Heterochronic growth of ostracods (Crustacea) from microbial deposits in the aftermath of the...

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Heterochronic growth of ostracods (Crustacea) frommicrobial deposits in the aftermath of the end-PermianextinctionMarie-Béatrice Forelaa State Key Laboratory of Geological Process and Mineral Resources, China University ofGeosciences, No. 388, Lumo Road, Wuhan 430074, ChinaPublished online: 14 May 2014.

To cite this article: Marie-Béatrice Forel (2014): Heterochronic growth of ostracods (Crustacea) from microbial deposits inthe aftermath of the end-Permian extinction, Journal of Systematic Palaeontology, DOI: 10.1080/14772019.2014.902400

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Heterochronic growth of ostracods (Crustacea) from microbial deposits in theaftermath of the end-Permian extinction

Marie-B�eatrice Forel*

State Key Laboratory of Geological Process and Mineral Resources, China University of Geosciences, No. 388, Lumo Road,Wuhan 430074, China

(Received 14 May 2013; accepted 14 November 2013)

The Kokarkuyu Formation (Early Triassic) from the C€ur€uk Da�g section, Western Taurus, Antalya Nappes, Turkey wascarefully sampled for a taxonomic study of the ostracods. A total of 57 species belonging to 15 genera are recognized.Three species are newly described: Monoceratina hussonae sp. nov., Reviya sylvieae sp. nov. and Eukloedenellaadcapitisdolorella sp. nov. The present data are the first Neo-Tethyan illustration of ostracod survival in the aftermath ofthe end-Permian extinction in a refuge of microbial origin. Ostracods are abundant within the microbialites at the base ofthe formation and allow the reconstruction of ontogenetic series for nine species. Shape variations through ontogeny aredescribed for seven species: Bairdia? kemerensis, Praezabythocypris? ottomanensis, Liuzhinia antalyaensis, Paracyprisgaetanii, Reviya curukensis, R. sylvieae and Eukloedenella adcapitisdolorella. Paedomorphosis through deceleration andperamorphosis through acceleration are identified as secondary survival strategies following the end-Permian extinction.

http://zoobank.org/urn:lsid:zoobank.org:pub:6849AB0D-B085-47BE-950A-BC98CC50C4E3

Keywords: Crustacea; ostracods; Early Triassic; microbialites; heterochrony; C€ur€uk Da�g

Introduction

The Permian–Triassic Boundary (PTB; �252 Ma) is

marked by the most dramatic extinction of the Phanero-

zoic, when 80–96% of marine species went extinct (e.g.

Sepkoski 1984; Erwin 1993; Benton & Twitchett 2003).

According to some authors, global ocean anoxia was

widespread during the latest Permian (Changhsingian),

increasing close to the main extinction and continuing

during the Early Triassic (e.g. Bond & Wignall 2010). An

important trigger of the extinction was the eruption of the

Siberian Traps, causing global warming, ocean acidifica-

tion and possible destruction of atmospheric ozone (e.g.

Svensen et al. 2009). Subsequent enhanced weathering

and nutrient run-off might have increased the pre-existing

ocean anoxia (e.g. Payne & Clapham 2012). Marine eco-

systems saw the replacement of the Late Permian benthic

shelly communities dominating shallow marine environ-

ments by microbial communities. These microbialites

were abundant in low-latitude, shallow-marine carbonate

shelves in the central Tethys where they occupied envi-

ronments formerly populated by Late Permian reefs but

extending also into deeper waters (see Kershaw et al.

2012 for a synthesis).

Current evidence regarding oxygen levels associated

with microbial expansions is conflicting, with evidence of

both low-oxygen markers (Bond & Wignall 2010; Liao

et al. 2010; Chen et al. 2011) and abundant benthic shelly

faunas dominated by ostracods, and occasionally micro-

gastropods, micro-brachiopods, foraminifers, bivalves and

conodonts (e.g. Baud et al. 1997; Crasquin-Soleau & Ker-

shaw 2005; Groves et al. 2005; Richoz 2006; Br€uwhileret al. 2008; Ezaki et al. 2008; Forel et al. 2009; Song et al.

2009; Kaim et al. 2010; Hautmann et al. 2011; Yang et al.

2011; Forel 2012; Frisk et al. 2012; Forel et al. 2013a).

These unusual surviving faunas are explained by a two-

step oxygenation mechanism of surrounding waters by cya-

nobacterial activity (Forel et al. 2013b).

Through the end-Permian extinction (EPE), ostracods

(Crustacea) show specific extinction rates ranging from

74 to 100% (Crasquin & Forel 2013). Their recovery is

assumed to be complete in the Late Triassic when biodi-

versity returned to levels similar to those of Middle Perm-

ian faunas. Ostracod faunas described from Early Triassic

rocks are rare, so that survival mechanisms and refuge

zones at the origin of later recovery remain unclear. The

assemblages considered here from the C€ur€uk Da�g section

(Turkey) illustrate secondary survival mechanisms, based

on the modifications of the development (heterochronies),

in a refuge of microbial origin already known from sev-

eral localities from the Panthalassa and Palaeo-Tethys

Oceans (Forel 2012, 2013; Forel et al. 2013a, b).

*Corresponding author. Email: [email protected]

� The Trustees of the Natural History Museum, London 2014. All Rights Reserved.

Journal of Systematic Palaeontology, 2014

http://dx.doi.org/10.1080/14772019.2014.902400

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mailto:[email protected]

http://dx.doi.org/10.1080/14772019.2014.902400

Previous work on the PTB interval from C€ur€uk Da�gdocumented 38 ostracod species from 22 genera (Cras-

quin-Soleau et al. 2002, 2004a, b). To assess ostracod

populations during and after microbial build-up in the

aftermath of the EPE, the Kokarkuyu Formation of Early

Triassic age was carefully resampled. Fifty-seven species,

including three new species, belonging to 15 genera were

found and are reported in the present work. The data from

Crasquin-Soleau et al. (2002, 2004a, b) are included in

our database to provide the most complete record cur-

rently available for the ostracod biodiversity variations

and palaeo-environmental changes throughout the Kokar-

kuyu formation and more generally in the Neo-Tethys

area (Fig. 1).

Material and methods

The material described in the present work comes from

the C€ur€uk Da�g section (36� 410 32400 N, 30� 270 40.100 E),located in the Western Taurus, Antalya Nappes, Turkey

(Fig. 1). A succinct description of the lithological

succession is provided here: the reader is referred to Baud

et al. (1997, 2005), Crasquin et al. (2009) and Kershaw

et al. (2010) and references therein for further details. The

Upper Permian Pamucak Formation consists of wacke-

stones and grainstones/packstones. It is overlain by the

Kokarkuyu Formation, 40 metres thick in the C€ur€uk Da�gsection. The first 15 metres of this formation are com-

posed of cyclic microbial limestones, mostly made of stro-

matolites, thrombolites and hybrid microbialites (Kershaw

et al. 2010). The discovery of the conodonts Isarcicella

staeschei at the base of the formation and Hindeodus

parvus 55 cm higher up (Richoz 2006) indicates that

the base of the microbialites is already in the Triassic. It

is followed by 3 metres of calcilutites, overlain by

wackestones. The Upper Permian ostracod assemblages

have been described by Crasquin-Soleau et al. (2004a, b).

Samples spanning the entire exposed Kokarkuyu

Formation were collected to describe ostracod assemb-

lages from facies below, within and above the microbia-

lites (Fig. 2).

External features of carapaces are necessary to deter-

mine ostracods. Extraction of ostracods from the matrix

Figure 1. Middle Permian palaeogeographical map (modified from Crasquin-Soleau et al. 2001) and modern map showing the locationof the C€ur€uk Da�g section, Taurus, Turkey.

2 M. B. Forel

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using acid dissolution is precluded because their carbon-

ate carapaces are enclosed in calcareous rocks. Disaggre-

gation of dehydrated hard limestone and release of

ostracod shells were performed by the hot acetolysis tech-

nique (Lethiers & Crasquin-Soleau 1988; Crasquin-Sol-

eau et al. 2005). A total of 140 samples were collected

and processed, of which 45 yielded ostracods (Fig. 3).

The productive samples are located in the lowermost Tri-

assic microbialites; only two samples yielded specimens

above the microbial crust. Because of poor preservation,

many species are left in open nomenclature. However, it

is important to figure all the material of this particular

interval (Figs 4, 6–8, 11, 12, 14, 20).

Figure 2. Stratigraphical log of the C€ur€uk Da�g section (Taurus, Turkey) and location of studied samples: samples labelled CDxx arefrom the present work, while those labelled TKxx are from Crasquin-Soleau et al. (2004a, b). Abbreviations: L.P.: Late Permian; Pam.:Pamucak Formation. (continue)

Heterochronic growth of ostracods (Crustacea) from microbial deposits 3

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Ostracods have determinate growth: they display a step-

wise development resulting from moulting (ecdysis) and

stop moulting when they reach adult stage. Specimens are

here assigned to their corresponding instar group based on

size. The preservation makes it impossible to study hinge

structure and muscle scars. Lengths of specimens were

plotted against heights in scatter plots to distinguish size

clusters which discriminate successive juveniles and

adults. In the fossil record, the recognition of different

growth stages is difficult or impossible because of time-

averaging of specimens from different environments or

seasons (e.g. specimens of the same species tend to be

smaller in the spring than in the autumn; Morales-Ram�ırez& Jakob 2008). To overcome this difficulty, I computed

Kernel density maps (Gaussian Kernel distribution, col-

umns ¼ 100, rows ¼ 100, radius ¼ 15) using PAST soft-

ware (Hammer et al. 2001; Hammer & Harper 2005). For

each species yielding at least 15 specimens (from this

Figure 2. (Continued)

4 M. B. Forel

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Figure 3. Distribution of ostracod species through the productive portion of the C€ur€uk Da�g section, Taurus, Turkey.


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work and the literature), it allowed the discrimination of

density patterns of individuals corresponding to different

ontogenetical stages. Successive instars of nine species

are recognized and described here. Of the 57 species iden-

tified, the Systematic palaeontology section does not

include those species left in open nomenclature.

All specimens are deposited in the collections of the

Pierre et Marie Curie University, Paris, France, under

numbers P6Mxxx.

Systematic palaeontology

For the three new species a full description is given. For

other species only the synonymy and occurrences are spec-

ified. The classification employed is that of Moore (1961),

modified after Lethiers (1981) and Horne et al. (2002).

