction (b, middle Famennian), whereas those in the lower Famennian are not calculated due to the few genera documented as shown in Table 1. Blue circles denote sections 1−15 as indicated in Fig. 1, with the various sizes corresponding to generic richness; grey circles denote genera recorded at a single locality, yellow circles denote genera shared in two or more sections

Fig. 4

a Number of brachiopod genera (marked in circles) recorded in sections 1–15 (horizontal axis) in three time intervals (noted by different colours) across the extinction event. b Number of brachiopod genera in various ecological habitats I–V (palaeoenvironmental categories as indicated in the text and also in Fig. 1); TG represents number of total genera

We also analysed the endemism of brachiopod genera in the occurrence network. The network connectedness were calculated by the three measures of bipartite density (BC), network graphic density (GD), and modularity (MD), and suggest a change of biogeographic structure by increased endemism and accumulation after the biotic crisis. Our analysis reveals a decrease in bipartite density from 0.283 in the late Frasnian to 0.241 in the middle Famennian, in accordance with the trend of graphic density from 0.072 to 0.056. Correspondingly, with a modularity rise from 0.245 to 0.335, increased network clustering is revealed after the biotic crisis.

Moreover, frequency analysis based on occurrences data (Fig. 5) suggests the probability distribution of all taxa before and after the biotic crisis. In the diagram, frequencies of brachiopod occurrences occurring in the given range/group of values are depicted and subdivided. It shows that low frequency items, which with large geographic distribution among localities, had largely decreased in the survival and recovery intervals. Whereas the high frequency items, which contribute more to the diversity but with narrow geographic distribution among localities, have increased as shown in Fig. 5b, c. That is to say that most of the widespread taxa before the mass extinction showed a dramatic decrease in their distribution during the crisis, often being confined to a single facies or even a single locality.
Fig. 5

Diagram depicting the distributional frequency of brachiopod genera before and after the biotic crisis in South China (interval A, upper Frasnian (a); interval B, lower Famennian (b); interval C, middle Famennian (c)). Vertical axis represents percentage. Horizontal axis represents distribution of occurrences of brachiopod genera

Implications for brachiopod faunal turnover across the F-F biotic crisis

The main taxonomic loss have been depicted by a decrease of the total number of brachiopod genera in South China, as have been estimated approximately 60% generic extinction rate by previous studies, with pentamerides and atrypides dis-appearing during the biotic crisis (Chen and Ma 2004; Chen 2006). Here, we document a sharp decrease of brachiopod genera from the 15 sections immediately after the biotic crisis and the progressive recovery in the subsequent period. More importantly, our study on brachiopod faunas from different sedimentary facies reveals varied patterns of brachiopod distributions and communication through the biotic crisis.

The pre-extinction brachiopods referred to the Tentico-spirifer tenticulum-Cyrtospirifer Assemblage (Ma and Zong 2010; Hou et al. 2017) contain more widespread taxa in the upper Frasnian as suggested by the higher proportion of low frequency taxa which have wide geographic ranges. The proportion, however, largely decreased after the biotic crisis, with only thirteen genera are recorded in Shaodong, and rare genera from other three localities (see data given in Fig. 4a). In addition, the ecological prominence due to habitat loss for post-extinction faunas are emphasised in most the habitats of South China from littoral clastic, open platform, peritidal to peri-platform slope facies, with decrease of generic connections among localities in the brachiopod occurrence network. It readily suggests that refugees during the survival interval were probably located in the deeper water zones, especially in the inter-platform basin, but also the peritidal zone and the open carbonate platform.

The post-extinction brachiopods represented by the Yunnanella-Nayunnella Assemblage (which could be traced in the Palmatolepis triangularis conodont zone, and exploded mainly in the Pal. crepida to Pal. marginifera zones) has been correlated to the brachiopod recovery, measured by the rapid increase in generic diversity in previous studies (Chen and Ma 2004; Ma and Zong 2010; Hou et al. 2017). Our data for the middle Famennian supports the proposal of a rapid brachiopod recovery in South China (see data given in Fig. 4). Nevertheless, significant biogeographic variations and distributions have been emphasised by the change of proportion of narrow-distributed taxa and widespread taxa after the biotic crisis. The post-extinction faunas are composed of a large proportion of high frequency items (which are rarely found due to restricted geographic distribution), since the proportion of low frequency taxa decreased notably in the recovery interval. This may be partly owing to early diversification of a large number of new taxa that colonised mainly in the open and restricted carbonate platforms, and also in the littoral clastic, inter-platform basin, and peritidal facies.

