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Evolutionary patterns of Productida (Brachiopoda) morphology during the Permian in South China ZHANG Yang1† & HE WeiHong1,2 1 2

Faculty of Earth Sciences, China University of Geosciences, Wuhan 430074, China； Key Laboratory of Biogeology and Environmental Geology of Ministry of Education, China University of Geosciences, Wuhan 430074, China

The evolutionary patterns of Productida (brachiopod) morphology throughout the Permian show that while the percentage proportion of Productida (brachiopod) with strongly concentric and radial ornamentation declined from the Cisuralian to the Guadalupian, and then increased towards the Changhsingian via Wuchiapingian, the percentage proportion of Productida (brachiopod) with fine concentric and radial ornamentation distinctly increased from the Cisuralian to the Guadalupian, slightly declined towards the Wuchiapingian, and then increased towards the Changhsingian. From the Cisuralian to the Changhsingian, the percentage proportion of brachiopods with spinose ornamentation shows a persistent declining trend. The shell size generally indicates a miniaturization trend at species level during the Wuchiapingian to Changhsingian (including the transitional bed). These evolutionary patterns of brachiopod ornamentation and size are possibly related to the anoxia, food shortage, sea-level fluctuation, and change of substrate in the Permian (including the Permian-Triassic transitional interval) in South China. brachiopod, ornamentation, size, Permian, South China

Animal morphology and structure bear on biology, behaviour and ecology[1] as they evolve through changing environments. Shell ornamentation and size are the key indicators for morphological and environmental evolution. Most studies on brachiopod ornamentation have focused on the Early Paleozoic, Devonian, and Carboni― ferous, or on modern brachiopods[2 15]. Permian bra-

mechanisms and possible causes. This paper describes the evolutionary patterns of Productida (brachiopod) ornamentation and size during the Permian in South China and discusses their possible causes.

1 Materials

[16－20]

chiopod ornamentation has not been well studied . Additionally, most studies on brachiopod miniaturization have focused on post Permian-Triassic extinction brachiopods, with little attention being paid to pre-extinction brachiopods except a few recent reports[20,21]. Understanding the evolutionary patterns of brachiopod ornamentation and size is of great importance, as it not only allows us to understand how brachiopods evolved and how species adjusted to environmental stressors before and even during mass extinctions, but also explains factors associated with extinction

Productida is described as follows: small to gigantic strophomenates with dorsal valves markedly smaller than ventral; corpus cavity in profile commonly concavo-convex to planoconvex; spines tubular and normally covering ventral valves, but may be restricted to Received June 5, 2008; accepted September 9, 2008 doi: 10.1007/s11430-008-0131-3 † Corresponding author (email: [email protected]) Supported by NSFC Innovation Research Group Program (Grant No. 40621002), MOE Innovative Research Team Program (Grant No. IRT0546), the National Natural Science Foundation of China (Grant Nos. 40502001 and 40872008) and the “111” Project (Grant No. B08030)

Sci China Ser D-Earth Sci | Nov. 2008 | vol. 51 | no. 11 | 1589-1600

hingeline, and may be also present on dorsal valves, rarely totally absent; radial ribbing common, rugae less so[22,23]. Productida is the most dominant brachiopod group in the Permian, accounting for 45%―70%[24,25] of all species. The specific number of Productida increased

from the Early Permian to the Late Permian in South China[24,25]. We collected nearly all references from the Cisuralian to the earliest Triassic (Induan) brachiopods taxonomy － of South China[26 55] (Figure 1), made statistics for the