Morphological abbreviations: RV, right valve; LV, left

valve; AB, anterior border; PB, posterior border; DB, dor-

sal border; VB, ventral border; ADB, anterodorsal border;

Figure 4. Ostracods from the C€ur€uk Da�g section, Taurus, Turkey. All images are right lateral views of complete carapaces. A, Acratia?sp. 1, P6M3151. B–I, Bairdia? kemerensis Crasquin-Soleau, 2004b; B, P6M3152; C, P6M3153; D, P6M3154; E, P6M3155; F,P6M3156; G, P6M3157; H, P6M3158; I, P6M3159. J, K, Bairdia sp. 1 sensu Crasquin-Soleau et al., 2006b; J, P6M3160; K,P6M3161. L–N, Bairdia sp. 2; L, P6M3162;M, P6M3163; N, P6M3164. O, P, Bairdia sp. 3; O, P6M3165; P, P6M3166. Q–U, Bairdiasp. 4; Q, P6M3167; R, P6M3168; S, P6M3169; T, P6M3170; U, P6M3171. V–X, Bairdia sp. 5; V, P6M3172; W, P6M3173; X,P6M3174. Scale bars are 100 mm.

6 M. B. Forel

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AVB, anteroventral border; PDB, posterodorsal border;

PVB, posteroventral border; ACA, anterior cardinal angle

(for Palaeocopida only); PCA, posterior cardinal angle

(for Palaeocopida only); ADA, anterodorsal angulation;

PDA, posterodorsal angulation; AVA, anteroventral angu-

lation (angle between AVB and VB); PVA, posteroventral

angulation (PVA as angle between VB and PVB); S1,

anterior sulcus; S2, median sulcus; H, height; Hmax, max-

imal height; L, length; Lmax, maximal length.

Class Ostracoda Latreille, 1806

Subclass PodocopaM€uller, 1894Order PodocopidaM€uller, 1894Suborder Podocopina Sars, 1866

Superfamily Bairdioidea Sars, 1887

Family Bairdiidae Sars, 1887

Genus BairdiaMcCoy, 1844

Bairdia? kemerensis Crasquin-Soleau, 2004

(Fig. 4B–I)

2004b Bairdia? kemerensis Crasquin-Soleau in Crasquin-

Soleau et al.: 285, pl. 2, figs 1–5.

Occurrence. Samples S4, S3, S7, 08CD17, 20, 26, 27,

29, 31, TK55b-58, TK120, TK121, TK123–126, C€ur€ukDa�g section (Fig. 3), Western Taurus, Turkey, Kokarkuyu

Formation, Griesbachian, Induan, Early Triassic (this

work; Crasquin-Soleau et al. 2004a, b)

Remarks. H versus L of all Bairdia? kemerensis speci-

mens are shown in Figure 5. They cluster into five differ-

ent ontogenetic stages, labelled A-4, A-3, A-2, A-1 and

Adults, in ascending order. The shape of B.? kemerensis

appears as relatively conservative through ontogeny. The

carapace becomes globally stockier but the modifications

mostly affect the PB that enlarges with the radius of cur-

vature migrating dorsally. Sexual dimorphism of modern

Bairdioidea is generally poorly expressed on the lateral

outline, contrary to soft parts and internal structures (e.g.

Maddocks & Iliffe 1986; Maddocks 1991, 2013). In the

absence of preserved soft parts and because of overall

poor preservation, only the lateral view is available. The

adult specimen in Crasquin-Soleau et al. (2004b, pl. 2, fig.

5) displays the largest posterior-half of the entire lineage

and could therefore be interpreted as heteromorph (,).This sexual dimorphism pattern is similar to that observed

for Bairdia nishiwakii in the Middle Permian of Japan

(Tanaka et al. 2013) and is therefore the second record of

possible sexual dimorphism for fossils of the genus

Bairdia.

Genus Liuzhinia Zheng, 1976

Liuzhinia antalyaensis Crasquin-Soleau, 2004

(Fig. 8P–Z)

2004b Liuzhinia antalyaensis Crasquin-Soleau in

Crasquin-Soleau et al.: 286, pl. 3, figs 6–13.

2006a Liuzhinia antalyaensis Crasquin-Soleau; Crasquin-

Soleau et al.: 62, pl. 3, figs 12, 13.

2008 Liuzhinia antalyaensis Crasquin-Soleau; Crasquin

et al.: 249, pl. 4, figs 9, 12.

2012 Liuzhinia antalyaensis Crasquin-Soleau; Forel: 19,

fig. 11J–N.

Occurrence. Samples 08CD03A, C, D, D1-H, 04, 05, 15,

23, 27, 28, TK55–56, C€ur€uk Da�g section (Fig. 3), Western

Figure 5. Length/height scatter plot for Bairdia? kemerensis. Specimens are from C€ur€uk Da�g section (this work and Crasquin-Soleauet al. 2004b). All specimens are to the same scale.


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Figure 6. Ostracods from the C€ur€uk Da�g section, Taurus, Turkey. All images are right lateral views of complete carapaces. A–C, Bair-dia sp. 6; A, P6M3175; B, P6M3176; C, P6M3177. D, Bairdia sp. 7, P6M3178. E, F, Bairdia sp. 8; E, P6M3179; F, P6M3180. G–S,Bairdia sp. 9; G, P6M3181; H, P6M3182; I, P6M3183; J, P6M3184; K, P6M3185; L, P6M3186; M, P6M3187; N, P6M3188; O,P6M3189; P, P6M3190; Q, P6M3191; R, P6M3192; S, P6M3193. T, U, Bairdia sp. 10; T, P6M3194; U, P6M3195. V–A0, Bairdia sp.11; V, P6M3196;W, P6M3197; X, P6M3198; Y, P6M3199; Z, P6M3200; A0, P6M3201. Scale bars are 100 mm.

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Taurus, Turkey, Kokarkuyu Formation, Griesbachian,

Induan, Early Triassic (this work; Crasquin-Soleau et al.,

2004b). Jinya/Waili section, Fengshan area, Guangxi Prov-

ince, South China (Crasquin-Soleau et al. 2006a). Bulla

section, Dolomites, Southern Alps, Northern Italy (Cras-

quin et al. 2008). Dajiang section, Daye Formation,

Guizhou Province, South China (Forel 2012). All occur-

rences from Griesbachian, Induan, Early Triassic.

Remarks. Fig. 9 plots H versus L for all specimens of

Liuzhinia antalyaensis. Five successive ontogenetic stages

are distinguished, from A-4 to Adults. Modifications

Figure 7. Ostracods from the C€ur€uk Da�g section, Taurus, Turkey. All images are right lateral views of complete carapaces. A–D, Bair-dia sp. 12; A, P6M3202; B, P6M3203; C, P6M3204; D, P6M3205. E–G, Bairdia sp. 13; E, P6M3206; F, P6M3207; G, P6M3208. H–L, Bairdia sp. 14; H, P6M3209; I, P6M3210; J, P6M3211; K, P6M3212; L, P6M3213. M–Q, Bairdia sp. 15; M, P6M3214; N,P6M3215; O, P6M3216; P, P6M3217; Q, P6M3218. R, S, Bairdia sp. 16; R, P6M3219; S, P6M3220. T–V, Bairdia sp. 17; T,P6M3221; U, P6M3222; V, P6M3223.W, Bairdia sp. 18, P6M3224. X, Bairdia sp. 19, P6M3225. Scale bars are 100 mm.


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Figure 8. Ostracods from the C€ur€uk Da�g section, Taurus, Turkey. All images are right lateral views of complete carapaces. A, Bairdiasp. 20, P6M3226. B, Bairdia sp. 21, P6M3227. C, D, Bairdia sp. 22; C, P6M3228; D, P6M3229. E, Bairdia sp. 23, P6M3230. F–H,Praezabythocypris? ottomanensis (Crasquin-Soleau, 2004b); F, P6M3231; G, P6M3232; H, P6M3233. I, J, Bairdiacypris sp. 1; I,P6M3234; J, P6M3235. K, Bairdiacypris sp. 2, P6M3236. L, M, Bairdiacypris sp. 3 sensu Forel, 2012; L, P6M3237; M, P6M3238. N,Bairdiacypris sp. 4, P6M3239. O, Bythocypris? sp. 1, P6M3240. P–Z, Liuzhinia antalyaensis Crasquin-Soleau, 2004; P, P6M3241; Q,P6M3242; R, P6M3243; S, P6M3244; T, P6M3245; U, P6M3246; V, P6M3247; W, P6M3248; X, P6M3249; Y, P6M3250; Z,P6M3251. Scale bars are 100 mm.

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mainly affect the posterior half of the carapace, which

becomes higher with a larger radius of curvature through

ontogeny. No dimorphic features are recognized.

Genus Praezabythocypris Kozur, 1985

Praezabythocypris? ottomanensis (Crasquin-

Soleau, 2004)

(Fig. 8F–H)

2004b Bairdiacypris ottomanensis Crasquin-Soleau in

Crasquin-Soleau et al.: 285, pl. 2, figs 13–24.

2005 Bairdiacypris ottomanensis Crasquin-Soleau;

Crasquin-Soleau & Kershaw: pl. 1, figs 10–12.

2009 Bairdiacypris ottomanensis Crasquin-Soleau; Forel

et al.: 819, fig. 4(1).

2008 Praezabythocypris sp. 1; Mette: pl. 2, figs 10, 11.

Figure 10. Length/height scatter plot for Praezabythocypris? ottomanensis. Specimens come from C€ur€uk Da�g (this work and Crasquin-Soleau et al. 2004b), Dajiang, Guizhou, China (Forel 2012), and Iran (Mette 2010). All the specimens are to the same scale.

Figure 9. Length/height scatter plot for Liuzhinia antalyaensis. Specimens come from C€ur€uk Da�g (This work andCrasquin-Soleau et al. 2004b), Dajiang, Guizhou, China (Forel 2012), Bulla, Italy (Crasquin et al. 2008) and Guangxi, China (Crasquin-Soleau et al. 2006a). All specimens are to the same scale.


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Figure 11. Ostracods from the C€ur€uk Da�g section, Taurus, Turkey. All images are right lateral views of complete carapaces. A, Liuzhi-nia cf. guangxiensis Crasquin-Soleau, 2006a, P6M3252. B, Liuzhinia sp. 1, P6M3253. C, Liuzhinia sp. 2, P6M3254. D–F, Liuzhinia sp.3; D, P6M3255; E, P6M3256; F, P6M3257. G–I, Liuzhinia sp. 4; G, P6M3258; H, P6M3259; I, P6M3260. J, Petasobairdia sp. 1,P6M3261. K–N, Praezabythocypris cf. pulchra Kozur, 1985; K, P6M3262; L, P6M3263; M, P6M3264; N, P6M3265. O–Q, Rectobair-dia sp. 1; O, P6M3266; P, P6M3267; Q, P6M3268. R–T, Rectobairdia sp. 2; R, P6M3269; S, P6M3270; T, P6M3271. U, V, Silenites?sp. 1; U, P6M3272; V, P6M3273.W–Z, Silenites sp. 2;W, P6M3274; X, P6M3275; Y, P6M3276; Z, P6M3277. Scale bars are 100 mm.

