European Journal of Taxonomy 19: 1-34

http://dx.doi.org/10.5852/ejt.2012.19

This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2012 • M.B. Forel

Research article

Ostracods (Crustacea) associated with microbialites across

the Permian-Triassic boundary in Dajiang

(Guizhou Province, South China)

Marie-Beatrice FOREL

State Key Laboratory of Geological Process and Mineral Resources, China University of Geosciences,

No. 388, Lumo Road, Wuhan 430074, People’s Republic of China.

E-mail: [email protected]

Abstract. 26 samples were processed for a taxonomic study of ostracods from the Upper Permian

(Changhsingian) - Lower Triassic (Griesbachian) interval of the Dajiang section, Guizhou Province,

South China. 112 species belonging to 27 genera are recognized. Five new species are described:

Acratia candy ae sp. nov, Bairdia adelineae sp. nov., Bair did 1 , huberti sp. nov., Bairdia jeromei sp. nov.,

Orthobairdia jeanlouisi sp. nov. The unexpected survival faunas associated with microbial formations

in the aftermath of the end-Permian extinction are documented for the first time. Ostracod biodiversity

variations and palaeo-environmental modifications associated with microbial growth through the

Permian-Triassic boundary (PTB) are discussed.

Keywords. Ostracods, Permian-Triassic, Microbialites, Dajiang, South China.

Forel M.B. 2012. Ostracods (Crustacea) associated with microbialites across the Permian-Triassic boundary in

Dajiang (Guizhou Province, South China). European Journal of Taxonomy 18: 1-34. http://dx.doi.org/10.5852/

eit.2012.19

Introduction

The end-Permian extinction, about 252 My, decimated 80 to 96% of species and 52% of families in the

marine realm (e.g. Seplcoski 1984; Erwin 1993; Benton & Twitchett 2003). Several potential geological

triggers have been identified: bolide impact, oceanic anoxia and euxinia and flood basalt volcanism.

It is now accepted that ocean anoxia was widespread during the latest Permian (Changhsingian), its

prevalence increased near the time of the main extinction and continued during the Early Triassic.

Anoxic and sulfidic waters commonly also extended into shallow-marine enviro nm ents. The eruption

of the Siberian Traps is now considered as the primary trigger of extinction. Global warming, ocean

acidification, and possible destruction of atmospheric ozone were engendered by the release of volcanic

CQ, and volatilized sedimentary organic carbon and evaporite minerals. Enhanced weathering and

nutrient runoff increased pre-existing ocean anoxia (see Payne & Clapham 2012 and references therein

for details). The extinction marks a major transition in marine ecosystems: the Upper Permian benthic

shelly communities from shallow marine settings were replaced by widespread microbial communities.

They are abundant in low-latitude shallow-marine carbonate shelves in central Tethyan continents

where they occupied similar environments to Upper Permian reefs but extended into deeper waters

European Journal of Taxonomy 19: 1-34(2012)

Fig. i. Location of the Dajiang section in the southern Guizhou Province, South China.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

(Kershaw et al. 2007, 2012). Current evidence regarding oxygen levels associated with microbialites

growth is conflicting : (z) evidence of low-oxygen conditions (Bond & Wignall 2010; Liao et al. 2010;

Chen et al. 2011), (z'z) abundant benthic shelly faunas dominated by ostracods, and occasionally some

rare micro-gastropods, microbrachiopods, foraminifers and conodonts (e.g. Crasquin-Soleau & Kershaw

2005; Forel et al. 2009; Song et al. 2009; Yang et al. 2011; Forel et al. in press). A two-steps oxygenation

of the surrounding waters by cyanobacterial activity will be proposed as a possible mechanism for this

unexpected survival (Forel in press).

74 to 100% of ostracods became extinct through the Permian-Triassic boundary (PTB) and very little is

known about their presence and characteristics in the interval following the extinction event. In the lower

Griesbachian neritic enviromnents, they are only found in association with microbialites. The faunas

described in the present article are of primary importance because they exemplify a survival phenomenon

in a refuge of microbial origin documented from several localities worldwide (Forel in press; Forel et al.

in press). This paper describes the upper Changhsingian and lower Griesbachian ostracod faunas of the

Dajiang section (Wujiaping and Daye Formations respectively), Guizhou Province, South China. 112

species are recorded, including 5 new species, belonging to 27 genera.

Material and Methods

The Dajiang section (25 o 33’56”N-106°39’41”E) is located in the southern Guizhou Province, South

China, a few kilometres north of the city of Luodian (Fig. 1). Only a brief description of the lithological

succession in Dajiang is given here: the reader is referred to Forel et al. (2009) and references therein

for further details. At the base of the section are about 7 metres of open shallow marine thick, massive-

bedded skeletal lime-packstone of the Wujiaping Formation (Changhsingian age). They are followed

by about 17 metres of microbialite carbonate layers of the Daye Formation. The conodont Hindeodus

parvus (Kozur & Pjatakova, 1976) occurs throughout the lowermost metre-thick microbialite carbonate,

above the major facies change that marks the greatest loss in Permian fossils (the first occurrence of

H. parvus is the index of the base of the Triassic: Yin et al. 1996; Lehrmann 1999; Lehrmann et al.

2003). The PTB ‘event horizon’ corresponds to the contact between the Wujiaping Formation and the

microbialite (Lehrmann et al. 2003).

Samples spanning the Permian-Triassic event horizon were collected for ostracod analysis and labelled

as 05PAJxx (Fig. 2). Ostracods are determined on the basis of external and internal characters of the

carapace and have to be released from the enclosing matrix. Extraction by means of acid is precluded

because their carapaces are made of calcium carbonate and enclosed in calcareous rocks. Samples were

processed by hot acetolysis technique to disaggregate the dehydrated hard limestones and release the

ostracod shells (Lethiers & Crasquin-Soleau 1988; Crasquin-Soleau et al. 2005). Twenty-six samples

were collected and analysed, only one was barren (05PAJ34; Fig. 2). The faunas in Dajiang show high

intraspecific variability that hindered the clear differentiation between species in many cases. Due to the

poor preservation and availability of specimens, many species are left in open nomenclature and are not

listed in the systematic part (e.g. Bairdia sp. 1 to 36), although all the material of this important interval

is figured. All specimens are stored in the Pierre et Marie Curie University Collections (Paris, France;

collection numbers P6M2917-3150).

All the species belonging to the Bairdioidea (Sars, 1887) superfamily are shown in Figs 3 (genera Acratia

Delo, 1930 and Bairdia McCoy, 1844), 4, 7-9 (genus Bairdia ), 10 (genera Bairdia and Bairdiacypris

Bradfield, 1935), 11 (genera Bairdiacypris , Bythocypris Brady, 1880, Fabalicypris Brady, 1880,

Liuzhinia Zheng, 1976 and Orthobairdia Sohn, 1960) and 13 (genera Petasobairdia Chen, 1982,

Spinocypris Kozur, 1971, Kempfina Crasquin, 2010, Silenites Coryell & Booth, 1933 and Microcheili-

nella Geis, 1933). Species belonging to the Cypridoidea Baird, 1845 are gathered in Figs 13 and 14

(genus Paracypris Sars, 1866). Cytheroidea Baird, 1850 are shown in Figs 14 (genera Monoceratina

European Journal of Taxonomy 19: 1-34 (2012)

t n

c/3

-5

c/3

't—>

• i-H

'e?

&3d

species distribution through the section.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Fig. 3. Ostracods from the Dajiang section, South China. — A-D. Acratia candyae sp. nov. A. Holotype,

carapace, right lateral view, P6M2917. B. Carapace, dorsal view, P6M2918. C. Paratype, carapace,

right lateral view, P6M2919. D. Carapace, right lateral view, P6M2920. — E-F. Acratia subfusiformis

Wang, 1978. E. Carapace, right lateral view, P6M2921. F. Carapace, right lateral view, P6M2922. —

G. Acratia ? sp. 1, carapace, right lateral view, P6M2923. — H. Acratia ? sp. 2, carapace, right lateral

view, P6M2924. — I. Acratia ? sp. 3, carapace, right lateral view, P6M2925. — J-K. Acratia sp. 4. J.

Carapace, left lateral view, P6M2926. K. Carapace, right lateral view, P6M2927. — L. Acratia sp. 5,

carapace, right lateral view, P6M2928. — M. Acratiidae indet., carapace, right lateral view, P6M2929.