Figure 1 Localities of the Permian Productida species in South China. All data are from refs. [26－55]. Anhui Province: 1, Pingdingshan Section, Chaohu; 2, Majiashan Section, Chaohu; 3, Bailujian Section, Anqing; 4, Dushan Section, Guangde; 5, Longshanwa Section, Guangde; 6, Xiaodushan Section, Guangde; 7, Niutoushan Section, Guangde; 8, Chafeicun Section, Guangde; 9, Wangcunjing Section, Guangde; 10, Shuizhutang Section, Guichi; 11, Renxingshan Section, Guichi; 12, Tangtian Section, Guichi; 13, Qiu’erling Section, Jing; 14, Niuxingshan Section, Tongling; 15, Suishiling Section, Tongling; 16, Shuidong Section, Xuancheng. Jiangsu Province: 17, Zhengpanshan Section, Nanjing; 18, Guanshan Section, Nanjing; 19, Yashan Section, Nanjing; 20, Hushan Section, Nanjing; 21, Xin’an Section, Wuxi; 22, Baiyangtang Section, Yixing; 23, Baoqing Section, Changxing; 24, Meishan Section, Changxing; 25, Fengjing Section, Changxing; 26, Xinhuai Section, Changxing; Zhejiang Province: 27, Huangzhishan Section, Wuxing; 28, Hejiashan Section, Jiangshan; Fujian Province: 29, Yanshizhen Section, Longyan; 30, Yongding Section, Yongding. Jiangxi Province: 31, Chetouduan Section, Yudou; 32, Ji’an Section, Ji’an; 33, Xiafang Section, Anfu; 34, Shixingshan Section, Anfu. Hubei Province: 35, Shatian Section, Huangshi; 36, Guanyinshan Section, Puqi; 37, Mao’ershan Section, Songzi; 38, Sanxikou Section, Songzi; 39, Jiaxinggou Section, Zigui; 40, Xintan Section, Zigui; 41, Guangjiayan Section, Zigui; 42, Daxiakou Section, Xingshan; 43, Xiaoxingchuan Section, Yunxi. Hunan Province: 44, Sansheng Section, Shimen; 45, Muyupu Section, Cili; 46, Rencunping Section, Sangzhi; 47, Nandouxi County, Sangzhi; 48, Yunanxi Section, Sangzhi; 49, Taozixi Section, Yongshun; 50, Leiyang Section, Leiyang; 51, Tongmuqiao Section, Chen; 52, Sanhe Section, Chen; 53, Dapaichong Section, Chen; 54, Yuanjia Section, Jiahe; 55, Xiaoyuanchong Section, Jiahe; 56, Meitian Section, Yizhang. Guangdong Province: 57, Gedingzhai Section, Renhua; 58, Meigang Section, Mei; 59, Wenfu Section, Mei; 60, Baokeng Section, Mei; 61, Chashan Section, Qvjiang; 62, Shuangshan Section, Yangshan; 63, Liyutian Section, Lian; 64, Sankoujiang Section, Lian; 65, Shuizhutang Section, Lian; 66, Damaikong Section, Lian; 67, Bao’an Section, Lian; 68, Yatian Section, Liannan; 69, Ma’an Section, Liannan. Guangxi Province: 70, Dongpan Section, Nanning; 71, Matan Section, Laibin; 72, Paoshui Section, Laibin; 73, Penglaitan Section, Laibin; 74, Heshan Section, Laibin; 75, Wuguiling Section, Laibin; 76, Longlin Section, Longlin. Yunnan Province: 77, Xiaobaihu Section, Luliang; 78, Qingyun Section. Guizhou Province: 79, Yangchang Section, Pan; 80, Huoshaopu Section, Pan; 81, Zhongying Section, Qinglong; 82, Langdai Section, Liuzhi; 83, Zhichang Section, Liuzhi; 84, Lanba Section, Shuicheng; 85, Zhongling Section, Nayong; 86, Damaochang Section, Zhijin; 87, Pingqiao Section, Zhijin; 88, Yanbeihou Section, Zhijin; 89, A’gong Section, Zhijin; 90, Zhuzang Section, Zhijin; 91, Liuchang Section, Zhijin; 92, Bulang Section, Puding; 93, Jiaozishan Section, Puding; 94, Liuchang Section, Qingzhen; 95, Wenjiangsi Section, Guiding; 96, Shizipu Section, Zunyi; 97, Tongzi Section, Tongzi. Chongqing City: 100, Yutangjiao Section, Niejiang; 101, Jijiang Section, Niejiang; 102, Banzhuyuan Section, Neijiang; 103, Taiping Section; 104, Yutianbao Section; 105, Beifengjing Section; 106, Liangfengya Section; 107, Beipei Section; 108, Yanjingxi Section, Hechuan; 109, Liziya Section; 110, Mulongdong Section. Sichuan Province: 98, Daijiagou Section, Luzhou; 99, Chuanyan Section, Luzhou; 111, Guang’an Section, Guang’an; 112, Shangsi Section, Guangyuan; 113, Xindianzi Section, Guangyuan. Shaanxi Province: 114, Xikou Section, Zhen’an. 1590