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2010 Praezabythocypris? ottomanensis Crasquin-Soleau;

Mette: 28, pl. 4, figs 1–4.

non 2010 Bairdiacypris ottomanensis Crasquin-Soleau;

Liu et al.: fig. 3(1).

2012 Bairdiacypris ottomanensis Crasquin-Soleau; Forel:

13, fig. 10E–H.

Occurrence. Samples 08CD04, 09, 10, S4, S3, S10,

08CD14’, S9, 08CD22, 31, TK120, 125, 126, C€ur€uk Da�gsection (Fig. 3), Western Taurus, Turkey, Kokarkuyu For-

mation, Griesbachian, Induan, Early Triassic (this work;

Crasquin-Soleau et al. 2004a, b). Laolongdong section,

Feixianguan Formation, Griesbachian, Induan, Early

Triassic, Sichuan Province, South China (Crasquin-Soleau

& Kershaw 2005). Zal section, Ali Bashi and Zal forma-

tions, Late Permian–Early Triassic, north-west Iran

(Mette 2008, 2010). Dajiang section, Daye Formation,

Griesbachian, Induan, Early Triassic, Guizhou Province,

South China (Forel et al. 2009; Forel 2012).

Remarks. Figure 10 plots H versus L for all known Prae-

zabythocypris? ottomanensis specimens. They cluster into

four successive ontogenetic groups, in ascending order:

A-3, A-2 (Crasquin-Soleau et al. 2004b, pl. 2, fig. 13),

A-1 (Fig. 8H; Forel 2012, fig. 10E–G), and Adults

(Fig. 8F, G). Through ontogeny, the carapace becomes

more preplete and PB more rounded with a maximum

of curvature located higher. The specimen identified as

Figure 12. Ostracods from the C€ur€uk Da�g section, Taurus, Turkey. All images except R, are right lateral views of complete carapaces. A–C,Silenites sp. 3; A, P6M3278; B, P6M3279; C, P6M3280. D, E, Silenites sp. 4; D, P6M3281; E, P6M3282. F–H, Silenites sp. 5; F, P6M3283;G, P6M3284; H, P6M3285. I, J, Silenites sp. 6; I, P6M3286; J, P6M3287. K, L, Microcheilinella sp. 1; K, P6M3288; L, P6M3289. M,Microcheilinella sp. 2, P6M3290. N–Q, Paracypris gaetanii Crasquin-Soleau, 2006a; N, P6M3291; O, P6M3292; P, P6M3293; Q, P6M3294.R–T, Paracypris sp. 1; R, left lateral view of a complete carapace, P6M3295; S, P6M3296; T, P6M3297. Scale bars are 100 mm.


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Praezabythocypris? ottomanensis by Liu et al. (2010) is

excluded because it clearly displays a bairdian shape with

short DB and no angulation at dorsum.

Praezabythocypris cf. pulchra Kozur, 1985

(Figs 11K–N)

cf. 1985 Praezabythocypris pulchra Kozur: 84, taf. 17,

fig. 13.

Occurrence. Samples 08CD04, 05, S1, 08CD10, 14’, S8,

08CD26, 28, C€ur€uk Da�g section (Fig. 3), Western Taurus,

Turkey, Kokarkuyu Formation, Griesbachian, Induan,

Early Triassic (this work).

Remarks. The present species differs from Praezabytho-

cypris pulchraKozur, 1985 from the Middle and Late Perm-

ian of Hungary (Kozur 1985) by its steeper PDB, a more

inclined DB at RV, a longer and less rounded AVB, and the

posterior maximum of curvature located more ventrally.

Superfamily Cypridoidea Baird, 1845

Family Paracyprididae Sars, 1923

Genus Paracypris Sars, 1910

Paracypris gaetanii Crasquin-Soleau, 2006

(Fig. 12N–Q)

1992 Paracypris sp.; Hao: 42, pl. 1, fig. 24.

2005 Paracypris sp. sensu Hao 1999 [sic]; Crasquin-

Soleau & Kershaw: pl. I, figs 7–9.

2006a Paracypris gaetanii Crasquin-Soleau in Crasquin-


2008 Paracypris gaetanii Crasquin-Soleau; Crasquin

et al.: 249, pl. 4, fig. 11.

2009 Paracypris gaetanii Crasquin-Soleau; Forel et al.:

819, fig. 4(5).

2011 Paracypris gaetanii Crasquin-Soleau; Forel &

Crasquin: figs 3F’–4A.

2012 Paracypris gaetanii Crasquin-Soleau; Forel: 22,

fig. 13T–X.

Occurrence. Samples 08CD03F, 09, 10, S4, S10,

08CD15, S7, 08CD22, 26–31, C€ur€uk Da�g section (Fig. 3),

Western Taurus, Turkey (this work; Crasquin-Soleau

et al. 2004a, b). Kokarkuyu Formation, Griesbachian,

Induan, Early Triassic. Feihsienkuan Formation, Griesba-

chian, Induan, Early Triassic, Zhenfeng, Guizhou Prov-

ince, South China (Hao 1992). Laolongdong section,

Feixianguan Formation, Griesbachian, Induan, Early

Triassic, Sichuan Province, South China, Induan, Early

Triassic (Crasquin-Soleau & Kershaw 2005). Jinya/Waili

section, Griesbachian, Induan, Early Triassic, Fengshan

area, Guangxi Province, China (Crasquin-Soleau et al.

2006a). Bulla section, Griesbachian, Induan, Early Trias-

sic, Dolomites, Southern Alps, Northern Italy (Crasquin

et al. 2008). Dajiang section, Wujiaping and Daye Forma-

tions, Guizhou Province, China, Late Permian–Early Tri-

assic (Forel et al. 2009; Forel 2012). Meishan section,

Figure 13. Length/height scatter plot for Paracypris gaetanii. Specimens come from C€ur€uk Da�g (this work), Dajiang, Guizhou, China(Forel 2012), Meishan, Zhejiang, China (Forel & Crasquin 2011), Bulla, Italy (Crasquin et al. 2008), Guangxi, China (Crasquin-Soleauet al. 2006a), Laolongdong, Sichuan, China (Crasquin-Soleau & Kershaw 2005), and Zhenfeng, Guizhou, China (Hao 1992). All speci-mens are at the same scale.

14 M. B. Forel

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Griesbachian, Induan, Early Triassic, Zhejiang Province,

China (Forel & Crasquin 2011).

Remarks. Figure 13 plots H versus L for all Paracypris

gaetanii specimens. They scatter into five ontogenetic

stages, labelled A-4 to Adults. Changes through growth

stages are mainly located at the posterior-half of the cara-

pace, becoming higher and more rounded. Interestingly,

some of the larger specimens display distinct morpholo-

gies at PB: (i) tapered with a narrow radius of curvature,

maximum located relatively ventrally (e.g. Fig. 12N;

Crasquin-Soleau et al. 2006a, pl. 4, fig. 2; Hao 1992, pl. 1,

fig. 24); (ii) rounded with a larger radius of curvature,

maximum located higher (e.g. Forel 2012, fig. 13T; Cras-

quin-Soleau & Kershaw 2005, pl. 1, figs 7–9). These two

groups could respectively correspond to tecnomorphs (<)and heteromorphs (,).

Superfamily Cytheroidea Baird, 1850

Family Bythocytheridae Sars, 1928

GenusMonoceratina Roth, 1928

Monoceratina hussonae sp. nov.

(Fig. 14A–J)

Diagnosis. Subtriangular species of Monoceratina with

well-expressed S2 and faint S1; adventral alae faint; AB

and PB rimmed; PB rounded; lateral surface reticulated.

Derivation of name. Dedicated to Dr Doroth�ee Husson,

Northwestern University, Chicago, USA.

Material. Holotype: P6M3300, one complete carapace

(Fig. 14C), sample S3. Paratype: P6M3305, one complete

carapace (Fig. 14H), sample 08CD10. All from the C€ur€ukDa�g section, Western Taurus, Turkey, Kokarkuyu Forma-

tion, Griesbachian, Induan, Early Triassic. Additional

material: 152 complete carapaces and several fragments.

Occurrence. Samples CD08A7, 08CD03A-C,

08CD03D1-H, 04, 05, S1, 08CD07, S2, 08CD10, 11, 13,

14, S3, 08CD17, 20, 23, 26, 29–31, C€ur€uk Da�g section

(Fig. 3), Western Taurus, Turkey, Kokarkuyu Formation,

Griesbachian, Induan, Early Triassic.

Description. Carapace subtriangular, elongated, rela-

tively massive for the genus (H/L generally �0.5); both

valves similar in shape, size and ornamentation, no over-

lap; Hmax at or in front of anterior one-third of Lmax;

Lmax slightly above midH.

DB: long (�65–80% of Lmax), straight to slightly con-

vex; eventual presence of faint shoulders anteriorly to

PDA and posteriorly to ADA (e.g. Fig. 14F, H, J); PDA

�100�, ADA �130�.VB: long, oblique (angle with horizontal line �15�),

straight to slightly concave.

AB: broadly rounded as half-circle, maximum of curva-

ture at midH; triangular with angulation (�120–125�)located above midH at some specimens (e.g. Fig. 14F, G).

PB: tapered, rounded with a narrower radius of curva-

ture, maximum of convexity in the upper 1/3 of Hmax.

Lateral surface of the valves evenly swollen; smooth

rim along AB and PB; 2 sulci at dorsal part: S2 character-

istic of the genus well expressed in the upper-third of

Hmax, S1 poorly defined in the upper quarter of Hmax;

adventral alae short and faint, slightly pointing outward;

swollen surface faintly reticulated.

Dimensions. H ¼ 142–198 mm; L ¼ 249–342 mm; H/L

¼ 0.51–0.64 (Fig. 15).

Remarks. Until now, the only Griesbachian record of the

Monoceratina genus is an unconfirmed occurrence from

the Late Griesbachian of Pakistan (Sohn 1970). The pres-

ent species is therefore the first known occurrence of the

genus in the Early Griesbachian, directly after the end-

Permian extinction. Because of its bisulcate carapace, M.

hussonae sp. nov. differs from most species of the genus.