— N-Q. Bairdia adelineae sp. nov. N. Holotype, carapace, right lateral view, P6M2930. O. Carapace,

dorsal view, P6M2931. P. paratype, carapace, right lateral view, P6M2932. Q. Carapace, right lateral

view, P6M2933. — R-V. Bairdia gaelleae Crasquin, 2010. R. Carapace, dorsal view, P6M2934. S.

Carapace, right lateral view, P6M2935. T. Carapace, left lateral view, P6M2936. U. Carapace, right

lateral view, P6M2937. V. Carapace, right lateral view, P6M2938. - Scale = 100 pm.

European Journal of Taxonomy 19 : 1-34 ( 2012 )

Roth, 1928, Basslerella Kellett, 1935 and Callicythere Wei, 1981) and 15 (genus Callicythere).

Cavellinidae Egorov, 1950 (genus Sulcella Coryell & Sample, 1932), Polycopidae Sars, 1866 (genus

Polycope Sars, 1866), Aparchitidae Jones, 1901 (genus Cyathus Roth & Skinner, 1930), Kirlcbyoidea

Ulrich & Bassler, 1906 (genera Amphissites Girty, 1910, Kirkbya Jones, 1859 and Shleesha Sohn, 1961),

Kloedenellidae Ulirch & Bassler, 1908 (genus Oliganisus Geis, 1932) and Paraparchitidae Scott, 1959

(genera Paraparchites Ulrich & Bassler, 1906 and Shemonaella Sohn, 1971) are shown in Fig. 15.

Abbreviations used in the text

ADB

AVB

PDB

PVB

Um ax

Lmax

Wmax

right valve

left valve

anterior border

posterior border

dorsal border

ventral border

antero-dorsal border

antero-ventral border

postero-dorsal border

postero-ventral border

height

maximal height

length

maximal length

width

maximal width

Results

Taxonomic descriptions

Class Ostracoda Latreille, 1802

Order Podocopida Muller, 1894

Sub-order Podocopina Sars, 1866

Superfamily Bairdioidea Sars, 1888

Family Acratiidae Griindel, 1962

Genus Acratia Delo, 1930

Acratia candyae sp. nov.

Fig. 3A-D

Diagnosis

Species of Acratia similar to Microcheilinella in dorsal view, with FV larger than RV and AB broadly

rounded in 1/4 circle.

Etymology

Personal dedication to Candy Boiteux.

Material examined

Holotype

One carapace (Fig. 3A), Dajiang section, sample 05PAJ28, collection number P6M2917.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Paratype

One carapace (Fig. 3C), Dajiang section, sample 05PAJ17, collection number P6M2919.

Other material

4 carapaces, several fragments. The species is known from its type locality only.

Type locality

Wujiaping Formation (Samples 05PAJ17, 25, 28, 29), Dajiang section (25°33 , 56 ,, N-106°39 , 4r , E),

Guizhou Province, South China, Changhsingian, Late Permian.

Measurements

L = 323 - 464 pm; H = 140 - 200 pm; H/L = 0.43 - 0.44.

Description

Carapace elongated and subovoid in lateral view; surface smooth; carapace slightly preplete to amplete;

LV strongly overlaps RV all around the carapace; Lmax below mid-H; dorsum long and slightly rounded

in both valves, angles between PDB and DB not distinct; rounded angle between DB and ADB (-140°);

PDB short and steep; DB long and gently bent backward; ADB short, steep and convex; ventral margin

flat and straight at both valves; AVB, VB, PVB not distinct; AVB straight and slightly bent toward AB; AB

broadly rounded in 1/4 circle; acratian beak clear but not pronounced, located at VB; PB sharpen with quite

large radius of curvature for the genus, maximum slightly below mid-H; angle between AVB-AB (>90°)

larger than angle between PVB-PB (<90°); dorsal view similar to the one of the genus Microcheilinella

with carapace thick and Wmax located in the posterior 1/3 of L, hinge line slightly incised.

Through the ontogeny, the carapace becomes more stocky, more amplete at RV, with DB shorter and

more rounded.

Remarks

The strong overlap of LV on RV all around the carapace and characteristics of dorsal view are reminiscent

of the genus Microcheilinella. However the acratian beak clearly precludes Microcheilinella attribu¬

tion and indicates the genus Acratia. Acratia candyae sp. nov. is very close to Acratia sp.2 sensu

Crasquin, 2008 from the Late Permian of Dolomites (Crasquin et al. 2008). However, Acratia sp.2

sensu Crasquin, 2008 has longer ADB and well expressed angles between PDB, DB and ADB.

Acratia subfusiformis Wang, 1978

Figs 3E-F

Acratia subfusiformis Wang, 1978: 294-295, pi.4, figs 3-4.

Acratia subfusiformis - Shi & Chen 1987: 49, pi. 11, figs 19-22, pi. 12, figs 1-2 — Shi & Chen 2002: 82,

pi.20, figs 1-9. — Crasquin et al. 2010: 357, figs 23I-L.

? Acratia subfusiformis - Shi & Chen 2002: 82, pi. 20, figs 12-13.

Not Acratia subfusiformis - Shi & Chen 2002: 82, pi. 20, figs 10-11.

Localities

- Wujiaping Formation (samples 05PAJ22, 26), Dajiang section (25 o 33 , 56 ,, N-106°39 , 41 ,, E), Guizhou

Province, South China, Changhsingian, Late Permian.

- Longtan Formation, Northern Guizhou and Southern Yunnan Provinces, South China, Wuchiapingian,

Late Permian (Wang 1978).

European Journal of Taxonomy 19 : 1-34 ( 2012 )

- Baoqing Member, Changxing Formation, Meishan section, Zhejiang Province, South China,

Changhsingian, Late Permian (Shi & Chen 1987).

- Matan and Pingding sections, Guangxi Province, South China Wuchiapingian and Changhsingian,

Late Permian (Shi & Chen 2002).

- Baoqing and Meishan Members, Changxing Formation, Meishan section, Zhejiang Province, South

China, Changhsingian, Late Permian (Crasquin et al. 2010).

Family Bairdiidae Sars, 1887

Genus Bairdia McCoy, 1844

Bairdia adelineae sp. nov.

Fig. 3N-Q

Diagnosis

Stocky species of Bairdia with strong dorsal overlap, laterally flattened plateau-like DB, AB and PB

with large radius of curvature.

Etymology

Personal dedication to Adeline Bienvenu.

Material examined

Holotype

One carapace (Fig. 3N), sample 05PAJ42, collection number P6M2930.

Paratype

One carapace (Fig. 3P), sample 05PAJ42, collection number P6M2932.

Other material

4 carapaces, several fragments. The species is known from its type locality only.

Type locality

Daye Formation (Sample 05PAJ42), Dajiang section (25 o 33 , 56 ,, N-106°39 , 41 ,, E), Guizhou Province,

South China, Griesbachian, Early Triassic.

Measurements

L = 513 - 655 pm; H = 292 - 347 pm; H/L = 0.53 - 0.58.

Description

Carapace elongated and massive, subovoid in lateral view; surface smooth; carapace amplete to preplete;

LV overlapping RV all around the carapace, dorsal overlap stronger than ventral one; dorsum subdivided

into 3 distinct parts at both valves; PDB straight to slightly convex; DB straight at both valves, flattened

laterally at RV; ADB straight to slightly concave at anterior extremity; DB and ADB longer than PDB;

VB long and slightly concave in median part at RV, straight to slightly concave at LV; AVB broadly

rounded and long (~ 75% of Hmax); PVB straight to convex and short (<50% of Hmax); AB rounded

with large radius of curvature, maximum in the upper 1/3 of H, bairdian beak poorly expressed but

distinct; PB pointed with large radius of curvature, maximum at or slightly below mid-H; in dorsal view,

carapace lenticular and thick with anterior extremity slightly tapered, Wmax located around mid-L,

plateau-like laterally flattened at RV.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Fig. 4. Ostracods from the Dajiang section, South China. —A-F. Bairdia? huberti sp. nov. A. Carapace,

right lateral view, P6M2939. B. Carapace, right lateral view, P6M2940. C. Paratype, carapace, right

lateral view, P6M2941. D. Holotype, carapace, right lateral view, P6M2942. E. Carapace, right lateral

view, P6M2943. F. Carapace, right lateral view, P6M2944. — G-P. Bairdia jeromei sp. nov. G.