ZHANG Yang et al. Sci China Ser D-Earth Sci | Nov. 2008 | vol. 51 | no. 11 | 1589-1600

analysis of shell ornamentation and size only based on those references where Productida specimens were illustrated, and selected some species which were found to range at least in two stages to make statistics for the size variation. In order to study the morphological variation of the Permian Productida of South China, we classified those species from the selected literatures (see above) into four categories in referring to the work of Leighton[56]. Class 1, weak ornamentation: ornamentation is either absent (smooth shell) except for growth lines, or capillate; Class 2, costate, rugose ornamentation; Class 3, strongly costate, rugose, or plicate ornamentation; and Class 4, spinose ornamentation anterior to the hingeline (see Figure 2). Chonetidines posses a row of posteriorly directed spines near their ventral hingelines, but the surface of the shell is not spinose. Consequently, no Chonetidines was involved in Class four. Additionally, length and width of brachiopods species were measured based on the selected refs. [26－55]. The chronostratigraphic assignment used for brachiopod statistics in this paper mainly follows the － works of Jin et al.[63 65], Gradstein et al.[66], and Ogg et al.[67] (Table 1). Because of the lack of biostratigraphic data in several papers[28,43,44], the chronostratigraphic assignment used for brachiopod statistics from these papers follows the roughly corresponding relationship between the chronostratigraphic and lithostratigraphic units of the Permian in South China. For example, The Changxing Formation, Yanshi Formation and Wangpanli

Formation are roughly corresponding to the age of Changhsingian, and the Wujiaping Formation, Shuizhutang Formation and Yundoutan Formation are roughly corresponding to the age of Wuchiapingian. We select the Cisuralian Epoch, Guadalupian Epoch, WuchiapinTable 1

Permian chronostratigraphic framework[63

―67]

Figure 2 Examples of four ornamentation classes: Class 1, smooth or weakly ornamented ((a)[57]); Class 2, costate ((b)[58] and (c)[59]); Class 3, plicate or rugose ((d)[60] and (e)[61]); Class 4, spinose ((f)[62]). Width, the maximum width of shell; Length, the maximum length of shell. ZHANG Yang et al. Sci China Ser D-Earth Sci | Nov. 2008 | vol. 51 | no. 11 | 1589-1600

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gian Stage, Changhsingian Stage and Induan Stage as the time units for the statistics of brachiopod morphology. The Cisuralian, Guadalupian, Wuchiapingian, and Changhsingian are differently-ranked time units. These differently-ranked time units are used because the frequency of occurrence of some species in the Cisuralian and Guadalupian epochs is too low for statistical significance at stage level. In addition, the frequency of occurrence of some species in each stage of the Lopingian Epoch is relatively higher and therefore acceptable for detailed statistics at the stage level. In some papers dealing with species described from the Permian-Triassic interval[29,31,48,54,55] (i.e., transitional bed[68]), it is unclear whether the species were from the top of the Changhsingian or the bottom of the Induan and these species have been assigned to the Permian-Triassic transition in this paper. The classification of brachiopods referred to in this study largely follows Williams et al.[23].

2 Results Based on integrated statistics for shell ornamentation of the above-mentioned four categories, the percentage proportion of Class 1 is very low and does not represent conspicuous changes during the Permian (Figure 3). The percentage proportion of Class 2 distinctly increased from the Cisuralian to the Guadalupian, slightly declined towards the Wuchiapingian, and then increased towards the Changhsingian (Figure 3). The percentage propor-

tion of Class 3 declined from the Cisuralian to the Guadalupian, and then attained an increasing trend from the Wuchiapingian to the Changhsingian (Figure 3). The percentage proportion of Class 4 exhibited a distinctly declining trend during the Permian (Figure 3). Fifteen species’ geometric size (length and width) from the Productida brachiopods are analysed and illustrated (Figures 4 and 5). Among these species, thirteen show a distinct decrease in size from the Wuchiapingian to the Changhsingian (Figure 4 (a)－(k), (n), (o); Figure 5 (a)－(k), (n), (o)). Only two species increased in size (lengths) during that time (Figure 4 (l) and (m)). In summary, the evolutionary pattern of concentric and radial ornamentation of brachiopods could be described as follows: during the first phase, the percentage proportion of Class 3 declined from the Cisuralian to the Guadalupian, and then increased towards the Wuchiapingian; the percentage proportion of Class 2 shows an opposite trend from the Cisuralian to the Wuchiapingian compared with the changes of Class 3. During the second phase, the percentage proportion of classes 2 and 3 shows an increasing trend towards the Changhsingian via Wuchiapingian. The evolutionary pattern of spinose ornamentation of brachiopods is characterized by that the percentage proportion of Class 4 shows a persistent declining trend during the Permian. The size generally shows a miniaturization trend from the Wuchiapingian to the Changhsingian, except the lengths of a few species showed an increasing trend.

