Capitanian (Middle Permian) Mass Extinction and Recovery in Western Tethys: A Fossil, Facies, and 13C Study From Hungary and Hydra Island (Greece)

PALAIOS, 2012, v. 27, p. 78–89

Research Article

DOI: 10.2110/palo.2011.p11-058r

CAPITANIAN (MIDDLE PERMIAN) MASS EXTINCTION AND RECOVERY IN WESTERN TETHYS:A FOSSIL, FACIES, AND d13C STUDY FROM HUNGARY AND HYDRA ISLAND (GREECE)

PAUL B. WIGNALL,1* DAVID P. G. BOND,1 JANOS HAAS,2 WEI WANG,3 HAISHUI JIANG,4 XULONG LAI,4

DEMIR ALTINER,5 STEPHANIE VEDRINE,6 KINGA HIPS,2 NORBERT ZAJZON,7 YADONG SUN,4 andROBERT J. NEWTON 1

1School of Earth and Environment, University of Leeds, Leeds LS2 9JT, United Kingdom, [email protected], [email protected], [email protected];2Geological Research Group of the Hungarian Academy of Sciences, Budapest, Hungary, [email protected], [email protected]; 3Nanjing Institute of Geology and

Paleontology, Chinese Academy of Sciences, 39 East Beijing Road, Nanjing 210008, People’s Republic of China, [email protected]; 4Key Laboratory of

Geobiology and Environmental Geology, China University of Geosciences, Wuhan 430074, People’s Republic of China, [email protected], [email protected];5Marine Micropalaeontology Research Unit, Department of Geological Engineering, Middle East Technical University, TR-06531 Ankara, Turkey, [email protected];

67 rue Albert 1er, 45000 Orleans, France, [email protected]; 7Department of Mineralogy and Petrology, University of Miskolc, Miskolc, Hungary,

[email protected]

ABSTRACT

The Capitanian (middle Permian) extinction and recovery event isexamined in carbonate platform settings from western Tethys (Hungaryand Hydra, Greece). The age model for these sections is poorly resolvedand we have constructed a d13C chemostratigraphic correlation scheme,supported by conodont and foraminifer data, which attempts correlationwith the well-dated events in China. This reveals the timing of events wassimilar in all Tethyan regions: extinction losses in the middle of theCapitanian produced late Capitanian assemblages in Hungary and Hydrawith a distinctive late Permian character (for example, they lack largefusulinaceans). There is no evidence for an extinction event at the end ofthe Guadalupian (Capitanian) suggesting that previous claims for an end-Guadalupian mass extinction are based on poorly dated records of a mid-Capitanian event. Base level was stable through much of the middle–latePermian transition with the exception of a major regression within theCapitanian Stage. The subsequent transgression established widespreadshallow-water carbonate deposition, such as the Episkopi Formation inHydra and the Nagyvisnyo Limestone Formation in Hungary.

INTRODUCTION

Until 20 years ago the Permian mass extinction was viewed as aninterval of protracted diversity decline throughout the final ,10 millionyears of the Permian (Newell, 1967; Teichert, 1990; Erwin, 1993) butmost subsequent studies favor rapid extinction in the latest Permian(Wignall and Hallam, 1992; Hallam and Wignall, 1997; Rampino andAdler, 1998). The discovery of a precursor extinction event at the end ofthe middle Permian (Guadalupian) suggests that claims for a prolongeddecline may be the result of conflating the extinction losses of twodiscrete events (Stanley and Yang, 1994; Jin et al., 1994). However,recent analysis of middle and late Permian marine invertebratecommunities led Clapham and Bottjer (2007) to conclude that diversitywas in long-term decline from the middle to the end of the Permian; afinding that reasserts the pre-1990s views of Newell and Teichert. LatePermian communities are also claimed to show a progressively moreMesozoic appearance as mollusk groups (bivalves and gastropodsespecially) became proportionally more important at the expense ofrhynchonelliform brachiopods (Clapham and Bottjer, 2007; Isozakiet al., 2007; Clapham et al., 2009). The status of a discrete Guadalupianmass extinction has thus been questioned (Fraiser et al., 2011).However, detailed studies of sections in South China have shown thatthere is a sharp extinction crisis for many groups within the middle ofthe Capitanian, the final stage of the Guadalupian (Wang andSugiyama, 2000; Wignall et al., 2009a, 2009b; Shen and Shi, 2009;