Holotype, carapace, right lateral view, P6M2945. H. Carapace, dorsal view, P6M2946. I. Carapace,

right lateral view, P6M2947. J. Paratype, carapace, right lateral view, P6M2948. K. Carapace, right

lateral view, P6M2949. L. Carapace, right lateral view, P6M2950. M. Carapace, right lateral view,

P6M2951. N. Carapace, right lateral view, P6M2952. O. Carapace, right lateral view, P6M2953. P.

Carapace, right lateral view, P6M2954. — Q-T. Bairdia limatusformis Forel, 2010. Q. Carapace,

right lateral view, P6M2955. R. Carapace, left lateral view, P6M2956. S. Carapace, sub-dorsal view,

P6M2957. T. Carapace, right lateral view, P6M2958. — U-V. Bairdia sp. 1. U. Carapace, right lateral

view, P6M2959. V. Carapace, right lateral view, P6M2960. — W-X. Bairdia sp. 2. W. Carapace, dorsal

view, P6M2961. X. Carapace, right lateral view, P6M2962. - Scale =100 pm.

European Journal of Taxonomy 19 : 1-34 ( 2012 )

Remarks

This new species can be related to Bairdia paussi Crasquin, 2010 from the Late Permian of the Meishan

section, Zhejiang Province, South China (Crasquin et al. 2010). However Bairdia adelineae sp. nov. is

more elongated, with stronger overlap and posterior maximum of convexity located closer to VB. It can

also be compared to Orthobairdia texana (Harlton, 1927) sensu Shi & Chen, 2002 from the Permian of

Heshan and Yishan, Guangxi Province, South China (Shi & Chen 2002). However, the bairdioid dorsal

view precludes generic attribution to Orthobairdia. Bairdia adelineae sp.nov is also close to Bairdia

chasae Kellett, 1934 sensu Wang, 1978 from the Late Permian of the Western Guizhou and Northeastern

Yunnan Provinces, South China (Wang 1978). However the new species has a less pointed PB, a thinner

ventral overlap and the dorsal view clearly shows a bairdioid shape. It is noteworthy that Bairdia chasae

Kellett, 1935 sensu Wang, 1978 seems wrongly attributed to the genus Bairdia: the dorsal view shows

a typical orthobairdian shape.

Bairdia gaelleae Crasquin, 2010

Fig. 3R-V

Bairdia gaelleae Crasquin, 2010: 344-346, figs 13W-Z.

? Bairdia hassi - Shi & Chen 2002: 66, pi. 4, figs 11-15.

Localities

- Wujiaping Formation (samples 05PAJ, 28,29), Dajiang section (N25 o 33 , 56 ,, -E106 o 39 , 41 ,, ), Guizhou

Province, South China, Changhsingian, Late Permian.

- Matan and Pingding sections, Guangxi Province, South China, Wuchiapingian, Late Permian (Shi &

Chen 2002).

- Baoqing and Meishan members, Changxing Formation, Meishan section, Zhejiang Province, South

China, Changhsingian, Late Permian (Crasquin et al. 2010).

0 )

300 400 500 600

Height (pm)

Fig. 5. Height/length diagram of Bairdia Ihuberti sp. nov.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Bairdia ? huberti sp. nov.

Fig. 4A-F

Diagnosis

Species attributed with doubt to the genus Bairdia , with relatively elongated carapace, ADB and DB not

distinct, very large AB.

Etymology

Personal dedication to Hubert Colas.

Material examined

Holotype

One carapace (Fig. 4D), sample 05PAJ28, collection number P6M2942.

Paratype

One carapace (Fig. 4C), sample 05PAJ28, collection number P6M2941.

Other material

10 carapaces, several fragments. The species is known from its type locality only.

Type locality

Wujiaping Formation (samples 05PAJ20, 21, 23, 26-28), Dajiang section (25°33 , 56 ,, N-106°39 , 4r , E),

Guizhou Province, South China, Changhsingian, Late Permian.

Measurements (Fig. 5)

L = 581 - 837 pm; H = 332 - 512 pm; H/L = 0.53 - 0.64.

Description

Carapace relatively elongated, ovoid in lateral view; PDB long, straight to slightly convex; PDB-DB

angle around 110°; DB and ADB not clearly distinct and along relative constant H, close to Hmax; VB

concave; AVB rounded and long (~ 75% of Hmax); PVB very short and rounded; AB with large radius

of curvature, maximum located on the upper 1/4 of H; PB with narrow radius of curvature, maximum

located in the lower 1/4 of H; Lmax located around mid-H.

Remarks

High intraspecific variations that mainly affect the ADB and AB are observed in Bairdia ? huberti

sp. nov. They create continuum between Cryptobairdia- like (ABD not distinct: Fig. 4C-E) and Bairdia-

like specimens ( ABD differentiated: Fig. 4A, B, F). The new species is therefore attributed with doubt to

the genus Bairdia. Bairdia 1 . huberti sp. nov. is similar to Bairdia broutini Crasquin, 2010 from the Late

Permian of the Meishan section, Zhejiang Province, South China (Crasquin et al. 2010) because of the

relative constant and high H. However, the new species is more elongated, with longer DB.

Bairdia jeromei sp. nov.

Fig. 4G-P

Diagnosis

Massive and subrectangular species of Bairdia, with DB and VB subparallel, long DB (~ 60% of Lmax),

rhombic dorsal view.

European Journal of Taxonomy 19: 1-34 (2012)

Etymology

Personal dedication to Jerome Cougoul.

Material examined

Holotype

One carapace (Fig. 4G), sample 05PAJ43, collection number P6M2945.

Paratype

One carapace (Fig. 4J), sample 05PAJ43, collection number P6M2948.

Other material

10 carapaces, several fragments. The species is known from its type locality only.

Type locality

Wujiaping and Daye Formations (samples 05PAJ23, 25, 26, 28, 32, 42, 43), Dajiang section

(25°33 , 56”N-106°39 , 41”E), Guizhou Province, South China, Changhsingian and Griesbachian, Late

Permian and Early Triassic.

Measurements (Fig. 6)

L = 722 - 1024 pm; H = 410 - 597 pm; H/L = 0.56 - 0.64.

Description

Carapace massive, elongated and subrectangular in lateral view; surface smooth; LV larger than RV,

overlapping at dorsum and VB; bairdian beaks at AB and PB poorly expressed; PDB, DB and ADB

distinct at both valves; PDB steeply bent backward, long (~ 65% of Hmax) and slightly convex; DB

long (-60% of Lmax) and gently convex; ADB slightly concave at RV, shorter than PDB; ventral

J -]-1-1

400 500 600 700

Height (gm)

Fig. 6. Height/length diagram of Bairdia jeromei sp. nov.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

part rounded; VB long, convex at LV, concave at RV; AVB and PVB rounded, AVB longer than PVB;

carapace slightly postplete to amplete; Lmax below mid-H; dorsal view rhombic with LV’s Wmax in

posterior 1/3 of Lmax, in anterior 1/3 at RV.

Remarks

In lateral view, Bairdia jeromei sp. nov. is similar to Orthobairdia sp.l sensu Crasquin et al. , 2010 from

the Permian of the Meishan section, Zhejiang Province, South China (Crasquin et al. 2010) but here the

dorsal view clearly precludes the generic attribution to Orthobairdia.

Bairdia limatusformis Forel, 2010

Fig. 4Q-T

Silenites limatus - Shi & Chen 1987: 62, pi. 15, figs 15-19.

Bairdia limatusformis - Crasquin et al. 2010: 346-347, figs 9E-G.

Not Silenites limatus - Shi & Chen 2002: 95, pi.27, figs 2-17.

Localities

- Daye Formation (sample 05PAJ43), Dajiang section (25 o 33 , 56”N-106°39 , 41 ,, E), Guizhou Province,

South China, Griesbachian, Early Triassic.

- Baoqing and Meishan members, Changxing formation, Meishan section, Zhejiang Province, South

China, Changhsingian, Late Permian (Shi & Chen 1987; Crasquin et al. 2010).

- Matan and Pingding sections, Guangxi Province, South China, Wuchiapingian, Late Permian (Shi &

Chen 2002).