It is close to M.? exiqua Styk, 1972 from the Ladinian

(Middle Triassic) of Poland (Styk 1972). However, the

new species is bisulcate, ornamented and DB is not paral-

lel to VB. The new species can also be compared to M.

kozuri Styk, 1972 from the same locality (Styk 1972) but

this species is bisulcate, less elongated, and DB and VB

are not parallel. It differs from M.? buekkensis Kozur,

1985, from the Middle and Late Permian of Hungary

(Kozur 1985), in being shorter, higher, bisulcate, orna-

mented and bearing no denticulation. The new species

also differs from M.? gheorghianae Crasquin-Soleau &

Gradinaru, 1996, from the Early Anisian (Middle Triassic)

of Dobrogea in Romania (Crasquin-Soleau & Gradinaru

1996), in its stockier carapace, PB larger, S2 more

expressed and ornamentation of lateral surface. The

weakly expressed S1 is observed only rarely inMonocera-

tina species and seems to be related to the attachment of

antennulae and antennae (Schornikov 1990).

The size variations of M. hussonae sp. nov. are limited

compared with the other species described here and no

growth stages can be distinguished (Fig. 15). Several

Late Triassic species of Monoceratina are known but only

consist of single specimens with no information on ontog-

eny (e.g. Ye et al. 1977; Wei et al. 1983). The only avail-

able ontogenetic series of Monoceratina species is from

two species from the Maastrichtian (Late Cretaceous) of

Poland (Szczechura 1964): M. umbonata (Williamson,

1848) and M. laevioides similis Szczechura, 1964. The

dimensions of other species from the literature are com-

pared. A compilation of the size distribution of these

Monoceratina species clearly shows that M. hussonae is

much smaller and has more restricted H and L variations.

Monoceratina data reporting only one specimen also show

that M. hussonae is much smaller than other species

(Fig. 16). In this locality, this characteristic is shared at the

superfamilial level and will be further discussed in the

Remarks for Callicythere postiangusta Wei, 1981.


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Figure 14. Ostracods from the C€ur€uk Da�g section, Taurus, Turkey. A–J, Monoceratina hussonae sp. nov.; A, right lateral view of acomplete carapace, P6M3298; B, left lateral view of a complete carapace, P6M3299; C, holotype, right lateral view of a complete cara-pace, P6M3300; D, left lateral view of a complete carapace, P6M3301; E, right lateral view of a complete carapace, P6M3302; F, rightlateral view of a complete carapace, P6M3303; G, right lateral view of a complete carapace, P6M3304; H, paratype, right lateral viewof a complete carapace, P6M3305; I, right lateral view of a complete carapace, P6M3306; J, right lateral view of a complete carapace,P6M3307. K, L, Basslerella sp. 1; K, right lateral view of a complete carapace, P6M3308; L, right lateral view of a complete carapace,P6M3309. M–R, Callicythere postiangusta Wei, 1981; M, left lateral view of a complete carapace, P6M3310; N, right lateral view of acomplete carapace, P6M3311; O, right lateral view of a complete carapace, P6M3312; P, right lateral view of a complete carapace,

16 M. B. Forel

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Family Cytherissinellidae Kashevarova, 1958

Genus CallicythereWei, 1981

Callicythere postiangustaWei, 1981

(Fig. 14M–R)

1981 Callicythere postiangusta Wei: 504, pl. 1, figs 19–22.

2005 Callicythere postiangusta Wei; Crasquin-Soleau &

Kershaw: pl. 1, figs 1–6.

2012 Callicythere postiangusta Wei; Forel: 25, figs 14T–

W, 15A, B.

Occurrence. Samples CD08A7, 08CD03A-C, 03D-G,

04, 05, S1, 08CD07, S2, 08CD09–11, 13, 14, S4, S3, S10,

08CD14’-16, S7, S8, S9, 08CD18, 20, 21–23, 25–30,

C€ur€uk Da�g section (Fig. 3), Western Taurus, Turkey,

Kokarkuyu Formation (this work). Sichuan Province,

South China (Wei 1981). Laolondong section, Sichuan

Province, South China (Crasquin-Soleau & Kershaw

2005). Dajiang section, Daye Formation, Guizhou Prov-

ince, South China (Forel 2012). All occurrences are from

the Griesbachian, Induan, Early Triassic.

Remarks. Figure 17 plots H versus L for all specimens of

Callicythere postiangusta Wei, 1981. As observed for

Monoceratina hussonae sp. nov., the size distribution of

the present species is narrower than all other documented

species. Few species of Callicythere have been docu-

mented in the literature, making comparisons of size pat-

terns difficult. However, since the narrow size range only

affects species of the superfamily Cytheroidea, tapho-

nomic sorting can be excluded. Owing to the fact that

other Monoceratina species do not show this phenome-

non, the observed limitation of size variation is not a gen-

eral superfamilial characteristic but is restricted to the

Cytheroidea of the studied area, in space and time. As a

first approximation, I consider that this conclusion is also

valid for Callicythere. It could be the result of a specific

physiological response to an environmental stress.

Order Palaeocopida Henningsm€oen, 1953Suborder Kirkbyocopina Gr€undel, 1969

Superfamily Kirkbyoidea Ulrich & Bassler, 1906

Family Kirkbyidae Ulrich & Bassler, 1906

Genus Reviya Sohn, 1961

Reviya sylvieae sp. nov.

(Fig. 14S–Z)

Diagnosis. Species of Reviya with ridge joining cardinal

angles and parallel to free margin; surface reticulated.

Derivation of name. Dedicated to Dr Sylvie Crasquin,

CNRS, Universit�e Pierre et Marie Curie, Paris, France.


(heteromorph; Fig. 14S), sample S8. Paratype: P6M3319,

one complete carapace (heteromorph; Fig. 14V), sample

S10. All from the C€ur€uk Da�g section, Western Taurus,

Turkey, Kokarkuyu Formation, Griesbachian, Induan,

Early Triassic (this work). Nineteen complete carapaces,

several fragments.

Occurrence. Samples 08CD03D1, 14, S3, S10, 08CD16,

17, S8, 08CD20–23, C€ur€uk Da�g section (Fig. 3), Western

Taurus, Turkey, Kokarkuyu Formation, Griesbachian,

Induan, Early Triassic (this work).

Description. Kirkbyan pit poorly defined, visible on

some broken specimens. Lateral surface of the valves: sur-

face reticulated; thin ridge parallel to free margin extend-

ing from ACA to PCB.

Heteromorphs (,; Fig. 14S, V): carapace subrectangu-

lar, slightly preplete; Lmax above midH; LV larger than

RV, slight overlap all along the free margin; DB straight

and long, �80–85% of Lmax; cardinal angles sharp,

PCA�100–110�, ACA�120�; PDB and ADB, short,

faintly convex and sharply bent ventrally; AB and PB

rounded with a smaller maximum of curvature at PB; ante-

rior maximum of convexity located around midH, posterior

maximum of convexity slightly above; VB straight, slightly

bent toward AB, �60% of Lmax; ventral angles blunt,

AVB�PVB�130�; AVB longer and more convex than

PVB; lateral surface with occasional thin ridges.

Tecnomorphs (<; Fig. 14T, U, W–Z): carapace subtrian-

gular, highly preplete with Hmax in the anterior one-third

of midL; Lmax above midH; LV larger than RV, slight

overlap all along the free margin; DB long and straight,

slightly shorter than Lmax; cardinal angles well expressed,

ACA �120–140�, PCA �90–110�; VB straight to slightly

convex, sharply bent toward AB; AB large and broadly

rounded, maximum of curvature above midL; PB narrow

finely reticulated with no accessory ridges.

Dimensions (heteromorphs and tecnomorphs). H ¼146–391 mm; L ¼ 220–708 mm; H/L ¼ 0.55–0.72

(Fig. 18).

P6M3313; Q, right lateral view of a complete carapace, P6M3314; R, right lateral view of a complete carapace, P6M3315. S–Z, Reviyasylvieae sp. nov.; S, holotype, right lateral view of a complete carapace, heteromorph, P6M3316; T, left lateral view of a complete cara-pace, tecnomorph, P6M3317; U, left lateral view of a complete carapace, tecnomorph, P6M3318; V, paratype, right lateral view of acomplete carapace, heteromorph, P6M3319; W, right lateral view of a complete carapace, tecnomorph, P6M3320; X, right lateral viewof a complete carapace, tecnomorph, P6M3321; Y, right lateral view of a complete carapace, tecnomorph, P6M3322; Z, left lateral viewof a complete carapace, tecnomorph, P6M3323. A0–B0, Reviya curukensis Crasquin-Soleau, 2004b; A0, left lateral view of a completecarapace, P6M3324; B0, right lateral view of a complete carapace, P6M3325. Scale bars are 100 mm.

3


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Figure 15. Length/height scatter plot for Monoceratina hussonae sp. nov. All specimens are from C€ur€uk Da�g section (Taurus, Turkey)and are at the same scale.

Figure 16. Comparative height/length distributions for species of Monoceratina: M. hussonae sp. nov.; M. umbonata (Williamson,1848) (Szczechura, 1964; Late Cretaceous, Maastrichtian of Poland); M. laevioides similis Szczechura, 1964 (Late Cretaceous, Maas-trichtian of Poland); M.? gheorghianae Crasquin-Soleau & Gradinaru, 1996 (Middle Triassic, Early Anisian of Romania; Crasquin-Sol-eau & Gradinaru 1996 and Sebe et al. in 2013); 1, M. scrobiculata Triebel & Bartenstein, 1938 (Bate et al. 1979; Early Jurassic,Sinemurian of leg ODP79, site 547); 2, M. vulsa (Jones & Sherborn, 1888) (Bate et al. 1979; Early Jurassic, Pliensbachian of legODP79, site 547); 3, ?M. sp. (Bate et al. 1979; Early Jurassic, Pliensbachian of leg ODP79, site 547); 4, M. subscaphodea (Huang &Gou, 1977 in Ye et al. 1977; Late Triassic, Sichuan Province, China); 5,M. subtriangularis (Huang & Gou, 1977 in Ye et al. 1977; LateTriassic, Yunnan Province, China); 6,M. scaphodea (Huang & Gou, 1977 in Ye et al. 1977; Late Triassic, Yunnan Province, China).

18 M. B. Forel

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Remarks. Reviya sylvieae sp. nov. is close to R. curuken-

sis Crasquin-Soleau, 2004a from the Induan (Early Trias-

sic) of the C€ur€uk Da�g section (Crasquin-Soleau et al.

2004a, b), but displays an adventral ridge. Three ontoge-

netic stages are identified for the new species, labelled A-

2, A-1 and Adults (Fig. 18).

Reviya curukensis Crasquin-Soleau, 2004

(Fig. 14A0–B0)

2002 Reviya? sp. 1. Crasquin-Soleau et al.: 493,

fig. 4.6–4.8.