It is much more interesting when comparing the Chinese data with those outside South China. The Frasnian-Famennian biotic crisis has been debated in recent years for less taxonomic severity of biodiversity loss in view of low origination and speciation rates (e.g. Bambach et al. 2004; Stanley 2007). Stigall (2010, 2012) suggested that brachiopod speciation rate was reduced in the period in which widespread taxa prevailed, as widespread taxa have more probability to disperse and invade between basins, and consequently contributed more to the diversity of post-extinction faunas.

In South China, high frequency taxa which had a narrow geographic distribution among localities (see Fig. 5) had largely increased after the biotic crisis. The result of frequency analysis of brachiopod faunas in South China shows that 25.9 and 37% of the genera confined within a single locality or only a single depositional facies respectively in the upper Frasnian, compared with 34.6 and 53.8% in the middle Famennian. This suggests that generic endemism increased in South China, accompanied with many widespread taxa narrowing in their distribution and surviving only within one or two refuges during the mass extinction. This has been observed both in generic richness and ecological preference in inter-platform basin of South China. Most of the survivors, such as Schuchertella and Cyrtospirifer which were listed in 1 or 2 localities in the lower Famennian and 3 or 4 localities in the middle Famennian, were cosmopolitan prior to the mass extinction. After then, as shown by the frequency analysis, the proportion of high frequency taxa went up and experienced a rapid increase in diversity in restricted zones of carbonate platform, which would benefit from the ecological vacancy after the biotic crisis, and colonised progressively in both shallow and deeper facies, which have largely contributed to the brachiopod recovery in the middle Famennian.

In conclusion, it is clear that the biogeographic structure underwent remarkable changes in the distribution area, range size, connections, and endemism during the Late Devonian biotic crisis, and exerted a significant impact on biodiversity loss and evolutionary dynamics. What is more interesting is that the biogeographic distribution notably changed even at the time of faunal cosmopolitism during the Late Devonian, with collapse of the cosmopolitan atrypid-cyrtospiriferid assemblage and rise of the post-extinction yunnanellid-cyrtospiriferid assemblage. The pre-extinction low frequency genera in South China (widespread taxa or “ecological generalist”) have been removed and overwhelmed during the extinction event. The brachiopod faunas in the survival and recovery intervals contain higher proportions of high frequency taxa (comparable to “ecological specialist”) and experienced a rapid increase in diversity, colonising many ecological vacancies in different depositional settings including littoral clastic, open and restricted platform, peritidal and lagoon facies in the middle Famennian. This pattern apparently can be correlated with sea-level fluctuation across the F-F biotic crisis in South China (Wu et al. 1997; Liao 2001; Ma et al. 2009, 2016; Huang and Gong 2016; Zong et al. 2016) and emphasises the recession of the pre-extinction brachiopod faunas with the habitat loss and restoration of ecological stability towards offshore environments during the post-extinction faunas in the early-middle Famennian.

Notes

Acknowledgements

We are grateful to Dr. B. Huang for the advice on the visualisation technique for bipartite graphs and to Dr. Robert B. Blodgett for the revision on the language. We thank Ulrich Jansen and two other anonymous reviewers for their valuable comments and constructive suggestions.

Funding information

This work is supported by the National Natural Science Foundation of China (No. 41502005, 41772004).

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

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Copyright information

© Senckenberg Gesellschaft für Naturforschung and Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  1. 1.CAS Key Laboratory of Economic Stratigraphy and PalaeogeographyNanjing Institute of Geology and PalaeontologyNanjingChina
  2. 2.Center for Excellence in Life and PaleoenvironmentChinese Academy of SciencesNanjingChina

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