Genus Bairdiacypris Bradfield, 1935

Bairdiacypris ottomanensis Crasquin-Soleau, 2004

Fig. 10E-H

Bairdiacypris ottomanensis Crasquin-Soleau, 2004: 285-286, pi. 2, figs 13-24.

Bairdiacypris ottomanensis - Crasquin-Soleau & Kershaw 2005: pi. I, figs 10-12. — Forel et al. 2009:

819, fig. 4(1).

Localities

- Daye Formation (samples 05PAJ38, 41), Dajiang section (25°33 , 56 ,, N-106°39 , 41 ,, E), Guizhou

Province, South China, Griesbachian, Early Triassic (Forel et al. 2009; this study).

- Kokarkuyu Formation, Quruk dag section. Western Taurus, Antalya Nappes, Turkey, Early Triassic

(Crasquin-Soleau et al. 2004).

- Feixianguan Formation, Laolongdong section, Sichuan Province, South China, Induan, Early Triassic

(Crasquin & Kershaw 2005).

Genus Fabalicypris Brady, 1880

Fabalicypris parva Wang, 1978

Fig. Ill

Fabalicypris parva Wang, 1978: 293, pi. 2, figs 12a-b, 13a-b.

Fabalicypris hungarica Kozur, 1985: 82, pi. 2, figs 2, 9, 10.

Bairdiacypris opulenta- Shi & Chen 1987: 51, pi. 13, fig. 10.

European Journal of Taxonomy 19: 1-34 (2012)

Fig. 7. Ostracods from the Dajiang section. South China. — A-B. Bairdia sp. 3. A. Carapace, right lateral

view, P6M2963. B. Carapace, dorsal view, P6M2964. — C. Bairdia sp. 4, Carapace, right lateral view,

P6M2965. — D-J. Bairdia sp. 5. D. Carapace, right lateral view, P6M2966. E. Carapace, right lateral

view, P6M2967. F. Carapace, right lateral view, P6M2968. G. Carapace, right lateral view, P6M2969.

H. Carapace, right lateral view, P6M2970. I. Carapace, right lateral view, P6M2971. J. Carapace,

right lateral view, P6M2972. — K-M. Bairdia' ? sp. 6. K. Carapace, left lateral view, P6M2973. L.

Carapace, right lateral view, P6M2974. L. Carapace, right lateral view, P6M2975. — N-S. Bairdia sp.

7. N. Carapace, left lateral view, P6M2976. O. Carapace, right lateral view, P6M2977. P. Carapace,

right lateral view, P6M2978. Q. Carapace, right lateral view, P6M2979. R. Carapace, right lateral view,

P6M2980. S. Carapace, right lateral view, P6M2981. — T-U. Bairdia sp. 8. T. Carapace, right lateral

view, P6M2982. U. Carapace, right lateral view, P6M2983. — V-W. ? Bairdia sp. 8. V. Carapace, right

lateral view, P6M2984. W. Carapace, right lateral view, P6M2985. — X. Bairdia sp. 9, carapace, right

lateral view, P6M2986. - Scale =100 pm.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Fig. 8. Ostracods from the Dajiang section, South China. — A. Bairdia sp. 10, carapace, right lateral

view, P6M2987. — B. Bairdia sp. 11, carapace, right lateral view, P6M2988. — C. Bairdia sp. 12,

carapace, right lateral view, P6M2989. — D. Bairdia sp. 13, carapace, right lateral view, P6M2990. —

E. Bairdia sp. 14, carapace, right lateral view, P6M2991. — F. Bairdia sp. 15, carapace, right lateral

view, P6M2992. — G. Bairdia sp. 16, carapace, right lateral view, P6M2993. — H. Bairdia sp. 17,

carapace, right lateral view, P6M2994. — I-L. Bairdia sp. 18.1. Carapace, right lateral view, P6M2995.

J. Carapace, right lateral view, P6M2996. K. Carapace, right lateral view, P6M2997. L. Carapace, dorsal

view, P6M2998. — M-N. Bairdia sp. 19. M. Carapace, right lateral view, P6M2999. N. Carapace,

right lateral view, P6M3000. — O-P. Bairdia sp. 20. O. Carapace, right lateral view, P6M3001. P.

Carapace, right lateral view, P6M3002. — Q. Bairdia sp. 21, carapace, right lateral view, P6M3003. —

R-S. Bairdia sp. 22. R. Carapace, right lateral view, P6M3004. S. Carapace, right lateral view, P6M3005.

— T. Bairdia sp. 23, carapace, right lateral view, P6M3006. — U-W. Bairdia sp. 24. U. Carapace,

right lateral view, P6M3007. V. Carapace, right lateral view, P6M3008. W. Carapace, right lateral view,

P6M3009. —X. Bairdia cf. sp. 24, carapace, right lateral view, P6M3010. - Scale =100 pm.

European Journal of Taxonomy 19: 1-34(2012)

Fig. 9 . Ostracods from the Dajiang section, South China. — A-D. Bairdia sp. 25. A. Carapace, right

lateral view, P6M3011. B. Carapace, right lateral view, P6M3012. C. Carapace, right lateral view,

P6M3013. D. Carapace, right lateral view, P6M3014. — E-G. Bairdia sp. 26. E. Carapace, right lateral

view, P6M3015. F. Carapace, right lateral view, P6M3016. G. Carapace, right lateral view, P6M3017.

— H-l. Bairdia sp. 27. H. Carapace, right lateral view, P6M3018. I. Carapace, right lateral view,

P6M3019. — J-K. Bairdia sp. 28. J. Carapace, right lateral view, P6M3020. K. Carapace, right lateral

view, P6M3021. — L-N. Bairdia sp. 29. L. Carapace, right lateral view, P6M3022. M. Carapace, right

lateral view, P6M3023. N. Carapace, right lateral view, P6M3024. — O-Q. Bairdia sp. 30. O. Carapace,

right lateral view, P6M3025. P. Carapace, right lateral view, P6M3026. Q. Carapace, right lateral view,

P6M3027. — R-S. Bairdia sp. 31. R. Carapace, right lateral view, P6M3028. S. Carapace, right lateral

view, P6M3029. — T-W. Bairdia sp. 32. T. Carapace, right lateral view, P6M3030. U. Carapace,

right lateral view, P6M3031. V. Carapace, right lateral view, P6M3032. W. Carapace, left lateral view,

P6M3033. - Scale = 100 pm.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Fig. 10. Ostracods from the Dajiang section. South China. — A. Bairdia sp. 33, carapace, right lateral

view, P6M3034. — B. Bairdia sp. 34, carapace, right lateral view, P6M3035. — C. Bairdia sp. 35,

carapace, right lateral view, P6M3036. — D. Bairdia ? sp. 36, carapace, right lateral view, P6M3037. —

E-H. Bairdiacypris ottomanensis Crasquin-Soleau, 2004. E. Carapace, right lateral view, P6M3038. F.

Carapace, right lateral view, P6M3039. G. Carapace, dorsal view, P6M3040. H. Carapace, right lateral

view, P6M3041. — I. Bairdiacypris sp. 1, carapace, right lateral view, P6M3042. — J-K. Bairdiacypris

sp. 2. J. Carapace, right lateral view, P6M3043. K. Carapace, right lateral view, P6M3044. — L-M.

Bairdiacypris sp. 3. L. Carapace, right lateral view, P6M3045. M. Carapace, dorsal view, P6M3046. —

N-Q. Bairdiacypris sp. 4. N. Carapace, right lateral view, P6M3047. O. Carapace, right lateral view,

P6M3048. P. Carapace, right lateral view, P6M3049. Q. Carapace, right lateral view, P6M3050. —

R-T. Bairdiacypris sp. 5. R. Carapace, right lateral view, P6M3051. S. Carapace, right lateral view,

P6M3052. T. Carapace, left lateral view, P6M3053. — U. Bairdiacypris ? sp. 6, carapace, right lateral

view, P6M3054. —V-X. Bairdiacypris sp. 7. V. Carapace, right lateral view, P6M3055. W. Carapace,

right lateral view, P6M3056. X. Carapace, right lateral view, P6M3057. - Scale = 100 pm.

European Journal of Taxonomy 19: 1-34 (2012)

Fig. 11. Ostracods from the Dajiang section. South China. — A-B. Bairdiacypris sp. 8. A. Carapace,

right lateral view, P6M3058. B. Carapace, right lateral view, P6M3059. — C-D. Bairdiacypris sp. 9. C.