2004a Reviya curukensis Crasquin-Soleau in Crasquin-


2004b Reviya curukensis. Crasquin-Soleau in Crasquin-

Soleau et al.; Crasquin-Soleau et al.: 282, pl. 1, figs 5–7.

2011 Reviya curukensis Crasquin-Soleau in Crasquin-

Soleau et al.; Forel & Crasquin: 463, fig. 4E–H.

Figure 17. Length/height scatter plot for Callicythere postiangusta. Specimens come from C€ur€uk Da�g (this work), Dajiang, Guizhou,China (Forel 2012), Laolongdong, Sichuan, China (Crasquin-Soleau & Kershaw 2005) and Sichuan, China (Wei 1981). All specimensare at the same scale.

Figure 18. Length/height scatter plot for Reviya sylvieae sp. nov. Specimens are from C€ur€uk Da�g section (Taurus, Turkey) and are atthe same scale.


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2013a Reviya curukensis Crasquin-Soleau in Crasquin-

Soleau et al.; Forel et al.: fig. 4U, V.

Occurrence. Samples S7, TK55–58, TK117–119, 121,

122, C€ur€uk Da�g section (Fig. 3), Western Taurus, Turkey,

Kokarkuyu Formation (this work; Crasquin-Soleau et al.

2002, 2004a, b). Meishan section, Zhejiang Province,

South China (Forel & Crasquin 2011), B�alv�any North sec-tion, Gerennav�ar Formation, B€ukk Mountains, Hungary,

Griesbachian (Forel et al. 2013a). All occurrences are

from the Griesbachian, Induan, Early Triassic.

Remarks. Figure 19 plots H versus L for all specimens of

Reviya curukensis. They cluster into four ontogenetic stages,

A-3, A-2, A-1 and Adults. The shape of this species is rela-

tively conservative through ontogeny and the most visible

morphological differences are related to sexual dimorphism.

Suborder Kloedenellocopina Scott, 1961 emend.

Lethiers, 1978

Superfamily Kloedenelloidea Ulrich & Bassler,

1908

Family Kloedenelloidea Ulrich & Bassler, 1908

Genus Eukloedenella Ulrich & Bassler, 1923

emend. Lethiers, 1981

Eukloedenella adcapitisdolorella sp. nov.

(Fig. 20A–H)

Diagnosis. A species of Eukloedenelella with small cara-

pace for the genus; DB and VB straight and parallel to

subparallel; ACA > PCA; S2 faint; surface homo-

geneously punctuated.

Derivation of name. From the Latin ad capitis dolores

for the headache the author suffered when working on the

taxonomy of this species.


(heteromorph; Fig. 20A), sample S6A. Paratype:

P6M3327, one complete carapace (tecnomorph;

Fig. 20B), sample S6A. All from the C€ur€uk Da�g section

(Fig. 3), Western Taurus, Turkey, Kokarkuyu Formation,

Griesbachian, Induan, Early Triassic (this work). Twenty

complete carapaces, several fragments.

Occurrence. Samples S6A, 08CD51, 78, C€ur€uk Da�g sec-tion (Fig. 3), Western Taurus, Turkey, Kokarkuyu Forma-

tion, Griesbachian, Induan, Early Triassic (this work).

Description. Lmax located around midH; lateral surface

of the valves finely and homogeneously punctuated; faint

S2 in front of midL in the upper 1/4 of H (Fig. 20A, D–G);

thin overlap of LV over RV sometimes perceptible along

free margin (Fig. 20C, D).

Heteromorphs (,; Fig. 20A, G): carapace of parallelo-

gram shape, amplete with H�Hmax all along DB; DB

parallel to VB, �40–45% of Lmax, straight with slight

invagination around midL marking S2; ACA and PCA

more rounded and larger than at tecnomorphs, PCA

�120�, ACA �140�; PDB long and convex, representing

Figure 19. Length/height scatter plot for Reviya curukensis. Specimens are from C€ur€uk Da�g (this work and Crasquin-Soleau et al.2004a), B�alv�any North, Hungary (Forel et al. 2013a), Dajiang, Guizhou, China (Forel 2012), and Meishan, Zhejiang, China (Forel &Crasquin 2011). All specimens are at the same scale.

20 M. B. Forel

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� two-thirds of Hmax; PB more rounded and prominent

than at tecnomorphs, with a narrower radius of curvature,

maximum located slightly above midH; ADB strongly

convex and long, being two-thirds of Hmax; AB rounded

with a narrower radius of curvature, maximum below

midH, �40% of Hmax; VB somewhat shorter than DB,

straight to slightly concave at middle part; PVA �110–

120�, AVA �140–150�; PVB long, convex and parallel to

ADB; AVB short parallel to PDB; sexual dimorphism

strong and atypical on the entire ventral half of the cara-

pace, becoming larger and massive with rectangular out-

line, lowering of the anterior maximum of convexity

resulting in a ventrally twisted aspect of the anterior por-

tion of the carapace.

Tecnomorphs (<; Fig. 20B–F, H): carapace subovate,

slightly tapered posteriorly, preplete, Hmax around ante-

rior one-third of Lmax; DB straight to slightly convex,

long, about 70% of Lmax; ACA and PCA relatively sharp,

PCA > 90�, ACA �130�; PDB short; PB rounded with

maximum of curvature located in the upper third of

Hmax; ADB gently convex and long representing 40–

50% of Hmax; AB rounded with a large radius of convex-

ity, maximum located around midH; VB straight to

slightly convex, parallel to subparallel to DB, slightly

bent toward AB; AVA�PVA�150�; AVB and PVB

rounded; PVB narrower and longer than AVB, �65% of

Hmax.

Dimensions. L ¼ 513–795 mm; H ¼ 294–483 mm;

H/L ¼ 0.55–0.68 (Fig. 21).

Remarks. Until now, only one genus of the superfamily

Kloedenelloidea was known from the Early Triassic

(Langdaia, family Knoxitidae, see Crasquin & Forel

2013 for details). The last occurrence of the genus

Eukloedenella was reported, without illustration, from

the Late Permian of Western Guizhou and Eastern

Yunnan, South China (Yao et al. 1980). Eukloedenella

adcapitisdolorella sp. nov. is the first lower Triassic

occurrence of the genus and of the family Kloedenelli-

dae, which are therefore recognized for the first time

through the PTB.

The 19 Eukloedenella species originally described

from Silurian deposits were gathered into five groups

based on the surface markings of their valves (Ulrich &

Bassler 1923). Although S2 is faint in E. adcapitisdolor-

ella sp. nov., this excludes it from the E. indivisa group.

Because it displays no projecting AV flange, and no

swollen or anterior third segregated by ventral depres-

sion, it cannot be assigned to the E. sinuata or E. bulbosa

groups. The homogeneous convexity of the carapace

Figure 20. Eukloedenella adcapitisdolorella sp. nov. from the C€ur€uk Da�g section, Taurus, Turkey. A, holotype, right lateral view of acomplete carapace, heteromorph, P6M3326; B, paratype, left lateral view of a complete carapace, tecnomorph, P6M3327; C, right lat-eral view of a complete carapace, tecnomorph, P6M3328; D, right lateral view of a complete carapace, tecnomorph, P6M3329; E, rightlateral view of a complete carapace, tecnomorph, P6M3330; F, right lateral view of a complete carapace, tecnomorph, P6M3331; G, leftlateral view of a complete carapace, heteromorph, P6M3332; H, right lateral view of a complete carapace, tecnomorph, P6M3323. Scalebars are 100 mm.


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relates the new species to the E. umbilicata group. How-

ever, all species assigned to this group show a more

depressed S2 and E. adcapitisdolorella sp. nov. seems to

be a transitional form between non-sulcate species of the

E. indivisa group and the clearly monosulcate species

belonging to the E. sinuata group. Eukloedenella adcapi-

tisdolorella sp. nov. is distinguished from all known

Eukloedenella species by the weakness of its S2 and its

punctuated ornamentation. In an evolutionary context, it

is puzzling that this transitional type corresponds to pos-

sibly the last occurrence of the group in the Early Trias-

sic. This paradox could be explained by the lack of data

or poor fossil record. A phylogenetic analysis of all

known specimens of Euckloedenella would help clarify

the question and determine whether the five groups of

Ulrich & Bassler (1923) have any real phylogenetic

meaning.

The observed unusual sexual dimorphism affecting the

entire ventral part of the carapace seems to indicate that

the brooding chamber was not confined to the PV part but

relatively long, extending all along VB.

Discussion

Reconstructions of ontogeny and growth ratesThe shapes of the carapaces of ostracods can be defined as

a function of three major parameters: ontogenetic stage,

sexual dimorphism and intraspecific variability. In this

paper, variations of shape attributable to ontogeny are

described and quantified. Intraspecific variations will be

discussed in detail in a forthcoming analysis. Sexual

dimorphism is well known and expressed in the carapaces

of most Palaeocopida, often observed in Recent

Bairdiidae (e.g. Maddocks & Iliffe 1986; Maddocks 1991;

Maddocks 2013) and is here recorded for the second time

in fossil Bairdia (Bairdia? kemerensis). Sexual dimor-

phism is related to the presence of breeding chambers in

females (e.g. palaeocopids, platycopids) and of sexually

dimorphic appendages. In the fossil record, the morpho-

logical characteristics of sexual dimorphism in the cara-

pace have been observed as early as A-3 in various

palaeocopid species (e.g. Martinsson 1956; Jaanusson

1956; Guber 1971; Whatley & Stephens 1977). This pre-

cocious dimorphism is related to the appearance of anla-

gen of sexually dimorphic appendages, not yet fully

functional (e.g. Smith & Kamiya 2002), and sexual matu-

rity is acquired during the final moult to adult stage. For

the present discussion, I exclude Cytheroidea for which

no larval stages were distinguished (Callicythere postian-

gusta and Monoceratina hussonae). Consequently, seven

species are considered: Bairdia? kemerensis, Liuzhinia

antalyaensis, Praezabythocypris? ottomanensis, Paracyp-

ris gaetanii (podocopids), Reviya curukensis, R. sylvieae

and Eukloedenella adcapitisdolorella (palaeocopids).

Analysis of ontogeny and associated shape variations

entails: (i) ‘qualification’ of growth patterns with eventual

heterogeneities at different zones of the carapaces; and

(ii) ‘quantification’ of growth from H versus L diagrams.