Bond et al., 2010a, 2010b). The loss of many fusulinacean foraminifersis constrained to the mid-Capitanian Jinogondolella altudaensisconodont Zone, an interval that coincides with the onset of eruptionof the Emeishan large igneous province in southwestern China (Wignallet al., 2009a). Similarly, many of the dominant large brachiopods of theearly Capitanian were extinct by the late Capitanian (Shen and Shi,2009). Brachiopod diversity recovered significantly in the Lopingian(e.g. Shen et al., 2010). The data compilation exercise of Clapham andPayne (2011) reveals that 41% of the 286 extant Capitanianbrachiopods failed to survive beyond the Stage. The extinction levelamong other groups is less well known but the generic extinctionintensity of rugose corals was twice that of the brachiopods (Wang andSugiyama, 2000).

Eustatic sea level probably reached its Phanerozoic low point aroundthe Guadalupian-Lopingian boundary (Hardenbol et al., 1998; Hallamand Wignall, 1999). It marks, for example, the point of peak regressionin the Paleozoic, carbonate-dominated successions of the Middle East(Weidlich and Bernecker, 2003; Al-Husseini and Matthews, 2010). Theapparent close temporal association between the extinction losses andthis major regression has led many to attribute the marine crisis to theloss of shallow shelf habitat area (Jin et al., 1994; Hallam and Wignall,1997, 1999; Lai et al., 2008; Bond and Wignall, 2009; Chen et al., 2009).However, recent conodont biostratigraphic dating in China has shownthe age of this major sequence boundary to be precisely constrained towithin the Jinogondolella xuanhanensis Zone of the middle Capitanian(Wignall et al., 2009b; Bond et al., 2010a; Sun et al., 2010; Fig. 1), this isone zone later than the mass extinction level in this region. Otherstudies have suggested cooling-driven extinction (Isozaki et al., 2007)or a crisis directly linked to the onset of Emeishan volcanism (Wignallet al., 2009a, 2009b).

This paper aims to examine the nature of faunal and sedimentarychanges in the Capitanian to Wuchiapingian interval in westernTethyan locations of Hungary and Greece (Fig. 2) and compare themwith changes in the well-known sections of South China. The nature ofthe post-extinction assemblages is also considered; are they typicalPermian types, or do they have high proportions of mollusks suggestinga transitional Mesozoic character (Clapham and Bottjer, 2007;Clapham et al., 2009)?

Detailed facies changes have been studied in four sections to evaluaterelative sea level and provide a context for fossil range charts: both areimportant factors in understanding this extinction event. Foraminifersand calcareous algae are the dominant bioclasts in Guadalupian Tethyancarbonates and their occurrence is strongly controlled by paleobathy-metry (e.g., Vachard et al., 2003; Insalaco et al., 2006). Consequently,rapid facies changes produce rapid compositional changes, and it istherefore important to constrain facies variation when evaluating rangecharts of these groups.* Corresponding author.

Copyright G 2012, SEPM (Society for Sedimentary Geology) 0883-1351/12/0027-0078/$3.00

Facies control of fossil distribution has also bedevilled biostrati-graphic study of many Permian limestone formations because dating isoften attempted using benthic groups such as foraminifers andostracodes. The result is conflicting age determinations. Carbon isotopechemostratigraphy offers an alternative approach to dating thanks tothe recent establishment of a Capitanian-Wuchiapingian d13C curvefrom carbonates in South China and the Panthalassa Ocean (Bondet al., 2010a). Thus, the d13C record of our study sections has beenanalyzed in combination with conodont sampling. Conodonts providethe most widely applicable and highest resolution zonal scheme for the

Permian and, although they are generally rare in the shallow-watercarbonate facies studied here, the occasional find has provided tiepoints for our chemostratigraphic correlation.

STUDY AREAS

Bukk Mountains, Hungary

The ,250-m-thick Nagyvisnyo Limestone Formation of the BukkMountains in northern Hungary contains abundant middle–late

FIGURE 1—Chemostratigraphic correlation scheme for the Permian study sections in Hungary (Mihalovits) and Hydra (Episkopi) based on correlation with d13C values from

conodont-zoned data from South China (Bond et al., 2010a). Thus, heavy (4%o–5%o) early Capitanian values are inferred to be present in the Marmari Formation and heavier

values (.5%o) of the later Capitanian are seen in the Episkopi and Nagyvisnyo Limestone formations. The level of Emeishan flood volcanism, marine extinction and the major

middle Permian sequence boundary in South China (Wignall et al., 2009a) are shown for comparison. These occur in the J. xuanhanensis–J. prexuanhanensis Zone. The

conodont genera are Jinogondolella (J) and Clarkina (C).