Figure 3 Relationship among species diversity, number of species with different classes of ornamentations, percentage proportion of species with different classes of ornamentations and sea-level changes[69] of South China. 1592


Figure 4 Shell lengths of brachiopods species (Productida) from South China during the Permian. Vertical axis represents the shell length (mm); abscissa represents the geological age. The midpoint, upper end, and lower end of the vertical line respectively represent the average length, the maximum length, and the minimum length of each species. Black solid dots are based on single datum points. A, Cisuralian; B, Guadalupian; C, Wuchiapingian; D, Changhsingian; E, Induan. The erect and short lines between D and E represent the latest Changhsingian or the Permian-Triassic transition. The mass extinction occurred at the end-Changhsingian (between D and E).


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Figure 5 Shell widths of brachiopods species (Productida) from South China during the Permian. Vertical axis represents the shell width (mm); abscissa represents the geological age. The midpoint, upper end, and lower end of the vertical line respectively represent the average width, the maximum width, and the minimum width of each species. Black solid dots are based on single datum points. A, Cisuralian; B, Guadalupian; C, Wuchiapingian; D, Changhsingian; E, Induan. The erect and short lines between D and E represent the latest Changhsingian or the Permian-Triassic transition. The mass extinction occurred at the end-Changhsingian (between D and E).

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3 Discussion 3.1 Ornamentation As mentioned above, the evolutionary pattern of concentric and radial ornamentation of brachiopods is very complex in the Permian. The percentage proportion of Class 3 declined from the Cisuralian to the Guadalupian, and then increased towards the Wuchiapingian; the percentage proportion of Class 2 shows an opposite trend compared with the changes of Class 3. In short, the concentric and radial ornamentation of Productida became weaker from the Cisuralian to the Guadalupian, and then stronger towards the Wuchiapingian. The acquisition of stronger concentric and radial ornamentation needs more nutrition (including oxygen and available food), and presumably the evolutionary pattern of concentric and radial ornamentation of brachiopods may be a result of the fluctuation of nutrition. This fluctuation is possibly caused by sea-level changes. This presumption is supported by the study of sequence stratigraphy and sea-level changes throughout the Permian. The sea level rose from the Cisuralian to the Guadalupian, fell toward the Wuchiapingian, and then rose again toward the － Changhsingian[69 71] (Figure 3). With the rise of sea level, the water became deeper, the available food for filter-feeder brachiopods was reduced, and the shells of brachiopods thinned and their ornamentation weakened; meanwhile, with the rise of sea level, the hydrostatic pressure increased, the thinning of shell and the weakening of ornamentation made it easier for brachiopods to open their shells to get nutrition. These are consistent with the fact that the ornamentation of brachiopods lived in the shallow-water carbonate setting is stronger than the ornamentation of brachiopods lived in the deep-water siliceous basin in South China. Therefore, the evolutionary pattern of concentric and radial ornamentation of Productida reflects the sea-level changes from the Cisuralian to the Wuchiapingian. From the Wuchiapingian to the Changhsingian, the percentage proportion of classes 2 and 3 shows an increasing trend and the species-level diversity of Productida brachiopods is also increased. The statistics show that the genera with higher species-level diversity mainly include Leptodus, Oldhamina, Richthofenia, Edriosteges, Haydenella, Spinomarginifera, Cathaysia, Neochonetes, Paryphella, and Tethyochonetes during the Wuchiapingian to Changhsingian, and thus the increase