Podocopids typically have 8 or 9 instars from egg to

adult (e.g. Cohen & Morin 1990; Smith & Kamiya 2002;

Mesquita-Joanes et al. 2012). Therefore, although clusters

of successive juveniles and adults have been described

above (Figs 5, 9, 10, 13, 15, 17, 18–20), ontogenies seem

truncated at the base with at least three stages missing:

smaller juveniles of the first stages might have been

removed taphonomically or lost during washing and siev-

ing. In the absence of internal features of carapaces, the

Figure 21. Length/height scatter plot for Eukloedenella adcapitisdolorella sp. nov. All specimens are from the C€ur€uk Da�g section (Tau-rus, Turkey) and are at the same scale.

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cluster of the largest specimens of each reconstructed line-

age are considered to be adults.

Variation in shape through ontogeny. Shape variations

through ontogeny are studied by comparison of the shapes

of individuals of each growth stage in the seven selected

species (Fig. 22). Bairdia? kemerensis (Fig. 22A) shows a

relatively conservative contour. The dominant modifica-

tions are located at both extremities, with the progressive

upward migration of the posterior and anterior maxima of

convexity. This pattern results in an overall change of the

lateral outline of the carapace: from diamond-shaped in

A-4 (Lmax below Hmax) to close to rectangular in adults

(Lmax around Hmax). Liuzhinia antalyaensis and Praeza-

bythocypris? ottomanensis exhibit stronger shape modifi-

cations distributed all around the carapace. The latter

(Fig. 22B) shows enlargement of PB with the upward

migration of the maximal convexity, an increase of round-

ness at AB with a downward shift of the maximum con-

vexity, differentiation of an anteriorly inclined ADB, and

the appearance of convexity at VB. These morphological

changes mostly occur between A-3 and A-2; later stages

only showing strengthening of these characteristics. Liuz-

hinia antalyaensis (Fig. 22C) displays a relatively conser-

vative VB, becoming flatter and longer through ontogeny.

Most of the changes are located in the upper part of the

carapace, which becomes more symmetrical with wider

PB and more constant Hmax. AB becomes wider and

more rounded. Compared to Praezabythocypris? ottoma-

nensis, the observed changes are more gradual and homo-

geneously distributed amongst the successive larval

stages.

Figure 22. Mean shapes of each instar (right valves only) reconstructed for A, Bairdia? kemerensis, B, Praezabythocypris? ottomanensis,C, Liuzhinia antalyaensis, D, Paracypris gaetanii, E, Reviya sylvieae, F, R. curukensis and G, Eukloedenella adcapitisdolorella.


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The adult contour of Paracypris gaetanii has not been

represented because only left valves are available

(Fig. 22D). This species displays the most conservative

shape from A-4 to A-1: the shape of stage A-(nþ1) glob-

ally appears as an enlargement of A-n. Shape modifica-

tions mostly affect the roundness of the extremities, PB

becoming narrower and AB becoming larger. Therefore,

the dominant pattern of ontogeny for Paracypris gaetanii

is size increase, with secondary shape changes located at

both AB and PB. Most changes observed through the

ontogeny of Reviya sylvieae (Fig. 22E) affect the ventral

and posterior portions of the carapace, DB and AB being

very conservative. PB becomes larger and more rounded,

with the maximum of curvature migrating ventrally, while

VB becomes longer, straighter and parallel to DB. Reviya

curukensis displays slight modifications, with PB enlarg-

ing and VB lengthening to become parallel to DB

(Fig. 22F). The last moult from A-1 to Adult records very

different patterns between these two species: a high

increase of size for R. sylvieae but a small one for R.

curukensis.

Modifications of Eukloedenella adcapitisdolorella

(Fig. 22G) are observed mostly in PB, VB and AB. The

patterns are similar but more subtle than those described

for the species of Reviya: PB more rounded with the maxi-

mum of curvature migrating ventrally, elongation and hor-

izontalization of VB, less marked ACA, and AB

protruding anteriorly. In the three palaeocopids studied

here, DB shape is very conservative through ontogeny,

being only affected by strict elongation.

Growth rates. Brooks’s rule states that the volume of

crustaceans should double with each moult and linear

dimensions should consequently increase by 1.26 (Brooks

1886; Teissier 1960). This rule is a generalization of

growth and exceptions may provide insights into biology,

ontogeny and life cycle. Although processes accounting

for this pattern are still unknown, the growth of ostracods

follows Brooks’s rule with some amount of variability

(e.g. Hessland 1949; Sohn 1950; Kesling 1951a, b, 1952,

1953; Shaver 1953; Kesling & Takagi 1961; Kesling &

Crafts 1962). In their review, Danielopol et al. (2008)

documented values ranging from 1.21 to 1.37 for non-

marine species. Such information has not been available

previously for marine species.

The match to Brooks’s rule can be evaluated by com-

paring the observed and predicted growth rates. For the

following discussion, only specimens from the C€ur€uk Da�gsection are considered based on this work and Crasquin-

Soleau et al. (2002, 2004a, b). Lmean and Hmean for all

specimens of each ontogenetic stage of Bairdia? kemeren-

sis, Liuzhinia antalyaensis, Praezabythocypris? ottoma-

nensis, Paracypris gaetanii, Reviya sylvieae, R.

curukensis and Eukloedenella adcapitisdolorella were

determined. Growth rates of Hmean and Lmean were

calculated for each ontogenetic transition in the following

way (Fig. 23):

KH ¼ Hnþ1=Hn

KL ¼ Lnþ1=Ln;

where KH and KL are respectively Hmean and Lmean

growth rates; Hn and Ln are respectively Hmean and

Lmean at ontogenetic stage n; and Hnþ1 and Lnþ1 are

respectively Hmean and Lmean at ontogenetic stage nþ1.

For most of the species, KH and KL are generally lower

than predicted by Brooks’s rule and diminish through

ontogeny. Values range from 1.07 to 1.75 for KH, and

from 1.09 to 2 for KL. They record a decelerating and

sluggish growth, with an exception for the species Reviya

sylvieae. The slowest growth in length and height is

recorded in Bairdia? kemerensis and Eukloedenella adca-

pitisdolorella respectively. The growth of all podocopid

species displays the same decreasing pattern through

time: KH and KL decrease strongly to A-2, KH then stays

stable while KL increases before Adult stage. KH in Liuz-

hinia antalyaensis shows a constant decrease, while KL is

stable until A-2 and then decreases sharply. Although

only two transitions are considered for Paracypris gaeta-

nii, the same strong decreasing trend is recognized for

both KH and KL. All growth values for Bairdia? kemeren-

sis, Liuzhinia antalyaensis and Paracypris gaetanii are

below values predicted from Brooks’s rule. Praezabytho-

cypris? ottomanensis displays the only growth event

above Brooks’s rule, at the transition A-3 to A-2. It is then

followed by a sharp decrease and a slight increase before

Adult stage. For each ontogenetic transition in all species,

KL > KH: growth in length is greater and faster than in

height. Palaeocopid growth differs amongst the three spe-

cies under scrutiny and from podocopids. KH and KL of

the two Reviya species are below that predicted from

Brooks’s rule and anti-correlated: (i) from A-3 to A-1, KH

decreases for R. sylvieae but increases for R. curukensis,

while KL increases for R. sylvieae but decreases for R.

curukensis; (ii) from A-2 to Adult, KH increases for R. syl-

vieae but decreases for R. curukensis, while KL decreases

for R. sylvieae but increases for R. curukensis. However,

the general trend recognized for these two species is a

decrease in both KH and KL through ontogeny with values

below those predicted from Brooks’s rule. Eukloedenella

adcapitisdolorella shows a different and striking develop-

ment, with strong growth from A-2 to Adult for both KH

and KL, from values below the Brook’s rule prediction for

A-2 to A-1, to values far above it for A-1 to Adult.

Although early growth stages are missing, the observed

increase during final moulting is unusual and previously

unknown for any other ostracods.

Heterochrony. Heterochronies are generally defined as

changes in the timing and/or rates of processes underlying

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the ontogenetic formation of morphological traits (Gould

1977; Raff & Kaufman 1983; McKinney & McNamara

1991; Reilly et al. 1997). They involve three developmen-

tal parameters: rate of shape development, and onset and

offset times of characters. Perturbations of these parame-

ters can produce a change in the ontogenetic timing

relative to the ancestral conditions and three types of het-

erochrony are possible: (i) paedomorphosis, for which the

ontogeny of the studied character is truncated relative to

the ancestral species; (ii) peramorphosis, for which the

ontogeny is extended compared to the ancestral species;

and (iii) isomorphosis, when the character is the same in

Figure 23. Growth ratio of Hmean (upper) and Lmean (lower) of Bairdia? kemerensis, Praezabythocypris? ottomanensis, Liuzhinia ant-alyaensis, Paracypris gaetanii, Reviya sylvieae, R. curukensis and Eukloedenella adcapitisdolorella at each recognized phase of growth.The dashed line indicates a growth ratio of 1.26, predicted by Brooks’s rule.


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the ancestor and descendant but the descendant has fol-

lowed a different ontogenetic trajectory to reach the same

shape. Paedomorphosis can be produced by deceleration

(slower rate), hypomorphosis (earlier offset time) or post-

displacement (later onset time). Peramorphosis can result

from acceleration (faster rate), hypermorphosis (later off-

set time) or pre-displacement (earlier onset time) (see

Reilly et al. 1997 for further details). Harries et al. (1996)

identified paedomorphosis as an adaptative mechanism to

survive ecological and environmental perturbations asso-

ciated with mass extinctions, with examples of progenetic

organisms (hypomorphotic according to Reilly et al.

1997). Moderate thermal fluctuations might be associated

with fast growth, while the retardation of development

was described in fluctuating regimes encompassing

extreme temperatures (Thorp & Wineriter 1981; Worner

1992). Deceleration is known for molluscs under the influ-

ence of warming that led to nutrient-depleted waters dur-

ing the Miocene (Sch€one et al. 2004), while acceleration

in fluctuating temperatures has been reported for Daphnia

(Orcutt & Porter 1983) and rotifers (Halbach 1973). Pro-

genesis (e.g. hypomorphosis) and post-displacement are

shown in Recent ostracods of the continental shelf of the

Congo and Senegal and are related to strong upwelling

and increased sedimentation rates (Bertholon 1997). Pera-

morphic and paedomorphic evolution have been impli-

cated as speciation mechanisms in Loxoconcha species

during the Pliocene–Pleistocene of the Western Pacific

(Tanaka & Ikeya 2002). Deceleration of growth rates has

been observed for several Cypridoidea (podocopids) at

low temperatures (Ganning 1971). Despite these exam-

ples, data are more abundant for freshwater ostracods, for

which it is known that salinity modulates hatching phenol-

ogy, survival and moulting, too low conductivity having a

negative effect on ostracod survival and growth (De Deck-

ker 1983; Mezquita et al. 1999). The final moulting of the

Recent Heterocypris barbara is delayed and mean size of

the adult stage decreases with lowered salinity (Rossi

et al. 2013). On the other hand, many species display

faster developmental time at higher temperatures and at

optimal salinity (Cohen & Morin 1990).