Carapace, right lateral view, P6M3060. D. Carapace, right lateral view, P6M3061. — E-F. Bythocyprisl

sp. 1. E. Carapace, right lateral view, P6M3062. F. Carapace, right lateral view, P6M3063. — G.

Bythocypris sp. 2, carapace, right lateral view, P6M3064. — H. Bythocyprisl sp. 3, carapace, right

lateral view, P6M3065. — I. Fabalicypris parva Wang, 1978, carapace, right lateral view, P6M3066.

— J-N. Liuzhinia antalyaensis Crasquin-Soleau, 2004. J. Carapace, right lateral view, P6M3067. K.

Carapace, left lateral view, P6M3068. L. Carapace, left lateral view, P6M3069. M. Carapace, right lateral

view, P6M3070. N. Carapace, right lateral view, P6M3071. — O. ? Liuzhinia antalyaensis Crasquin-

Soleau, 2004, carapace, left lateral view, P6M3072. — P. Liuzhinia sp., carapace, right lateral view,

P6M3073. — Q-S. Liuzhinia sp. 2. Q. Carapace, right lateral view, P6M3074. R. Carapace, left lateral

view, P6M3075. S. Carapace, right lateral view, P6M3076. — T-X. Orthobairdia jeanlouisi sp. nov. T.

Holotype, carapace, right lateral view, P6M3077. U. Carapace, left lateral view, P6M3078. V. Paratype,

carapace, dorsal view, P6M3079. W. Carapace, right lateral view, P6M3080. X. Carapace, right lateral

view, P6M3081. - Scale =100 pm.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Fabalicypris parva - Crasquin-Soleau et al. 2004: 286, pi. 3, figs 4-5. — Mette 2008: pi. 2, fig. 8 -

Crasquin et al. 2010: 353, fig. 9A’-B\

Localities

- Wujiaping Formation (sample 05PAJ26), Dajiang section (25°33 , 56 ,, N-106°39 , 4r , E), Guizhou

Province, South China, Changhsingian, Late Permian.

- Longtan and Changxing Formations, Guizhou and Yunnan Provinces, South China, Wuchiapingian

and Changhsingian, Late Permian (Wang 1978).

- Biikk Mountains, Hungary, Late Moscovian, Carboniferous, Late Permian (Kozur 1985).

- Changxing formation, Meishan section, Zhejiang Province, South China, Changhsingian, Late

Permian (Shi & Chen 1987; Crasquin et al. 2010).

- £uriik dag section, Western Taurus, Turkey, Wuchiapingian and Changhsingian, Late Permian

(Crasquin-Soleau et al. 2004).

- Zal section, Iran, Changhsingian, Late Permian (Mette 2008).

Remarks

According to previously published records of Fabalicypris parva Wang, 1978, this species is known

from the Late Moscovian (Carboniferous; Kozur 1985) to the Changhsingian (Late Permian; Wang

1978; Shi & Chen 1987; Crasquin-Soleau et al. 2004; Mette 2008; Crasquin et al. 2010). This long

stratigraphic range is suspicious: most of the occurrences being late Permian in age, the Carboniferous

one seems questionable. When defining F. hungarica Kozur, 1985 as a synonym of A parva Wang, 1978,

Crasquin et al. (2010) stated that F. hungarica had a more rounded PB and a maximum of convexity

located higher than F. parva. However, because all intermediate shapes between the 2 species were

available, they considered all the specimens as belonging to the same species. This observation together

with the long stratigraphic repartition of the species seems to indicate an evolutionary trend within one

single species. More material is necessary to test this hypothesis.

Genus Liuzhinia Zheng, 1976

Liuzhinia antalyaensis Crasquin-Soleau, 2004

Fig. 11J-N

Liuzhinia antalyaensis Crasquin-Soleau, 2004: 286, pi. 3, figs 6-13.

Liuzhinia antalyaensis - Crasquin-Soleau et al. 2006: 62, pi. 3, figs 12-13. — Crasquin et al. 2008: 249,

pi. 4, figs 9, 12.

Localities

- Samples 05PAJ30-33, 42, Daye Formation, Dajiang section (25 o 33 , 56 ,, N-106°39 , 41 ,, E), Guizhou

Province, South China, Griesbachian, Early Triassic.

- Kokarkuyu Formation, C^uruk dag section. Western Taurus, Turkey, Early Triassic (Crasquin-Soleau et

al. 2004).

- Jinya / Waili section, Fengshan area, Guangxi Province, South China, Griesbachian, Early Triassic

(Crasquin-Soleau et al. 2006).

- Bulla section. Dolomites, Southern Alps, Northern Italy, Early Triassic (Crasquin et al. 2008).

European Journal of Taxonomy 19: 1-34 (2012)

Genus Orthobairdia , Sohn, 1960

Orthobairdia jeanlouisi sp. nov.

Fig. 11T-X

Diagnosis

Species of Orthobairdia with preplete carapace, sharp angles between PDB, DB and ADB (-150°) and

large radius of curvature at AB and PB.

Etymology

Personal dedication to Jean-Louis Forel.

Material examined

Holotype

One carapace (Fig. 11T), sample 05PAJ43, collection number P6M3077.

Paratype

One carapace (Fig. 11V), sample 05PAJ43, collection number P6M3079.

Other material

13 carapaces, several fragments. The species is known from its type locality only.

Type locality

Daye Formation (samples 05PAJ43-45), Dajiang section (25°33 , 56 ,, N-106°39 , 4r , E), Guizhou

Province, South China, Griesbachian, Early Triassic.

200 300 400 500 600

Height (pm)

Fig. 12. Height/length diagram of Orthobairdia jeanlouisi sp. nov.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Measurements (Fig. 12)

L = 422 - 918 jam; H = 250 - 546 jim; H/L = 0.50 - 0.75.

Description

Carapace subtriangular in lateral view; surface smooth; LV overlapping RV all around the carapace,

dorsal overlap thicker; angles between PDB, DB and ADB ~ 150° at both valves; PDB straight with

steep backward slope; DB and ADB long, together ~ 60% of Lmax; DB straight at both valves, bent

backward; ADB straight at LV, slightly convex at RV, less steep than PDB; VB concave; AVB long

and gently rounded; PVB shorter, gently rounded; AB subtriangular, with large radius of convexity,

maximum located above mid-H; PB subtriangular, sharply pointed, with narrower but still large radius

of curvature, maximum located slightly below mid-H; carapace preplete; Lmax located at mid-H or

slightly below.

Remarks

Orthobairdia jeanloaisi sp. nov. can be compared to Orthobairdia texana (Harlton, 1927) sensu Shi

& Chen, 2002 from the Late Permian of Heshan and Yishen, Guangxi Province, South China (Shi &

Chen 2002) but here the species is shorter, posterior maximum of convexity is clearly situated more

dorsally, anterior part is more elongated, ADB steeper. Orthobairdia jeanlouisi sp. nov. is also close

to Orthobairdia exilimarginata Chen, 1987 from the Changhsingian (Late Permian) of the Meishan

section, Zhejiang Province, South China (Shi & Chen 1987), but the new species is less elongated and its

DB is shorter and bent backward. This species can also be related to Orthobairdia meishanensis Chen,

1987 from the Permian of the Meishan section, Zhejiang Province, South China (Shi & Chen 1987),

however, Orthobairdia jeanlouisi sp. nov. has the PB located higher, a shorter DB and a less steep PDB.

Family Pachydomellidae Berdan & Sohn, 1961

Genus Microcheilinella Geis, 1933

Microcheilinella cf. venusta Chen, 1958

Fig. 13Q

Locality

- WujiapingFormation (samples 05PAJ22,24,25,28,29), Dajiang section (25°33 , 56 ,, N-106°39 , 41 ”E),

Guizhou Province, South China, Changhsingian, Late Permian.

Remarks

This species is closely related to Microcheilinella venusta Chen, 1958 from the Early Permian of

Lungtan, Qixia Formation, Nankin (Chen 1958) but here the specimens are more elongated and thinner,

with a more rounded PB. It can also be compared to Microcheilinella cf. venusta Chen, 1958 sensu

Crasquin-Soleau, 2006 from the Early Triassic of the Jinya/Waili section, Guangxi Province, South

China (Crasquin-Soleau et al. 2006). However here the species is more elongated, with longer DB and

no lateral compression in the posterior part.