of the percentage proportion of classes two and three is related to the diversification of these genera. Leptodus, Oldhamina, and Richthofenia that have a bigger shell size and abundant body spines were commonly found in a substrata of carbonate or in a setting of carbonate platform[30]. Cathaysia, Neochonetes, Paryphella, and Tethyochonetes were commonly found in a setting where siliceous materials deposited[29,30], and these genera have common characters such as abundant radially- and densely-arranged pseudopunctates (the shells of Productida brachiopods are filled with pseudopunctates, but the pseudopunctates of these genera are more abundant and more densely arranged), spines developed along their hingelines, and thin shells. When there is a shortage of nutrition in a setting, denser pseudopunctates could increase the area of mantle of brachiopods and the mantle has the function of respiration[4,72], so these brachiopods can make use of nutrition more effectively. Besides, the species with thinner shells consume less food than those species with thicker shells. Additionally, the species with spines along their hinglines can hug to floating wood or sea algae by spines[18,72], improving brachiopods mobility for obtaining food in shortage of nutrition. Therefore, the diversification of Cathaysia, Neochonetes, Paryphella, and Tethyochonetes is possibly related to the shortage of nutrition. As analyzed above, the increase of percentage proportion of brachiopods with strong concentric and radial ornamentation is related to the change of substrata, i.e., the widespread of platform and reefs from the Wuchiapingian to the Changhsingian[73,74], and the increase of percentage proportion of brachiopods with fine concentric and radial ornamentation indicates the shortage of nutrition in the siliceous basin during the Late Permian. From the Cisuralian to the Changhsingian, the percentage proportion of Productida with spinose ornamentation showed a persistent decline trend. From the Cisuralian to the Guadalupian, this decline was probably － related to the sea-level rise of Guadalupian[69 71]. In a setting of shallow water the spinose ornamentation of brachiopods became strong for anchoring themselves to the substrata because of the higher energy. Reversely, if the sea level rises, the spinose ornamentation of brachiopods would become weaker. From the Wuchiapingian to the Changhsingian, the decline was possibly resulted from the diversification of Cathaysia, Neochonetes, Paryphella, and Tethyochonetes but not from the


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decline of diversity of Class 4. It is because that when we classified species (Productida) into four categories, Class 4 (with strong spinose ornamentation) does not include the species with spines along their hingelines. Thus, the increase of percentage proportion of Classes 2 and 3 (including those species with spines along hingelines) surely led to the decline of percentage proportion of Class 4. The phenomenon that Class 4 increased in proportion but its species diversity did not decline towards the Changhsingian also supports this argument (Figure 3). As mentioned above, the decline of the percentage proportion of Class 4 was related to the diversification of Cathaysia, Neochonetes, Paryphella, and Tethyochonetes towards the Changhsingian via Wuchiapingian, and it also indicated the shortage of nutrition in the siliceous basin during the Late Permian. 3.2 Size The statistical results related to species-level size of brachiopods indicate a miniaturization (Lilliput) trend from the Wuchiapingian to the Changhsingian, and even possibly into the Induan. It may be argued that miniaturization is related to anoxia, increased temperature, or － decreased food supply[75 77]. We prefer to consider that anoxia probably caused the miniaturization from the Wuchiapingian to the Changhsingian. Absence of oxyTable 2

gen would directly affect metabolism for the brachiopods, reducing their calcium secretion and inhibiting － their growth[75 77]. The anoxic setting was expanded from the Wuchiapingian to the Changhsingian in South China, as evidenced by the wider distribution of basin-facies black shale in the Changhsingian, compared with the Wuchiapingian[78]. Changhsingian basin-facies black shale contained abundance pyrite and organic materials in South China and indicates an anoxic environment. It possibly resulted from a Changhsingian marine transgression of South China and global atmospheric anoxia[79,80]. Additionally, the miniaturization of most brachiopod species (studied in this paper) did not result from the changes of substrata, because they were found in the same lithofacies (either in limestone or in siliceous-rock facies) or the discovery frequency in limestone facies is similar to that in siliceous-rock facies from the Wuchiapingian to the Changhsingian (Table 2). However, Tethyochonetes wongiana and Tyloplecta yangtzeensis were commonly found in limestone facies in the Wuchiapingina but were very common in siliceous-rock facies in the Changhsingian (Table 2). To sum up, the miniaturization process from the Wuchiapingian to the Changhsingian is mainly resulted from marine transgression and anoxia. The causes of the

The frequency of measurement for each species and lithofacies of different species found (only including the species in Figures 4 and 5)

Species Cathaysia chonetoides Chao

Cisuralian

Compressoproductus compressus Waagen

Guadalupian 4 2 1

Wuchiapingian 8

32

4

2 1 2 1 12 4 21

Edriosteges poyangensis Kayser Edriosteges subplicatilis Frech Haydenella kiangsiensis Kayser Leptodus deminutus Liao Neochonetes zhongyingensis Liao Ogbinia dzhagrensis Sarytcheva Paryphella sulcatifera Liao

16 5 7 2 2 4

Transitional bed

3

Spinomarginifera kueichowensis Huang Spinomarginifera pseudosintanensis Huang Tethyochonetes liaoi Chen Tethyochonetes quadrata Zhan

Changhsingian

8 4 4

Tethyochonetes wongiana Chao 2 Tyloplecta yangtzeensis Chao

1 2 2 10 1 2 4 6 8 3 4

2 5 7 3

18

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Facies siliceous rock limestone siliceous rock limestone siliceous rock limestone limestone limestone limestone limestone limestone siliceous rock limestone siliceous rock limestone limestone limestone siliceous rock limestone siliceous rock limestone siliceous rock limestone

end-Permian or the Permian-Triassic transitional miniaturization are more complex and include a marine regression, decline in available food sources for brachiopods, or increased terrestrial input[21,70,81], probably similar to the causes of the end-Permian brachiopod miniaturization observed in the deep waters of South China[21]. The variations in size of Tethyochonetes quadrata Zhan and Tethyochonetes liaoi Chen did not show a miniaturization trend. Lengths of Tethyochonetes quadrata Zhan and Tethyochonetes liaoi Chen increased although their widths decreased from Late Changhsingian to transitional bed (latest Changhsingian or earliest Triassic), leading to their increased ratio of length and width. This change may be because that in a stressed ecologic pressure, these brachiopods were forced to change their life mode, from benthic to pseudoplankton. These species were forced to hug to the floating wood or sea algae by hingeline spines for a long time and their beak preferred to face upcurrent[5]. If the ratio of length and width of these species increases, then it would make the brachiopods apt to be line-shaped and thus easier to overcome the resistance of flowing water. This change would improve brachiopods’ mobility to capture food during shortage of nutrition, especially during or after the Permian-Triassic crisis. 3.3 Evolutionary patterns of Productida morphology and their implications for mass extinction In deep-water siliceous environment of the Late Permian in South China, available nutrition (including food and oxygen) for brachiopods was in a low level for a long time. In such an environment, commonly-found brachiopods were the ones with weak ornamentation, abundant pseudopunctate, and smaller and thinner shell, and good mobility. The shortage of nutrition kept the brachiopod fauna in a low diversity and caused the miniaturization of brachiopods. With further decline of nutrition, negative carbonate isotope, reduced organic carbon, distinct reduction of Protozoa (radiolarians and foraminifers), and collapse of reef ecosystem at the endPermian[21,73,82,83], most brachiopods suffered miniaturization and began to die out. In shallow-water carbonate environment, available nutrition for brachiopods was relatively abundant. In such an environment, brachiopods with strong ornamentation, larger sizes, and higher diversity were com-

monly found there, such as the diversification of Leptodus, Oldhamina, and Richthofenia. Nevertheless, towards the end-Permian or the Permian-Triassic transition, this brachiopod fauna was replaced by the fauna with weak ornamentation, abundant pseudopunctate, smaller and thinner shell, and good mobility[29]. The end-Permian or the Permian-Triassic transitional brachiopod fauna from the shallow-water carbonate environment is very similar to the Late Permian fauna from the deep-water siliceous environment in ornamentation, size and even species composition. Therefore, towards the end Permian or the Permian-Triassic transition, the decline of nutrition in the carbonate environment may have caused the faunal replacement and even may ultimately have caused the extinction of brachiopods. Additionally, the sea level rose from the Wuchiapingian towards the Changhsingian, and fell in the latest Changhsingian, and then rose again towards the earliest Triassic in South China[21,70,71]. The sea-level fluctuation is one of causes for the formation of evolutionary patterns of Productida morphology, and also one of causes for mass extinction of brachiopods. The regression opened up new shallow-water habitats, allowing brachiopod abundance to increase, which also resulted in an increase in population density and an increase in competition among brachiopods for available habitats and food resources[21,84]. The subsequent transgression further led to the reduction of nutrition for brachiopods. Shortage of food resources would have limited the metabolism of the brachiopods, reducing the secretion of calcareous materials, slowing their growth, weakening ornamentation, and reducing the size. The shortage of food resources would also lead to the mass extinction of some brachiopods that need more nutrition for their metabolism. The change of substrata is also possibly responsible for the forming of evolutionary patterns of brachiopods’ morphology from the Wuchiapingian to the Changhsingian. However, in the intervals where most brachiopods species disappeared, the substrata did not show a distinct change, such as at the Chaoxian section of Anhui, the Sangzhi section of Hunan and the Liangfengya section of Chongqing. The change of substrata therefore is possibly not closely related to the mass extinction of brachiopods. In summary, the shortage of food and oxygen, sea-level fluctuation and the change of substrata are


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causes for the forming of evolutionary patterns of brachiopods’ morphology from the Permian to the Permian-Triassic transition, and the shortage of food or oxygen and sea-level fluctuation possibly led to the end-Permian mass extinction of brachiopods in South

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1600


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