PALAIOS MIDDLE PERMIAN EXTINCTION WESTERN TETHYS 79

Permian taxa in dark, pyritic limestones (Fulop, 1994; Berczi-Makket al., 1995; Haas et al., 2004). In the lower member of the formation(50–100 m) dolomite and limestone beds alternate, showing an upward-decreasing trend of dolomitization. Hermatypic corals (Waagenophyl-lum) and calcareous sponges were found in the topmost bed of thelower member. The middle member (Mihalovits Member; 60–80 mthick) is characterized by dark-gray to black limestone, punctuated bythin shale layers. It is rich in calcareous algae, foraminifera, andostracodes; bivalves, gastropods, brachiopods, and nautiloids alsooccur. The upper member of the formation (60–80m) is typified by shaleand clayey nodular limestone interbeds. These beds are rich incalcareous algae, foraminifera, ostracodes, and in some beds, brachio-pods (Leptodus) and mollusks; trilobites also occur. The formation wasformed in a euphotic, shallow marine environment that was intermit-tently restricted. The faunal assemblage suggests usually normal marinebut periodically high salinity conditions. The rich biota and the low-energy depositional environment led to reducing diagenetic conditionsbelow the water/sediment interface and accumulation of organic-richand pyritic deposits. The precise age of this unit has proven difficultto constrain. Berczi-Makk et al. (1995) suggest a Capitanian toWuchiapingian age based on foraminifer species, whereas Kozur(1985) suggests a latest Permian (Changhsingian) age for the upperpart of the formation based on ostracodes. In contrast, Thery et al.(2007) have argued that the Nagyvisnyo Limestone belongs entirely tothe Changhsingian Stage based on the occurrence of Paradagmaritaand Multidiscus.

We have examined a 40-m-thick section in the Nagyvisnyo Limestonein an abandoned quarry at Nagyvisnyo (Mihalovits Quarry) (Fig. 3).This is one of the lowest sections available in the formation in the BukkMountains and it potentially records the Capitanian/Wuchipingiantransition. A total of 70 samples was collected for thin-section analysisof microfacies and microfossil identification, 90 samples were collected

for C and O isotope analysis, and 20 large samples were collected forconodont extraction.

Hydra Island (Greece)

Nearly 500 m of Permian strata, mostly limestone, are present on theGreek island of Hydra (Idhra) with the Guadalupian-Lopingianinterval reported from the Marmari and Episkopi formations(Romermann, 1968, 1969; Grant et al., 1991). Thanks to silicificationthe fauna of the Episkopi Formation is well preserved and thebrachiopods and ostracodes in particular are well studied (Grant et al.,1991; Grant, 1994; Crasquin-Soleau and Baud, 1998). The strataaround the boundary between the Marmari and Episkopi formationshave been logged in detail at Cap Bisti, Episkopi, and Lehusis on thesouthwest coast of the island (Fig. 4). A total of 71 thin sections wasanalyzed for their microfacies and microfossil content, and the d13C andd18O values of 18 samples straddling the formational boundary atEpiskopi were obtained.

Both the Marmari and Episkopi formations thicken toward thesouthwest of Hydra. Thus, the Marmari Formation is 117 m thick atLehusis, 145 m thick at Episkopi and 178 m thick 0.7 km to thesouthwest of this last location (Grant et al., 1991). The same thicknesstrend is seen in the Episkopi Formation: it is 200 m thick at its typelocality but thins to ,10 m over a distance of 10 km to the northwest(Grant et al., 1991). These thickness variations suggest half-graben stylestructural control on deposition. However, for much of the depositionalhistory sedimentation remained close to base level.

The Marmari Formation consists of two, extensively dolomitized,peritidal cycles (Baud et al., 1990) with an assemblage of brachiopods,calcareous algae and ostracodes that is of low diversity compared tothat found in the overlying Episkopi Formation (Grant et al., 1991;Crasquin-Soleau and Baud, 1998; Vachard et al., 2008). There are onlya few biostratigraphically useful taxa, but the presence of Neoschwa-

gerina margaritae in the middle and upper part of the formationsuggests a Wordian-Capitanian age (Baud et al., 1990; Grant et al.,1991), and also a pre-Capitanian extinction assemblage.