European Journal of Taxonomy 19: 1-34 (2012)

Superfamily Cypridacea Baird, 1845

Family Paracyprididae Sars, 1923

Genus Paracypris Sars, 1866

Paracypris gaetanii Crasquin-Soleau, 2006

Fig. 13T-X

Paracypris sp. Hao 1992: 42, pi. 1, fig. 24.

Paracypris gaetanii Crasquin-Soleau, 2006: 64, pi. 4, figs 1-4.

Paracypris sp. - Crasquin-Soleau & Kershaw 2005: pi. 1, figs 7-9.

Paracypris gaetanii - Crasquin et al. 2008: 249, pi. 4, fig. 12. — Forel et al. 2009: 819, fig. 4 (5). —

Forel & Crasquin 2011: figs 3F’, 4A.

Localities

- Wujiaping and Daye Formations (samples 05PAJ22, 31, 32, 35, 37-40), Dajiang section

(25°33 , 56”N-106°39 , 4r , E), Guizhou Province, South China, Changhsingian and Griesbachian, Late

Permian and Early Triassic (Forel et al. 2009; this study).

- Feihsienkuan Formation, Zhenfeng, Guizhou Province, South China, Early Triassic (Hao 1992).

- Feixianguan Formation, Laolongdong section, Sichuan Province, South China, Induan, Early Triassic

(Crasquin & Kershaw 2005).

- Jinya/Waili section, Fengshan area, Guangxi Province, South China, Griesbachian, Early Triassic

(Crasquin-Soleau et al. 2006).

- Bulla section, Dolomites, Southern Alps, Northern Italy, Early Triassic (Crasquin et al. 2008).

- Yinkeng Formation, Meishan section, Zhejiang Province, South China, Griesbachian, Early Triassic

(Forel & Crasquin 2011).

- Kokarkuyu Formation, £uriik dag section. Western Taurus, Turkey, Induan, Early Triassic (Forel in

progress).

Remarks

This taxon is one of the rare species which crosses the Permian - Triassic boundary.

Superfamily Cytheracea Baird, 1850

Family Cytherideidae Sars, 1925

Genus Basslerella Kellett, 1935

Basslerella tota Chen & Bao, 1986

Fig. 140-Q

Basslerella tota Chen & Bao, 1986: 123, pi. 1, figs 31-32; pi. 4, figs 7-8.

Basslerella tota - Crasquin-Soleau et al. 2004: 288, pi. 4, figs 9-10 — Yi 2004: pi. 2, fig. 20.

Localities

- Wujiaping Formation (samples 05PAJ24, 26, 28), Dajiang section (25°33 , 56 ,, N-106°39 , 41 ,, E),

Guizhou Province, South China, Changhsingian, Late Permian.

- Chisia formation, Jiangsu Province, South China, Early Permian (Chen & Bao 1986).

- Palmdale Formation, Quruk dag section, Western Taurus, Turkey, Late Permian (Crasquin-Soleau

et al. 2004).

- Kongtonshan section, Fujian Province, South China, Late Permian (Yi 2004).

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Fig. 13. Ostracods from the Dajiang section, South China. — A-B. Petasobairdia sp. 1. A. Carapace,

right lateral view, P6M3082. B. Carapace, right lateral view, P6M3083. — C. Petasobairdia sp. 2,

carapace, right lateral view, P6M3084. — D-E. Petasobairdia sp. 3. D. Carapace, right lateral view,

P6M3085. E. Carapace, right lateral view, P6M3086. — F. Petasobairdia ? sp. 4, carapace, right lateral

view, P6M3087. — G. Spinocypris sp. 1, carapace, right lateral view, P6M3088. — H-E Spinocyprisl sp.

2. H. Carapace, right lateral view, P6M3089.1. Carapace, right lateral view, P6M3090. — J. Kempfina

sp. 1, carapace, right lateral view, P6M3091. — K-L. Silenites sp. 1. K. Carapace, right lateral view,

P6M3092. L. Carapace, right lateral view, P6M3093. — M-N. Silenites sp. 2. M. Carapace, right lateral

view, P6M3094. N. carapace, right lateral view, P6M3095. — O. Silenites sp. 3, carapace, right lateral

view, P6M3096. — P. Silenites sp. 4, carapace, right lateral view, P6M3097. — Q. Microcheilinella

cf. venusta Chen, 1958, carapace, dorsal view, P6M3132. — R. Microcheilinella sp. 1, carapace,

right lateral view, P6M3131. — S. Cetollina ? sp. 1, carapace, right lateral view, P6M3098. — T-X.

Paracypris gaetanii Crasquin-Soleau, 2006. T. Carapace, right lateral view, P6M3099. U. Carapace,

right lateral view, P6M3100. V. Carapace, left lateral view, P6M3101. W. Carapace, left lateral view,

P6M3102. X. Carapace, right lateral view, P6M3103. - Scale =100 pm.

European Journal of Taxonomy 19: 1-34 (2012)

Fig. 14. Ostracods from the Dajiang section, South China. — A-B. Paracypris cf. gaetanii Crasquin-

Soleau, 2006. A. Carapace, right lateral view, P6M3104. B. Carapace, right lateral view, P6M3105.

— C-E. Paracypris sp. 6. C. Carapace, right lateral view, P6M3106. D. Carapace, right lateral view,

P6M3107. E. Carapace, right lateral view, P6M3108. — F. Paracypris sp. 1, carapace, right lateral

view, P6M3109. — G. Paracypris ? sp. 2, carapace, right lateral view, P6M3110. — H. Paracypris ?

sp. 3, carapace, right lateral view, P6M3111. — I. Paracypris ? sp. 4, carapace, right lateral view,

P6M3112. — J-M. Paracypris ? sp. 5. J. Carapace, right lateral view, P6M3113. K. Carapace, right

lateral view, P6M3114. L. Carapace, dorsal view, P6M3151. M. Carapace, right lateral view, P6M3115.

— N. Monoceratinal sp. 1, carapace, right lateral view, P6M3116. — O-R. Basslerella tota Chen &

Bao, 1986. O. Carapace, right lateral view, P6M3117. P. Carapace, right lateral view, P6M3118. Q.

Carapace, right lateral view, P6M3119. R. Carapace, right lateral view, P6M3120. — S. BasslerellaJ. sp.

1, carapace, right lateral view, P6M3121. — T-W. Callicythere postiangusta Wei, 1981. T. Carapace,

right lateral view, P6M3122. U. Carapace, right lateral view, P6M3123. V. Carapace, right lateral view,

P6M3124. W. Carapace, dorsal view, P6M3125. - Scale = 100 pm.

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Family Cytherissinellidae Kashevarova, 1958

Genus Callicythere Wei, 1981

Callicythere postiangusta Wei, 1981

Figs 14T-W, 15A-B

Callicythere postiangusta Wei, 1981: 504, pi. 1, figs 19-22.

Callicythere postiangusta - Crasquin-Soleau & Kershaw 2005: pi. 1, figs 1-6.

Localities

- Samples05PAJ30,31,33,35,37,38,44,DayeFormation,Dajiangsection(25°33 , 56 ,, N-106°39 , 4r , E),

Guizhou Province, South China, Griesbachian, Early Triassic.

- Weiyuan, Leikoupo Formation, Sichuan Province, South China, Middle Triassic (Wei, 1981).

- £uriik dag section, Western Taurus, Turkey, Early Triassic (Forel, in progress).

- Laolongdong section, Feixianguan Formation, Sichuan Province, South China, Early Triassic

(Crasquin-Soleau & Kershaw, 2005).

Ostracod biodiversity variations

Ostracods are found from the top of the Wujiaping formation (05PAJ20) to the base of the Daye

formation (05PAJ45). Of the 26 samples within this interval, 25 yielded ostracods. Species distribution

is summarised in Fig. 2 and abundance and species richness variations are presented in Fig. 16A. In the

productive samples, abundance varies from 1 (05PAJ36) to 2403 (05PAJ41) and species richness from 1

(05PAJ36) to 24 (05PAJ28). A taxa-specimen rarefaction analysis has been performed and suggests that

communities have been well sampled and little sampling bias is expected. Several peaks (P) and drops

(D) are distinguished on the biodiversity curves (Fig. 16A):

(1) Samples 05PAJ20 to 26 are relatively highly diversified, for both species richness and abundance

(Pi).