A key element in any discussion about heterochrony is

the definition of the terminal shape of the studied parame-

ter. Because the present discussion focuses on the general

ontogenetic trajectories of ostracods, the terminal shape

should be their shape at sexual maturity, i.e. at the adult

stage. The slow growth of podocopids and species of

Reviya could therefore be related to paedomorphosis by

deceleration or isomorphosis by deceleration and hyper-

morphosis, while the fast development of Eukloedenella

could result from peramorphosis by acceleration or iso-

morphosis by acceleration and hypomorphosis. Conse-

quently these patterns might result, respectively, from

paedomorphic growth by deceleration and peramorphic

growth by acceleration. As a first approximation, the

growth discrepancies between podocopids/Reviya and

Eukloedenella seem to be related to their different palae-

obiology. Deceleration of the growth of podocopids/

Reviya at A-3 or A-2 indicates that their optimal resilience

should have been reached at these stages. In contrast, the

strong acceleration of Eukloedenella growth to reach adult

stage seems to indicate that it was the most resilient for

this species.

Ostracods surviving to the EPE in association with the

PTBM (Permian-Triassic boundary microbialites) of the

C€ur€uk Da�g section are characterized by: (i) proliferation

of specimens; (ii) abundant small-sized individuals; and

(iii) developmental heterochronies for at least seven spe-

cies. Although they clearly record persistence of environ-

mental stress, how did such faunas maintain their viability

throughout the deposition of microbial deposits? To

address this problem, the way an organism acquires

energy from its environment and allocates it to different

biological processes should first be discussed. Organisms

extract resources from the environment, and use them for

three basic functions: somatic maintenance, growth and

reproduction. For a fixed amount of resource, there is a

trade-off between these functions (Roff 1992), depending

on: (i) the state of the organism, i.e. body volume

(described by size-related phenomena such as food uptake

and growth), energy reserves and ageing process; and (ii)

the environment (e.g. Lika & Kooijman 2003). Those

resources allocated to reproduction can sustain either the

production of numerous small eggs or a few large ones

(e.g. Oloffson et al. 2009). Life history theory is con-

cerned with the way individuals allocate resources to

growth and reproduction, based on the assumptions that

resources are limited and that physiological constraints act

on growth and reproduction (Roff 1992; Stearns 1992).

Ostracods do not exhibit simultaneous growth and

reproduction, except in cases of precocious sexual matu-

rity. Their reproductive effort is set to 0 until a certain age

at which growth ceases and energy is switched to repro-

duction (Perrin 1992; Bulmer 1994). As a consequence,

energy resources of eggs and juveniles are entirely dedi-

cated to maturation and conservation of the achieved

degree of maturity, while they are allocated to mainte-

nance and reproduction in adults (Lika & Kooijman

2003). There are few papers on the life history strategies

of marine ostracods but most of these organisms appear to

be iteroparous, with potentially more than one reproduc-

tive event during the adulthood of the females. It is inter-

esting to estimate how many eggs an adult female can

produce under normal and optimal conditions. To do so,

three parameters are important: the number of reproduc-

tive events per year, the number of eggs per clutch and the

lifetime of adult females. Since Bairdioidea dominate the

studied PTBM assemblages and the life cycles of

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palaeocopids are relatively unknown, I will only consider

Bairdioidea. Data on longevity, breeding cycles and num-

ber of generations per year (voltinism) is scattered for

Bairdioidea, but the literature does contain some informa-

tion on other marine podocopids. Cytheroidea with sea-

sonal reproduction produce from one to four generations

per year (Horne 1983). The number of eggs per clutch is

also poorly known but is 11–50 in Cytheroidea (Cohen &

Morin 1990) and 10 to more than 20 in Cypridoidea (Mad-

docks 1990). Data for Bairdioidea are rare but Maddocks

(2013, fig. 9D, E) shows at least eight eggs of Neonesidea

tenera (Brady) in uteri, while N. oligodentata lays eggs

either singly or in groups of 7–10 (Obato 1999). Adult life-

time is known to last from 7 days to 3 months in Cytheroi-

dea, and from 1.5 to 7 months in Cypridoidea (Cohen &

Morin 1990). In the absence of precise data for Bairdioi-

dea, mean values of data observed for other podocopids

(Cypridoidea, Cytheroidea) can be used to estimate the

fecundity of adult female Bairdioidea. Considering a life-

time of 7 months, 1–4 reproductive events per year (¼ 1–2

per lifetime), 7–10 eggs per clutch, then the fecundity of a

Bairdioidea female would be 7–20 eggs in a lifetime under

optimal environmental and ecological conditions.

The observed reduction of the body size of ostracods in

the PTBM implies a reduction of the space potentially

available for eggs. To cope with this phenomenon, two

adaptations of the production of eggs are conceivable,

known as bet-hedging: (i) the same quantity of smaller

size eggs, or (ii) fewer eggs of a larger size. They have

been reported frequently and studied for all kinds of

organisms, including arthropods (Fox & Czesak 2000).

Bet-hedging strategy has been evoked to describe the

asynchronous hatching of resting eggs in populations of

freshwater Eucypris virens (Martins et al. 2008). Modifi-

cations of the production of eggs in marine ostracods

were discussed by Heip (1976), without quantification, for

the Recent Cyprideis torosa. This author considered that

in stable and predictable environments, the total number

of eggs will be reduced and that under unstable and/or del-

eterious conditions, reducing the number of eggs would be

catastrophic. According to recent observations and mod-

els, both strategies are used by arthropods, although larger

eggs seem more suitable in unstable settings (Fox &

Czesak 2000; Oloffson et al. 2009). Such information is

lacking for ostracods; however, reduction in the size of

these post-extinction ostracods seems incompatible with

larger eggs. Hence, I consider here the production of

smaller and more abundant eggs as more suitable in the

PTBM setting following the EPE.

The persistence of stable populations implies that mor-

tality and birth rates are balanced, reproductive patterns

being linked to mortality levels. Following the EPE in the

C€ur€uk Da�g section, mortality might have been higher than

normal, with two consequences: (i) higher reproductive

rates; and (ii) reduction of the time before females begin

to reproduce. Time to reproduction is further decreased

for podocopids by deceleration, resulting in a longer time

spent for the maturation of the juveniles and the shorten-

ing of adulthood duration. In these conditions, the conclu-

sion of smaller and more abundant eggs laid by PTBM

ostracods is reinforced: populations could not have been

stable with the conjunction of less abundant larger eggs

and reduced time available for reproduction. In this bet-

hedging strategy, the augmentation of the abundance of

eggs produced has never been quantified. If it is hypothe-

sized that production doubled in PTBM ostracods, the

fecundity of ostracod females should have been raised to

14–40 eggs per lifetime to maintain population viability

throughout PTBM deposition. This doubling of reproduc-

tion is fairly probable since it has been observed under

fluctuating temperatures for the Recent freshwater Heter-

ocypris virens (Rossi et al. 2013).

For freshwater ostracods, the hypothesis has been made

that more generations in a given period would allow faster

adaptation (Rossi et al. 2003, 2013; Forrest & Miller-

Rushing 2010). Together with the augmentation of the

number of eggs produced in the PTBM, increase in the

frequency of reproductive events during the adult stage is

also envisaged. Temperature, food availability and sea-

sonal time constraints regulate the duration of the life

cycle in copepods that can alternate between univoltine or

multivoltine strategies (Allan & Goulden 1980; Chen &

Folt 1996; Twombly et al. 1998; Gerten & Adrian 2002;

Winder & Schindler 2004; Adrian et al. 2009). Salinity,

temperature and day length have a great impact on freshwa-

ter ostracod reproduction, which can be reduced in low sal-

inities and doubled under fluctuating temperatures (Ganning

1971; Hagerman 1978; Otero et al. 1998; Rossi et al. 2013).

As a consequence of these factors, it is obvious that

Bairdioidea have greater adaptive potential than is tradi-

tionally considered. The faunas of the C€ur€uk Da�g section

cannot be taken as surviving in the PTBM refuge in a

straightforward and unique way. Instead they exemplify

two fundamentally different types of adaptation, poten-

tially opening new questions on the palaeobiology of

podocopids and palaeocopids. The study of ontogenetical

development of other species from other PTBM will pro-

vide additional analogues. Regarding the deceleration

observed in podocopid species, this can result in dwarf-

ism when it co-occurs with decoupled size and shape

change so that the descendant completes the same degree

and manner of shape change as the ancestor but in a nar-

rower size range (Alberch et al. 1979). In contrast, pera-

morphosis by deceleration could result in gigantism.

These phenomena could explain the overall reduction in

the size of ostracods after the EPE (Forel et al. in prog-

ress) as well as being consistent with observations that

some species become smaller and others larger in the


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PTBM. It could also explain the anomalous size records

for Monoceratina hussonae and Callicythere postiangusta.

However, because no larval stage has been distinguished

for these two species, many questions are still pending: did

growth stop before terminal shape? And did they display a

very fast growth, or was it very slow? To answer these

questions, more data are required.

Ostracod assemblages

Diversity and abundance. Lower Triassic ostracods

from the C€ur€uk Da�g section are found at the base of the

Kokarkuyu Formation, which is of microbial origin, and

are very rare higher up. Species richness and abundance of

assemblages through the section can be divided into peaks

(P) and drops (D) (Fig. 24A). For clarity and because the

upper part of the section did not yield any ostracods, I

report only diversity variations up to sample CD31 (the

first 18 metres), in association with the different fabrics of

microbialites observed and described by Kershaw et al.

(2012). From the base to the top:

1. Assemblages TK51b to CD05 record increasing

richness and abundance (P1). Maximum abundance

is reached in CD05 with 1435 specimens. This inter-

val is mostly dominated by stromatolites and throm-

bolites at the top.

2. An important reduction in diversity and abundance

is observed from CD06 to CD08 (D1). This corre-

sponds to thrombolite at the base and hybrid micro-

bialites at the top.

3. Samples S2 to TK55b yielded rich and abundant

assemblages (P2), corresponding to thrombolites at

the base and hybrid microbialites at the top.

4. A diversity drop is recorded by assemblages CD12

to CD13 (D2). This event is correlated with a shift

to facies dominated by thrombolites.

5. This phase is followed by an important diversifica-

tion from CD14 to CD16 (P3). The maximum rich-

ness is recorded in S10, with 20 species.

6. Assemblages CD17 to CD25 record a long phase of

low richness and abundance (D3).

7. The interval from CD26 to CD29 records a slight

rediversification of assemblages characterized by

relatively higher richness and abundance (P4).

8. From TK125 onwards, samples are barren of ostra-

cods (D4). Only two low diversity and abundance

assemblages are found in CD51 and CD78.

Composition. The ostracods from the C€ur€uk Da�g section

belong to six superfamilies/families, the distributions of

which are shown in Figure 24B. The most common super-

family is Bairdioidea (Acratia, Bairdia, Bairdiacypris,

Bythocypris, Liuzhinia, Microcheilinella, Petasobairdia,

Praezabythocypris, Rectobairdia and Silenites). When

present, it comprises between 25% (CD03A, S1) and

100% of the assemblages (02TK51b, CD02, 02TK120,

123–126). The second most frequent family is Kirkbyidae

(Reviya), absent from specific peaks and drops (P3, D4

and P6). When present, Kirkbyidae varies between 5%

(S10) and 100% (02TK117, 122) of the recovered assemb-

lages. When present, Cytheroidea (Basslerella, Callicy-

there and Monoceratina) comprises between 6% (S4) and

100% (CD03B, 07) of the assemblages. Cypridoidea (gen-

era Macrocypris, Paracypris) are episodic components of

all diversity phases, making up between 5% (S10) and

17% of species (CD03E). Three superfamilies/families

have a unique occurrence in D3: (i) Kloedenelloidea

(genus Eukloedenella) – 50% of 08S6A, CD51, CD78

species; (ii) Cavellinidae (genus Sulcella) and (iii) Para-

parchitidae (Paraparchitidae indet.), each being 33% of

the 02TK118 assemblage.

Abundant small-sized specimens are found in some

assemblages, as in other PTBM deposits in South China

and Hungary (Forel 2012; Forel et al. 2013a). These small

specimens belong to the same genera as the larger forms:

the observed deviation of the size is not due to the pres-

ence of smaller genera per se. This aspect of ostracod fau-

nas associated with microbial deposits in the aftermath of

the EPE is the subject of a work in progress and will only

be discussed briefly below.

Palaeoenvironmental implications. In the present sec-

tion, three major units can be recognized based on both

sedimentology and composition of the ostracod faunas

(Table 1; Fig. 24B). The base of the section (up to �5 metres) is composed of stromatolites and hybrid micro-

bialites and is mostly dominated by Bairdioidea, Bythocy-

theridae and Cytherissinellidae. The middle portion of the

section (� 5 to 13.5 metres) records important facies and

faunal shifts: (i) facies shift to thrombolite deposits; and

(ii) faunal shift with the appearance and gradual increase

of Kirkbyidae. Important modifications also occur at the

top (from 13.5 to 18 metres): (i) transition from thrombo-

lites to oolites; and (ii) complete disappearance of Kirk-

byidae, together with the appearance of Cypridoidae and

the return of Cytherissinellidae. These patterns record the

forcing of environmental parameter(s) on the type of

deposits and composition of the faunas. Compared to

other PTBM (e.g. Forel 2009, 2012; Forel et al. 2013a),

faunas from the C€ur€uk Da�g section are atypical because

of the presence and abundance of Kirkbyidae, Cytherissi-

nellidae and Bythocytheridae. However, knowledge about

these last two families is still limited. Since they are key

elements of many assemblages recovered here, it would

be useful to give precise conclusions about parameters

forcing the observed patterns. A work based on the litera-

ture and new data is in progress to provide information on

Cytherissinellidae and Bythocytheridae that will allow the

28 M. B. Forel

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assessment of detailed environmental changes and pro-

pose palaeogeographical reconstructions.

One conclusion concerns oxygen levels at the base of

the water column. The oxygenation level of seawater is a

major challenge in understanding the mechanisms of EPE

and subsequent recovery patterns (see Payne & Clapham

2012 for a recent scenario). It has also been of great con-

cern in discussions of parameters controlling microbial

growth in devastated environments following extinction

(e.g. Kershaw et al. 2007). Ostracod faunas have tradition-

ally been used as tools to estimate quantitatively past oxy-

gen levels in marine environments (Lethiers & Whatley

1994). However, evidence from both the Recent and the

fossil record has called this tool into question (e.g.

Brand~ao & Horne 2009; Horne et al. 2011; Forel 2013),

although it is still possible to obtain a qualitative idea of

the oxygenation state of waters based on the ecology of

the superfamilies/families present. As stated above, most

of the assemblages from the C€ur€uk Da�g section are domi-

nated by Bairdioidea. Throughout its history, this super-

family has been typical of normal oxygen conditions (R.

Maddocks, pers. comm.). Its dominance testifies to good

oxygenation at the base of the water column. Although

palaeoecological requirements of Kirkyoidea are still

unclear, they are also major components of the assemb-

lages and seem to be associated with similar conditions to

Bairdioidea. Therefore, the overall dominance of these

two superfamilies throughout the microbial deposits in the

C€ur€uk Da�g section documents open marine, normal oxy-

gen conditions at the base of the water column. A possible

degradation of oxygenation might be recorded from P2 to

Table 1. Details of the three sedimentological and faunal unitsrecognized for the microbial deposits in the C€ur€uk Da�g section,Taurus, Turkey.

Facies Diversity Composition

Oolites P4 – D4 Decrease of KirkbyidaeIncrease of CytherissinellidaeCypridoidae

Thrombolites D2 – P3 – D3 Increase of KirkbyidaeStromatolites P1 – D1 – P2Hybrid

microbialites

Figure 24. Changes in the number of species (species richness), number of specimens (abundance) and relative proportions of eachsuperfamily/family found in ostracod assemblages through the lowermost Griesbachian microbialite sequence of the C€ur€uk Da�g section,Taurus, Turkey.


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the top of the section, but it is still difficult to distinguish

the respective influence of oxygenation and water depth.

The growth of microbialites is not influenced by water

salinity (McNamara 2009). Despite the relative instability

of the environment recorded in the C€ur€uk Da�g section

through microbial development, the continuous presence

of stable Bairdioidea/Kirkbyoidea assemblages reveals

adaptation to environmental conditions in the area. The

impacts of the variations of salinity on ostracods are gen-

erally observed on the morphology of the valves, such as

the variation of nodation with salinity changes of Recent

Cyprideis torosa (e.g. Van Harten 1975; Boomer & Frenzel

2011). No thickening or nodation of the carapaces was

observed in the PTBM here, and the range of fluctuation of

salinity may therefore have been relatively narrow, never

reaching lethal levels for these types of ostracods. We

recently proposed a two-step oxygenation process to explain

the co-existence of low-oxygenation proxies and faunas of

relatively normal oxygenation setting (Forel et al. 2013b)

and identified microbial deposits following the EPE as ref-

uge areas. In the present section, ostracods are still present

in the 4 metres following the microbial deposits, which are

of oolitic facies. This pattern could be explained by the

non-fossilization of microbial mats within the oolites, still

providing a temporary shelter to ostracod faunas.

The heterochronies discussed above record the persis-

tence of a relatively degraded environment, in spite of the

refuge characteristics provided by the microbial ecosys-

tem. Although it is unclear whether they were the conse-

quences of environmental stress or adaptations to cope

with, they provided both individual and community abili-

ties to survive and maintain the viability of populations.

However, the nature of this stress cannot be solved in our

present state of knowledge of ostracod ecology: factors

influencing progeny size are climatic conditions, location

where eggs are laid, population densities, predation, com-

petition, starvation, dessication, oxygen stress, tempera-

ture stress, nutritional stress and environmental toxins

(Fox & Czesak 2000 and references therein).

Conclusions

1. Like in other localities yielding microbial deposits

in the earliest Griesbachian, ostracods are abundant

at the C€ur€uk Da�g section throughout microbialites

and become extinct shortly after the end of micro-

bial growth.

2. Fifty-seven species were found, amongst which

three are new: Monoceratina hussonae sp. nov.,

Reviya sylvieae sp. nov. and Eukloedenella adcapi-

tisdolorella sp. nov. Possible sexual dimorphism for

fossils of the genus Bairdia is reported for the first

time in the species Bairdia? kemerensis. This obser-

vation is the first record of the family

Kloedenellidae and the genus Eukloedenella in the

Early Triassic, showing that these taxa crossed the

Permian–Triassic boundary.

3. Heterochrony by paedomorphosis (deceleration) is

identified in six species: Bairdia? kemerensis, Liuz-

hinia antalyaensis, Praezabythocypris? ottomanen-

sis, Paracypris gaetanii, Reviya curukensis and

Reviya sylvieae. Peramorphosis (acceleration) is rec-

ognized in Eukloedenella adcapitisdolorella. These

patterns are interpreted as secondary survival strate-

gies in the aftermath of the end-Permian extinction.

4. Because of the overall dominance of Bairdioidea/

Kirkbyidae, oxygen conditions are interpreted as

normal at the base of the water column during the

growth of microbial mats at the C€ur€uk Da�g section.

Acknowledgements

This work is part of IGCP 572 ‘Restoration of marine eco-

systems following the Permian–Triassic mass extinction:

lessons for the present’ and Chinese programs NSFC

(40621002) and 111 (B08030) and was funded by Actions

transversales du museum (ATM) Biodiversit�es actuelles

et fossiles. It was undertaken at the UMR 7207 Centre de

recherche sur la pal�eobiodiversit�e et les pal�eoenvironne-ments (CR2P), Paris. I thank Prof. Dave Horne (Univer-

sity College, London, UK), Dr Gengo Tanaka (Japan

Agency for Marine-Earth Science and Technology, Japan)

and an anonymous reviewer for their comments and cor-

rections that greatly improved an earlier version of the

manuscript. I am also grateful to Prof. Horne for his valu-

able advice. My gratitude is addressed to Dr Sylvie

Crasquin (CNRS, Pierre et Marie Curie University, Paris,

France) for her advice, trust, and motherly support; it will

always be remembered. I am indebted to Prof. Steve

Kershaw (Brunel University, UK) and Dr Pierre-Yves

Collin (Burgundy University, France) for their construc-

tive remarks, and to Aymon Baud (Lausanne, Switzer-

land) and Erdal Kosun (Akdeniz University, Antalya,

Turkey) for their help in the field. I thank Martine Fordant

and Alexandre Lethiers (Pierre et Marie Curie Univer-

sity), respectively, for their help with the processing of

samples and the drawings. Finally I thank Brigitte L�etang(France) for patiently correcting my ‘frenglish’.

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Heterochronic growth of ostracods (Crustacea) from microbial deposits in the aftermath of the...

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