The contact between the Marmari and Episkopi formations iserosional, and in southwestern Hydra the basal bed of the EpiskopiFormation is a calcirudite composed of reworked clasts of the MarmariFormation (Grant et al., 1991; Jenny et al., 2004). The duration of thehiatus at this contact is difficult to precisely constrain, but the Yabeina

globosa fusulinid Zone is absent, implying much of the CapitanianStage is missing (Grant et al., 1991; Jenny et al., 2004). The presence ofParaglobivalvulina mira and Codonofusiella aff. schubertelloides in thebasal 90 m of the Episkopi Formation is used by Baud et al. (1990,

FIGURE 3—Location of the middle-upper Permian Mihalovits Quarry section in

northern Hungary.

FIGURE 4—Location of middle-upper Permian study sections in Hydra, Greece.

FIGURE 2—Global paleogeography of the middle Permian showing location of the

two study areas in western Tethys.

80 WIGNALL ET AL. PALAIOS

p. 195) and Jenny et al. (2004) to suggest ‘‘a late? Midian’’(5Capitanian) age. However, Codonofusiella populations are typicallyrecorded from the Wuchiapingian, and conodonts diagnostic of thisStage were obtained in a sample 80 m from the top of the formation(Nestell and Wardlaw, 1987). The Marmari/ Episkopi erosional contacttherefore marks either an intra-Capitanian hiatus or a hiatus straddlingthe Capitanian-Wuchiapingian boundary. The level of the Capita-nian mass extinction, identified in South China at the end of the J.altudaensis Zone (Wignall et al., 2009a), could be recorded in the upperMarmari Formation or it may be missing at the unconformable contact.

AGE OF STUDY UNITS

Biostratigraphic Evidence

Attempts to extract conodonts from Nagyvisnyo Limestone sampleswere generally unsuccessful; only a few fragments were found, with theexception of a well-preserved specimen of Jinogondolella granti from theupper part of the section (Fig. 5). This indicates the presence of the lateCapitanian J. granti Zone at this level. The occurrence of Rectostipulinaquadrata around this level further supports this age assignment becausethis foraminifer appears and is abundant in post-extinction assemblagesof the J. granti Zone in South China (Wignall et al., 2009b; Bond et al.,2010a). R. quadrata was also found to be common in the basal EpiskopiFormation in all three study sections in Hydra.

Carbon Isotope Analysis and Results

Well-screened, whole-rock carbonate (mostly micrite) samples fromEpiskopi and Mihalovics were measured for C and O isotopes at thestable isotope laboratory of the Nanjing Institute of Geology andPalaeontology (MAT 253). CO2 was generated by the addition of an-hydrous phosphoric acid to around 5 mg of powdered whole rock in avacuum. Values have been corrected with standard methods and arereported relative to the Vienna Pee Dee belemnite (VPBD) standard.The analytical precision for this analysis, based on replicate analyses ofan in-house strontium carbonate standard, is ,0.1%o.

The d13C results from Mihalovits quarry show stable values around+4%o throughout much of the section, with values falling to +2%o–+3%o

between 11 and 17 m and above 35 m in the section (Figs. 1, 5). Thed18O values are also stable, and there is no correspondence between theminor fluctuations of the two isotopic curves, suggesting little dia-genetic influence.

The d13C results from the Marmari Formation of the Episkopi sectionshow an initial value of 2%o rising to stable values of 4.5%o in the topmost4 m (Figs. 1, 6). Values fall sharply to 3%o across the unconformablecontact at the base of the Episkopi Formation and then rise rapidly to5%o in the overlying 5 m of section. The O isotope values also show asharp fall across the formation boundary, but the second orderfluctuations do not correspond to variations in d13C values. The Cisotope record at Episkopi is therefore considered to be a primary one.