(2) This phase is followed by a sharp drop from 20 species in 05PAJ25 to 2 in 05PAJ27 (Dl). 05PAJ27

records the minimum of Permian ostracod diversity in Dajiang.

(3) An important diversification is observed just before the Permian - Triassic boundary (PTB) (P2:

05PAJ28, 29). This peak is the maximum of species richness in Dajiang.

(4) Samples 05PAJ28 to 05PAJ30 (D2) bracket the PTB and show an important reduction of specific

richness (from 24 to 3).

(5) Abundance and species richness then increase from 05PAJ31 to 05PAJ33 (P3). Species richness is

however lower than in the Permian assemblages.

(6) Another reduction of diversity, both abundance and specific richness, is recorded in 05PAJ34-36

(D3).

(7) A slight rediversification is observed from 05PAJ37 to 05PAJ40 (P4).

(8) A reduction of species richness, together with a high abundance, is recorded in 05PAJ41 (D4).

(9) 05PAJ42 to 05PAJ45 (P5) show a slight rediversification of the assemblages. During this phase,

species richness is relatively stable while abundance is low.

The biodiversity parameters in Dajiang show striking inversion through the PTB: high species richness/

low abundance for Permian assemblages, low species richness/ high to very high abundance in Triassic

microbialites.

European Journal of Taxonomy 19: 1-34 (2012)

Composition of assemblages

Ostracods in Dajiang belong to 10 superfamilies/families, the distribution of which through the PTB is

illustrated in Fig. 16B. The most abundant superfamily, both in the Permian and the Triassic, is Bairdioidea

(encompassing the genera Acratia , Bairdia , Bairdiacypris, Bythocypris , Fabalicypris , Kempfina ,

Liuzhinia , Microcheilinella, Orthobairdia , Petasobairdia , Silenites and Spinocypris). It is absent from

only 2 assemblages from D3 (05PAJ35, 36). Its proportions are between 20% (05PAJ37) and 100% of

species (05PAJ27, 29, 45). The second most important superfamily is Cypridoidea (genus Paracypris).

It is ubiquitous both in Permian and Triassic assemblages but shows higher proportions in Triassic

assemblages. It is absent from 05PAJ20, 21, 26-29, 38, 44, 45. When it is present, it represents between

Fjm-w

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

6% (05PAJ22) and 100% (05PAJ36) of the species. Cytheroidea (genera Basslerella , Callicythere and

Monoceratina) are found both in Permian and Triassic assemblages. When it is present, it represents

between 4% (05PAJ28) and 50% (05PAJ30, 35) of species. Paraparchitidae (genera Paraparchites ,

Shemonaella , Paraparchitiidae indet.) are present in 2 Permian assemblages and 1 Triassic: 05PAJ21

(11%), 05PAJ28 (4%), 05PAJ44 (9%). The following superfamilies/families only occur in the Permian:

Cavellinidae (genera Cavellina, Sulcella) are found in only 3 assemblages of PI (05PAJ21,24,25) where

they represent between 5% (05PAJ25) and 11% of species (05PAJ21). Kirkbyoidea (genera Amphissites,

Kirkbya, Shleesha ) is part of 3 assemblages from PI (05PAJ20, 22, 24) where it is respectively 14, 6 and

20% of the species. Polycopidae (genus Polycope) is present in 2 assemblages from PI: 05PAJ24 (7%)

and 05PAJ25 (10%). Kloedenellidea (genus Oliganisus) is found in only one assemblage (05PAJ21)

where it represents 11% of the species. Aparchitidae (genus Cyathus ) is present in 2 assemblages:

05PAJ21 (11%) and 05PAJ28 (4%). Finally, only one species belongs to an undetermined superfamily/

family (genus Cetollina). It is part of 05PAJ22 where it represents 6% of species.

In terms of species, the main extinction event occurs in the latest levels of the Wujiaping formation

and the extinction rate is of 98%. Two species cross the PTB in Dajiang: Bairdia jeromei sp. nov. and

Paracypris gaetanii Crasquin-Soleau, 2006 (Fig. 2). They are found in almost all microbialites, their last

occurrences are high in the section (respectively 05PAJ41 and 05PAJ43). Two of the ten genera found

in the microbialites have not been recorded in Permian strata before: Callicythere and Orthobairdia.

Consequently, the faunal turnover is nearly complete at the specific level (94% of new species), partial

at the generic level (20% of new genera) and no renewal is observed at the superfamily/family level.

Importantly, most assemblages from the microbialites show high proportions of very small specimens

(100 pm in length). These specimens belong to the same genera as the larger forms: hence the deviation

of size is not due to the presence of intrinsically smaller genera. This aspect of the faunas will be

addressed in future work.

Discussion

Palaeoenvironmental reconstruction

Most of the ostracods found in the Dajiang section are typical of intertropical warm settings (Crasquin-

Soleau et al. 1999,2004). The presence of Polycope could indicate the influence of cold waters (Kornicker

1959). The palaeoenvironmental preferences of Late Palaeozoic and Lower Triassic ostracods (Peterson

Fig. 15. (opposite page). Ostracods from the Dajiang section. South China. — A-B. Callicythere

postiangusta Wei, 1981. A. Carapace, left lateral view, P6M3126. B. Carapace, left lateral view,

P6M3127. — C-E. Callicythere sp. 1. C. Carapace, right lateral view, P6M3128. D. Carapace, right

lateral view, P6M3129. E. Carapace, dorsal view, P6M3130. — F. Sulcella sp. 1, carapace, right lateral

view, P6M3133. — G. Sulcellal sp. 2, carapace, right lateral view, P6M3134. — H-I. Polycope sp. 1.

I. Carapace, right? lateral view, P6M3135. J. Carapace, right? lateral view, P6M3136. — J. Polycope

sp. 2, carapace, right? lateral view, P6M3137. — K. Polycopel sp. 3, carapace, right? lateral view,

P6M3138. — L-M. Cyathus sp. 1. L. Carapace, right lateral view, P6M3139. M. Carapace, dorsal

view, P6M3140. — N. Cyathus sp. 2, carapace, right lateral view, P6M3141. — O. Amphissites ? sp.

1, carapace, right lateral view, P6M3142. — P. Amphissites ? sp. 2, broken carapace, right lateral view,

P6M3143. — Q. Shleeshal sp. 1, broken carapace, right lateral view, P6M3144. — R. Kirkbya ? sp. 1,

carapace, right lateral view, P6M3145. — S. Kirkbyal sp. 2, carapace, right lateral view, P6M3146. —

T. Oliganisus ? sp. 1, carapace, left lateral view, P6M3147. — U. Paraparchites sp. 1, carapace, right

lateral view, P6M3148. — V. Paraparchitidae indet., carapace, right lateral view, P6M3149. — W.

Shemonaella sp. 1, carapace, left lateral view, P6M3150. - Scale = 100 pm.

European Journal of Taxonomy 19: 1-34 (2012)

& Kaesler 1980; Costanzo & Kaesler 1987; Melnyk & Maddocks 1988a, b; Crasquin-Soleau et al. 1999)

found in Dajiang are summarized as follows (Fig. 16C):

(1) Euryhaline enviro nm ents on the proximal platform (PP): Aparchitidae, Kloedenelloidea, Kirkbyoidea.

(2) Euryhaline environments, shallow to very shallow waters on the intermediate platform (IP):

Cavellinidae, Cytherideidae, Paracyprididae, Paraparchitidae.

(3) Open carbonate environments with normal salinity and oygenation on the distal platform (DP):

Acratiidae, Bairdiidae, Pachydomellidae.

(4) External zone of the distal platform (EDP): Bythocytheridae, Cytherissinellidae.

(5) Any platform environment, in normal salinity: Polycope.

The palaeobathymetric setting is estimated by the composition of assemblages regarding the five

palaeoenvironmental groupings. Each bathymetrical zone is marked by the relative dominance of

the corresponding group, while relative abundances of the other groups give information about the

positioning within the zone determined by the dominant group.

DP forms are found all along the section and are always more than 50% of the assemblages (Fig. 16C).