Chemostratigraphic Correlation

Using the occurrence of J. granti (and R. quadrata) as a tie point, it ispossible to align the Mihalovits d13C curve with the late Capitanianrecord from South China (Fig. 1). In this tentative correlation, the minornegative excursion at the top of the Hungarian section equates with anexcursion of similar size and magnitude seen in the base of theWuchiapingian in China (Wang et al., 2004). No conodonts have beenobtained from the Hydra sections, but the presence of abundant R.quadrata at the base of the Episkopi Formation suggests a J. granti Zoneage for this level; a correlation that accords with the similar d13C record.

By Phanerozoic standards, the Capitanian Stage is marked by heavyd13C values, and Isozaki et al. (2007) has called the +4%o plateau theKamura event. The +4%o values in the Marmari Formation are taken to

indicate a Capitanian age (Fig. 1), in accord with the age determinedusing foraminifer taxa (Baud et al., 1990; Grant et al., 1991). The rapidrise of d13C values in the basal Episkopi Formation can be matchedwith the rise seen around the J. xuanhanensis–J. granti zonal boundaryin the Bond et al. (2010a) curve. However, the major intra-Capitaniannegative excursion of this curve is missing in Hydra, presumably be-cause of a hiatus at the formation contact (Fig. 1).

The chemostratigraphic correlation proposed here allows the baseEpiskopi unconformity to be correlated with the major J. xuanhanensissequence boundary in China (Wignall et al., 2009b; Bond et al., 2010a;Fig. 1). The subsequent transgression saw the re-establishment ofthe platform carbonates of the Episkopi Formation. The onset ofNagyvisnyo Limestone Formation deposition in northern Hungary isprobably a similar response to base-level rise in the late Capitanian.

HUNGARY: FACIES AND FOSSILS

The Nagyvisnyo Limestone Formation in the Mihalovits Quarryconsists of tabular bedded wackestone and packstones ranging from 0.2to 3.0 m thick, separated by thin (1–10 cm thick) beds of calcareousshales (Fig. 5). Most limestone beds are massive but some show gradingwith bioclasts concentrated at the base and more micritic tops bur-rowed by Psilonichnus. Four facies types are recognized: (A) mollusk-algal-echinoderm packstone (Fig. 7A), (B) algal wacke-packstone(Fig. 7B), (C) echinoderm wackestone/packstone (Fig. 7C), and (D)ostracode wackestone (Fig. 7D). Facies A–C are very similar, beingdistinguished by their varying proportions of the same bioclasts (seebelow). Facies D is distinctly different because it contains fewer bioclasttypes, predominantly ostracode valves, and has patches of anhydritecrystals replaced by calcite (Fig. 7D).

With the exception of facies D, the limestone beds contain a diverseassemblage consisting of foraminifers, calcareous algae, Tubiphytesobscurus, echinoderm material (especially crinoids), brachiopods, ostra-codes, microgastropods, and bivalves. The foraminifers include rarefusulinaceans, of which only Codonofusiella is identifiable, while smallervarieties, mostly lagenides and miliolinids, are much more abundant andinclude Agathammina, Diplosphaerina, Globivalvulina, Hemigordius,Nodosinelloides, and Pachyphloia (Fig. 5). These are all genera knownto have survived the Capitanian mass extinction (Bond and Wignall,2009). Calcareous algae are the most abundant bioclasts in theMihalovits Quarry section and assemblages are dominated by Permo-calculus spp., Mizzia spp., and Gymnocodium bellerophontis that rangethroughout the section (Figs. 5, 7B). Such dominance by calcareousalgae is typical of Permian carbonates, however gastropods and bivalvesare dominant in beds of facies A. Mollusk groups are more typical ofMesozoic assemblages and in this regard the Nagyvisnyo Limestone hassome beds with the mixed Permian-Mesozoic faunal character suggestedfor late Permian carbonates by Clapham and Bottjer (2007).

In summary, the Mihalovits Quarry record consists of moderatelydiverse fossil assemblages that lack Capitanian mass extinction victimssuch as schwagerinids and neoschwagerinid fusulinaceans. This suggeststhat the Capitanian crisis predates the J. granti Zone in Hungary.Bioclastic packstone deposition was punctuated by intervals ofevaporation and anhydrite growth supporting previous restricted,lagoonal interpretations (Berczi-Makk et al., 1995) as do the absenceof some stenohaline groups such as ammonoids and bryozoans and therare occurrence of corals (Fulop, 1994). However, echinoderms,conventionally regarded as a stenohaline group, are present throughout.

HYDRA: FACIES AND FOSSILS

Facies Trends

The uppermost Marmari Formation consists of pale gray–whitebeds that contrast with the darker gray limestones of the Episkopi


Formation. There is modest lateral facies variation between the threesections and all show low diversity compared with the overlying EpiskopiFormation (Figs. 6, 8–9). Algal-ostracode wackestones and packstonesdominate at Lehusis and Cap Bisti. These facies are also present at

Episkopi but beds of pelmicrite are more common. In the topmost bedsat Cap Bisti some bioclasts are replaced by chert-calcite intergrowths.

The nature of the transition between the Marmari and Episkopiformations varies substantially between sections. At Cap Bisti the

FIGURE 5—Section in the lowermost Nagyvisnyo Limestone Formation at Mihalovics quarry showing an alternating succession of the four facies: a 5 mollusk-algal-

echinoderm packstone, b 5 algal wacke-packstone, c 5 echinoderm packstone, d 5 ostracode wackestone; and C and O isotope data and range charts for foraminifers and

calcareous algae. Photograph shows Jinogondolella granti obtained from sample 68, scale bar is 100 mm.


uppermost bed of algal-ostracode wackestone in the MarmariFormation is deeply karstified with a network of vertical and horizontalfissures several decimeters in width penetrating up to 3 m downward(Fig. 8). This is infilled and overlain by a 2-m-thick bed of clast-supported breccia consisting of reworked clasts of the underlying beds.This bed is in turn partly truncated by erosion at the base of a furtherbed of clast-supported breccia with clasts approaching one meter indiameter. Reworked Marmari clasts continue to occur up to 4.5 mabove the base of the overlying Episkopi Formation where they arepresent as rounded, floating pebbles in a dark matrix of echinodermpackstones and pelsparites. Clast orientation varies from random, highangle orientations to bedding parallel pebble trains that do not showimbrication. Further east, at Episkopi, evidence for erosion of thetopmost Marmari Formation is less spectacular but reworked clasts arepresent in the uppermost part of the formation although coarse,angular pebbles are not present (Fig. 7E). Instead, rounded pebblesoccur either in sharp-based beds of conglomerate or as floating clastsin a bioclastic packstone or grainstone matrix. At the easternmost lo-cation (Lehusis), the formation contact is seen as an irregular surface,

with ,10 cm of erosional relief between the white Marmari Formationand the dark-gray Episkopi Formation. A few, small rounded Marmaripebbles are present immediately above this surface, but the conglom-erates and breccias of the more western localities are not seen at Lehusis(Fig. 9).

The onset of deposition of the Episkopi Formation saw the establish-ment of a uniform facies style in southwestern Hydra: foraminifer-algalpackstones dominate the basal beds at all three locations, althoughmollusk clasts are not uncommon (Fig. 7F). Higher levels, between 70and 100 m above the basal contact, were sampled at Episkopi. Thisrevealed that this facies persists, albeit with a larger component of largebioclasts (colonial Rugosa, calcareous sponges, and brachiopods).

Fossil Ranges

The uppermost Marmari beds sampled here contain a low diversityfauna of ostracodes, calcareous algae and foraminifers (Fig. 10). Theforaminiferal records are restricted to sparse occurrences of smaller

FIGURE 6—Episkopi section showing d13C and d18O fluctuations in the uppermost Marmari and lower Episkopi formations and foraminifer and calcareous algal range

charts. No extinction horizon is identifiable and so this section is considered to postdate the middle Capitanian crisis.


forms (mostly lagenides), including Dagmarita, Earlandia, Eotuberitina,Glomomidiella, Hemigordius, Nodosinelloides, and Pachyphloia. Anindeterminate staffellid fusulinacean (probably Staffella) was recoveredfrom sample EK6 at Episkopi, where it ranges up into the overlyingEpiskopi Formation (Fig. 10L); a unit with an abundant and diverseforaminifer assemblage including those genera listed above, pluscommon Agathammina pusilla, Climacammina, Diplosphaerina in-equalis, Geinitzina, Globivalvulina, Palaeotextularia, Pseudomidiella,Rectostipulina quadrata (and lesser R. pentamerata), Retroseptellina,as well as numerous less common genera (Fig. 10). Fusulinaceanforaminifera are limited to the smaller genera Codonofusiella andReichelina, plus Palaeofusulina and Staffella (?). Codonofusiella andReichelina are biostratigraphically useful because, although they arepresent before the Capitanian extinction in South China, they come todominate the fusulinacean components of post-extinction faunas in thatarea. Their presence in the Episkopi Formation, together with anabsence of fusulinacean extinction casualties, indicates that these strataare of post-extinction age, in keeping with its sequence- and chemo-stratigraphic age assignment. The extinction aftermath attributes of theHydra assemblages are further supported by the common Earlandia

and Diplosphaerina; a typical post-extinction disaster assemblage (e.g.,Groves and Altiner, 2005; Bond et al., 2010a, 2010b).

DISCUSSION

Relative Sea-Level Change

The overall uniformity of facies types in the Mihalovits sectionsuggests that there were no major changes of palaeobathymetry in thislate Capitanian record. The Hungarian record is here correlated withthe lower Episkopi Formation of Hydra (Fig. 1) that similarly showspersistent, uniform deposition in a shallow, platform setting. Incontrast, the sedimentological evidence from the Marmari-Episkopiboundary clearly records a major regression. At Cap Bisti, the mostwesterly location, sea level fell sufficiently for emergence and karsti-fication of the upper Marmari Formation to occur, but at the moreeasterly locations, the upper surface is only eroded to a depth of a fewcentimeters, which may have occurred in a shallow marine setting. Thecoarse breccia beds at Cap Bisti are probably a subaerial scree slopefacies, perhaps part of a fan conglomerate, but this facies is absent in

FIGURE 7—Photomicrographs of typical facies types at Mihalovits Quarry (A–D) and Hydra (E–F). Facies A) Mollusk-algal-echinoderm packstone, showing Permocalculus

cf. tenellus grains (5A), echinoderm grain (5E) and mollusk (5M), likely a gastropod. B) Algal wacke-packstone dominated by Gymnocodium bellerophontis, a complete thallus

is labelled A. C) Echinoderm wackestone. D) Ostracode wackestone, although the patch illustrated consists of anhydrite crystals now pseudomorphed by calcite and showing

relict zonation. E) The upper two-thirds of the field of view is a clast of calcareous algal packstone in a micrite matrix, topmost Marmari Formation, Episkopi. F) Bioclastic

packstone/grainstone showing mollusk clasts (thick-shelled bivalves (5B) and microgastropods (5M)), calcareous algae (5A) and peloids (5Pe), lower Episkopi Formation,

Episkopi. Width of field of view is 6 mm for all images.


the east. The scree slope facies is overlain by shallow marine limestoneswith rounded pebbles of Marmari Formation at Cap Bisti. This faciescharacterizes the uppermost Marmari Formation to the east. Therounding of pebbles suggests beach reworking prior to transport into ashallow marine setting.

A Capitanian Mass Extinction in Western Tethys?

The fossils of the Nagyvisnyo Limestone and Episkopi formationshave distinctive post-Capitanian extinction attributes, while extinctionvictims (large fusulinaceans) are only known from low in the Marmari

FIGURE 8—Sedimentary log of the Cap Bisti section, westernmost Hydra Island showing a deeply karstified surface in the uppermost Maramari Formation and microfossil range chart.


Formation (Baud et al., 1990; Grant et al., 1991; Jenny et al., 2004). Themass extinction level therefore occurs, either within the upper MarmariFormation, or at the boundary with the Episkopi Formation. UpperMarmari assemblages are impoverished and of similar composition topost-extinction assemblages in South China (Lai et al., 2008; Wignall

et al., 2009b; Bond et al., 2010a). This could reflect an unfavorableenvironment because the facies at this level record peritidal/supratidalconditions (Grant et al., 1991; Jenny et al., 2004). The carbon isotopevalues from the upper Marmari Formation suggest an early Capitanianage (J. altudaensis Zone or older), which places this strata below the

FIGURE 9—Sedimentary log and microfossil range chart from the Lehusis section, south Hydra Island.


extinction level seen in China (Fig. 1). Therefore, it is likely that theCapitanian mass extinction occurred during the hiatus recorded by thesequence boundary between the Marmari and Episkopi formations.The clear signal of a major intra-Capitanian regression, in a stageotherwise characterized by little eustatic change, occurs in both Chinaand Hydra. In Hydra the regression and extinction may coincide,suggesting a possible causal connection (e.g., Jin et al., 1994; Hallamand Wignall, 1999; Lai et al., 2008). However, in China, the regressionslightly post-dates the extinction event, and a more circumspect

conclusion to be drawn from the western Tethyan record would bethat the extinction occurs during the hiatus caused by base level fall.

The Nature of Late Permian Assemblages

The contention that post-Capitanian extinction assemblages weremore mollusk-rich than earlier Permian times (Clapham and Bottjer,2007) receives some support from this study, particularly in themollusk-dominated facies A of the Nagyvisnyo Limestone. However,

FIGURE 10—Foraminifera from Episkopi, Hydra. A) Dagmarita chanakchiensis (sample EK14); B) Diplosphaerina sp. (EK19); C) Diplosphaerina reitlingerae (EK14); D)

Geinitzina araxensis (EK14); E) several Hemigordiopsis sp. (EK14); F) Hemigordius irregulariformis (EK13); G) Pachyphloia ovata (EK14); H) Pseudomidiella sp. (EK16); I)

Rectostipulina pentamerata (EK15); J) Rectostipulina quadrata (EK15); K) Reichelina sp. (EK19); L) Staffellid fusulinacean (indet, EK14). All scale bars 5 100 mm except E, H,

and L 5 300 mm.


the occasional presence of evaporite pseudomorphs in this sectionsuggests salinity restriction occurred at least periodically: conditions thatwill have favored euryhaline mollusks. In other regards the microfossilcontent of the normal marine assemblages is dominated by typicalPermian bioclasts such as calcareous algae and foraminifers. Theabundant and diverse Episkopi ostracode fauna is similarly typical ofshallow-water assemblages of late Permian age (Crasquin-Soleau andBaud, 1998). Thus, the ecological transition proposed to have occurredbetween the middle and late Permian, as a possible consequence of theCapitanian mass extinction, is only manifest in the reported increase inthe dominance of mollusks in units such as the Episkopi Formation(Clapham and Bottjer, 2007). In all other regards, the palaeoecologicalattributes of the assemblages are clearly Permian. It has also been arguedthat the Episkopi brachiopods have peculiar attributes more typical ofolder Paleozoic assemblages (Grant et al., 1991, p. 491) but more recentappraisal shows that they are typical Wuchiapingian taxa with a strongsimilarity to eastern Tethyan assemblages of South China and with alesser number of Perigondwanan taxa (Shen and Clapham, 2009).

CONCLUSIONS

Western Tethyan Capitanian-Wuchiapingian sections of Hungaryand Hydra (Greece) reveal that a typical post-Capitanian massextinction fauna is developed from the late Capitanian and persistswithout faunal break into the Wuchiapingian. This is comparable to thepaleontological record seen in South China (where conodonts define arefined zonation scheme) that shows the extinction event happenedaround the top of the J. altudaensis Zone. Sporadic conodonts, fora-minifers, and correlation of d13C records reveal that the timing of thecrisis is likely to be similar in western Tethys, although the precise levelis difficult to pin down in the extensively dolomitized middle MarmariFormation of Hydra. In Hungary, the extinction occurs below theNagyvisnyo Limestone Formation studied here.

A major regression within the Capitanian, seen in China, is alsorecorded by the major base-level fall recorded at the boundary betweenthe Marmari and Episkopi formations in Hydra. This is the only majorbase-level fluctuation in the middle–late Permian in Hydra and itswidespread development (i.e., occurrence in Hungary, Greece, andChina) clearly argues for a eustatic origin. Subsequent base level risesaw the establishment of platform carbonate deposition with a diversefauna of typical Permian aspect seen in both Hydra and Hungary.

ACKNOWLEDGMENTS

We thank Helen Cope for field assistance, Csaba Pero for infor-mation that facilitated fieldwork in Hungary, and Daniel Vachard forhelp with identifying calcareous algae. Matthew Clapham and JohnGroves are thanked for their reviews. This research was supported bythe following grants: Natural Environment Research Council grantNE/D011558/1 to Paul Wignall; National Natural Science Foundationof China (NSFC) and BK2010022 (Jiangsu Province) grants to WangWei; Natural Science Foundation of China (grant 40872002) andChinese State Administration of Foreign Experts Affairs (GrantB08030) to Xulong Lai.

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Capitanian (Middle Permian) Mass Extinction and Recovery in Western Tethys: A Fossil, Facies, and 13C Study From Hungary and Hydra Island (Greece)