They indicate an open marine enviro nm ent both in Late Permian and Early Triassic microbialites in

Dajiang. PP forms only occur in Permian assemblages (05PAJ20-22, 24, 28). IP forms show the same

pattern (05PAJ21, 24-26, 28) with one Triassic occurrence (05PAJ44). This documents proximal and

variable conditions in the Permian of Dajiang, in the internal part of the circalittoral stage. EDP forms

appear significantly in the first Triassic assemblage and are found throughout the microbialites. This

suggests that the environmental stabilisation through the PTB can be related to a transgressive trend.

Polycope are found only in Permian assemblages together with the unique Permian occurrence of EDP

forms. This co-existence records a marked deepening of the area, associated with the possible influence of

cold waters. The quasi-absence of PP and IP forms in the microbialites indicates a relative environmental

stability in the deep external circalittoral zone, as revealed by the overwhelming dominance of DP forms

and with significant components of EDP forms.

Worthy to note is the co-occurrence in assemblages 05PAJ25 and 44 of IP and EDP forms, indicating

very different environmental settings. This record is best explained by considering that ecological classes

used in this study not only reflect bathymetry but more generally the stability of the environmental

parameters, here salinity, oxygenation and water depth. The punctual co-occurrence of these IP and EDP

therefore seems to be related to the decoupling of environmental factors.

In summary, ostracod data provide additional evidence for the open-marine context through the

entire PTB interval in Dajiang (Lehrmann et al. 2005). The shallow-subtidal, open-marine platform

environment with relatively low to moderately high current energy of the Wujiaping formation is

documented by ostracod faunas. However, the transgressive trend described here is discordant compared

to the sedimentological proxy that indicates enviro nm ents similar to the underlying Permian skeletal

packstone. This discrepancy can be related to the fact that wave action has been reconstructed based on

interbedded molluscan grainstone, which may only be episodic phenomenon (Lehrmann et al. 2005).

Contribution to the debate on oxygenation

As stated above, oxygenation of marine waters surrounding microbialites is an important challenge to

understand their build-up mechanisms and the survival/recovery phenomenon following the extinction.

In our former work addressing ostracod faunas of the PTB transition in Dajiang, we used the traditional

Lethiers & Whatley model (Lethiers & Whatley 1994) to estimate the oxygen concentration at the base

of the water column (Forel et al. 2009). This tool is now highly questioned and it is no longer possible

to use it (e.g. Brandao & Horne 2009), but we can get a qualitative idea of the water oxygenation

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

independently. Most Permian and Triassic assemblages from Dajiang are dominated by Bairdioidea

(being more than 50% of species, except in 05PAJ30, 35-37, 40, 41; Ford in press). This superfamily

has been associated throughout its history with normal oxygenation settings (Pr R. Maddocks, Univ.

of Houston, pers. comm.), so its dominance is a witness of good oxygenation at the base of the water

I' i' I bioclastic limestone

thrombolite

■ productive samples

— barren sample

Species richness —*—

0 5 10 15 20 25 30 0

Superfam ilies/fam i 1 ies

distribution (%)

25 50 75 100

0 600 1200 1800

Abundance ■ -*■ ■

2400

□ Aparchitidae Q Kirkbyoidea

■ Bairdioidea E3 Kloedenellidea

S3 Cavellinidae ESI Paraparchitiidae

□ Cypridoidea □ Polycopidae

H Cytheroidea ■ Indet

Palaeoecologial

affinities (%)

0 25 50 75 10C

□ PP forms

□ IP forms

E3 DP forms

□ EDP forms

■ Polycope

■ Indet

Fig. 16. Evolution of the ostracod faunas through the PTB in Dajiang. A. Evolution of the number of

species (species richness) and number of specimens (abundance). B. Evolution of the relative proportions

of each superfamily/family. C. Evolution of the relative proportions of each palaeoecological group.

European Journal of Taxonomy 19 : 1-34 ( 2012 )

column. The assemblages where Bairdioidea no longer represent the majority yielded high proportions

of Cypridoidea that have similar requirements regarding oxygen levels (Pr R. Maddocks, Univ. of

Houston pers. comm.). Although the setting through the PTB in Dajiang appears as normoxic, the

biodiversity changes described above undeniably indicate deep ecological modifications. They should

be linked to the relative degradation of the environment, buffered by the refuge created by the microbial

ecosystem (Forel in press). These peculiar assemblages are recognised in all microbialites bearing

sections analysed until now while they are absent from synchronous non-microbial deposits from deeper

settings such as at the Meishan section. Global Stratotype Section and Point of the PTB (Crasquin et

al. 2010; Forel & Crasquin 2011). They are intimately linked to the presence of microbialites (/) in

space, areas lacking microbial mats are devoid of ostracods, (//) in time, faunas disappearing with

microbialites (e.g. Forel et al. in press; Forel in press).

Note on palaeo-geographical distributions and marine ostracods dispersal ways

Several species of this study display a wide distribution around both the Palaeo-Tethys and Neo-Tethys

Oceans: Bairdiacypris Crasquin-Soleau, 2004 , Liuzhinia antalyaensis Crasquin-Soleau, 2004, Paracypris

gaetanii Crasquin-Soleau, 2006, Basslerella tota Chen & Bao, 1986, Callicythere postiangusta Wei,

1981. Because benthic ostracods have no pelagic stage, their migration is performed actively through

locomotion and/or passively by the bottom currents. Their carbonate carapaces exclude them from

depths below the lysocline, their migration is achieved on routes above this level. Their dispersal relies

on the constancy of favourable environmental parameters of water-masses on their route, e.g. oxygen

content, salinity, temperature.

However, active transoceanic dispersal potential of recent forms is relatively limited. Eggs, juveniles

and adults would therefore be passively carried by wind, birds, fish, drifting algae or oceanic currents

(Van Morkhoven 1962; Sandberg 1964; Teeter 1973; Whatley 1988; Babinot & Colin 1992; Tethiers

& Crasquin-Soleau 1995). The characteristics of eggs of marine ostracods set some limits to this

dissemination mode. Whereas eggs of freshwater ostracods are double-walled, those of marine ostracods

are single-walled and do not stand desiccation (Kesling 1961), excluding dispersal by birds and winds

(Teeter 1973). Larvae and adults are less sensitive to wind transportation because of their bigger size

but would probably not withstand it (Teeter 1973). Although Komicker & Sohn (1971) showed the

possibility for adult freshwater ostracods ingested by fish to survive, it is unlikely that marine forms

would have the same resistance. Aquatic plants could also be an important transportation vector for

ostracods. Teeter (1973) found living specimens of Hemicytherura cranekeyensis Puri, 1960 on the

marine algae Turbinaria Lamouroux, 1825 from Honduras. These algae often break into pieces that can

float very far from the coast. Living ostracods have also been observed on the algae Sargassum Agardh,

1820. Turbinaria and Sargassum are found in the tropical area of Atlantic and Pacific Oceans. Once

floating they could drift under the influence of surface currents to climatic zones favorable or not to the

survival of the commensal ostracods. Most of the living ostracods have a high degree of flexibility and

ecological tolerance so they should have a high capacity to successfully invade new environments.

Acknowledgements

This study is part of IGCP 572 ‘Restoration of Marine Ecosystems following the Permian-Triassic Mass

Extinction: lessons for the present" and was undertaken with the support of the French CNRS Research

Team UMR 7207 CR2P and the Chinese programs NSFC (40839903 and 40921062) and 111 (B08030).

I am deeply indebted to my PhD supervisor Dr. Sylvie Crasquin (CNRS - CR2P) for her availability

and great help in this study. I thank Prof. Feng Qinglai (China University of Geosciences, Wuhan) for

his help during fieldwork on the Dajiang section. I also thank Martine Fordant (UPMC, Paris) for the

processing of material and Alexandre Lethiers (UPMC, Paris) for his help with the illustrations. I am

also grateful to Dr. Carys Bennett (Universite de Lille, France) and Dr. Vincent Perrier (University of

FOREL M.B., Ostracods from the Permian-Triassic boundary in Guizhou

Tartu, Estonia) for their critical reviews and constructive suggestions, which have greatly improved the

quality of this analysis.

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Manuscript received: 3 April 2012

Manuscript accepted: 26 June 2012

Published on: 1 August 2012

Topic editor: Christian de Muizon

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Natural History Museum, London, United Kingdom; Royal Belgian Institute of Natural Sciences,

Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark.