Review of Palaeobotany and Palynology xxx (2014) xxx–xxx

PALBO-03575; No of Pages 12

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Review of Palaeobotany and Palynology

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Palynology of the Permian and Triassic of the Tesero and Bulla sections(Western Dolomites, Italy) and consideration about the enigmaticspecies Reduviasporonites chalastus

A. Spina a,⁎, S. Cirilli a, J. Utting b, J. Jansonius 1

a Dipartimento di Fisica e Geologia, Università di Perugia, Via Pascoli, 06123 Perugia, Italyb Geological Survey of Canada (Calgary), 3303 33rd St., N.W. Calgary, Alberta T2L 2A7, Canada

⁎ Corresponding author. Tel.: +39 0755852696.E-mail address: amalia.spina@unipg.it (A. Spina).

1 Deceased.

http://dx.doi.org/10.1016/j.revpalbo.2014.10.0030034-6667/© 2014 Elsevier B.V. All rights reserved.

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Article history:Received 10 April 2014Received in revised form 29 September 2014Accepted 17 October 2014Available online xxxx

Keywords:Permian–Triassic boundaryWestern DolomitesItalyTesero sectionBulla sectionReduviasporonites chalastus

A palynological study of two Permian–Triassic sections of the Southern Alps (Tesero and Bulla sections) wascarried out in order to correlate the palynological data with the available conodont zones. Both the sectionscrop out in the Western Dolomites, the Tesero section in Fiemme Valley and the Bulla section in Badia valley.The two sections include the uppermost Bellerophon Formation and the overlying Werfen Formation. In thelatter the Permian/Triassic boundary (Farabegoli et al., 2007) has been defined by conodonts. The palynologicalassemblages collected in both sections throughout the Bellerophon up to theWerfen formations show composi-tional variationmarked by: a) a dominance of taeniate and non-taeniate bisaccates in the Bellerophon Formationdated as Late Permian; b) a bloomof the organicmicrofossil incertae sedis Reduviasporonites chalastus in the lowerTesero Member (Werfen Formation), found only in the Tesero section. This interval should stratigraphically liebelow the Permian/Triassic boundary, as defined by conodonts recorded in the Bulla section; c) poorly preservedacavate and bisaccate taxa in the upper Tesero Member, above the Permian/Triassic boundary, and in the over-lying Mazzin Member of the Tesero section. The origin and the ecological preferences of Reduviasporoniteschalastus and the implications of its worldwide distribution across the Permian/Triassic boundary are alsodiscussed.

© 2014 Elsevier B.V. All rights reserved.

1. Introduction

In the Southern Alps of Italy, the Val Gardena Sandstone and theBellerophon Formation are the two principal lithostratigraphic unitsspanning from the Middle (?) to Upper Permian. The two units havebeen regarded as a composite transgressive sequence (Massari andNeri, 1997). The Val Gardena Sandstone is composed of siliciclasticsand red beds, deposited in alluvial fans, braided streams and meander-ing rivers under semiarid conditions (Massari and Neri, 1997). TheBellerophon Fm. is characterized by carbonates and evaporites, depositedin a coastal sabkha to shallow-shelf environments. The BellerophonFm. wedges out westward and disappears west of the Adige Valley bymerging into continental deposits of the Val Gardena Sandstone. Bothformations are overlain by the shallow-marine Lower Triassic WerfenFm.. The Tesero section includes the upper part of Bellerophon andWerfen formations. The area has been the subject of numerous studiese.g.: Bosellini, 1964; Assereto et al., 1973; Broglio Loriga et al., 1988;Magaritz et al., 1988; Perri, 1991; Neri et al., 1999; Nicora and Perri,1999; Farabegoli et al., 2007; Groves et al., 2007. The Tesero section is

nology of the Permian and Tlaeobot. Palynol. (2014), http:/

well known since the discovery of the so-called “mixed fauna”, character-ized by in situ Permian-type brachiopods and foraminifers located atabout 1.5–2.0 m above the Bellerophon–Werfen lithostratigraphicalboundary (Neri and Pasini, 1985). This finding allowed a reconsider-ation of the Permian/Triassic Boundary (PTB) position, traditionallyplaced at the base of Werfen Fm. Successively, the recording of cono-dont faunas in the Tesero section enabled the PTB to be preciselylocated. Five conodont zones have been documented (Nicora andPerri, 1999), including the Hindeodus parvus Zone (recorded withintheMazzinMember),which according to the International Subcommis-sion on Triassic Stratigraphy (STS)marks the base of the Triassic system(Yin et al., 2001).

In the Bulla section (including Bellerophon and Werfen formations),the first occurrence of Hindeodus parvus and thus the PTB was recordedat 1.3 m from the base of the Tesero Member (Farabegoli and Perri,1998; Farabegoli et al., 2007). According to the authors this intervalshould correspond to the uppermost part (subunit 5a) of the TeseroMember in the Tesero section,whereNicora and Perri (1999) document-ed some conodonts showing transitional features between Hindeoduspraeparvus and H. parvus. Based on these findings, the first occurrenceof H. parvus in the Tesero section has been considered below theMazzinMember (Nicora and Perri, 1999; Farabegoli et al., 2007). The Bulla sec-tion has been recommended as the PTB parastratotype for the shallow

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marinewestern Tethys (Farabegoli et al., 2007). In the present article,a palynological study of these two important and better knownPermian–Triassic independently dated sections of the Southern Alpswas carried out. Additionally, we discuss the origin and the ecologicalpreferences of the organic microfossil Reduviasporonites chalastus, andthe implications of its worldwide distribution across the PTB.

2. Stratigraphy

2.1. Tesero section

The Tesero section, about 40m thick, is located in the Fiemme Valley(Western Dolomites) where the Bellerophon and Werfen formationsare quite well exposed in a road-cut near the Tesero village (Fig. 1). Itincludes the uppermost Bellerophon and the lower Werfen formations(Fig. 2). The Bellerophon Fm. is mainly characterized by lagoonal toperitidal dolostones and marls (Neri et al., 1999). The exposed tractdoes not exceed 10 m. The formation has been divided into three infor-mal units (Neri et al., 1999; Perri and Farabegoli, 2003): the lowest,the Fossiliferous Dolomite facies, consists of dolostones and dolomiticlimestones (mudstones and wackestones) alternating with marls. Itsdepositional environment has been interpreted as a restricted, probablyhyperhaline, lagoon. This facies corresponds to Unit 1 of Neri et al.(1999). It is overlain by the Ostracod and peritidal dolomite unit (Unit2 of Neri et al., 1999) consisting of marly dolostone with thin peliteinterlayers. Mud-cracks and root-traces attest to subaerial exposuresurfaces. A sharp unconformable–paraconformable erosional surface(U1 of Farabegoli et al., 2007) separates this unit from the overlyingBulla Member. At the base of the Bulla Member (Unit 3 of Neriet al., 1999), blackish pelite has been recognised, grading upwardsinto calcareous marls and bioclastic wackestone and packstone. Asharp paraconformable–unconformable surface (U2 of Farabegoli

Fig. 1. Location map of Bul

Please cite this article as: Spina, A., et al., Palynology of the Permian and Tconsideration about the enigmatic ..., Rev. Palaeobot. Palynol. (2014), http:/

et al., 2007) separates the Bulla Member from the overlying TeseroMember at the base of the Werfen Fm. The Tesero Member has beensubdivided into three informal units (Neri et al., 1999; Perri andFarabegoli, 2003). Of these the lowest, Unit 4, is characterized by alower interval (subunit 4a, less than 10 cm in thickness), of grainstonesand packstones that grades upwards into the first oolitic bank of theTesero Member (subunit 4b, about 90 cm in thickness). The overlyingUnit 5, characterized by a lateral variability, has been subdivided intothree subunits: subunit 5a consisting of oolitic calcarenite beds withmarly interlayers; subunit 5b mainly characterized by bioclastic wacke-stone and packstone; and subunit 5c is marked by oolitic–bioclasticcalcarenites, with mudstone/wackestone intercalations. The uppermostinformal Unit 6 consists of two coarsening upwards parasequences,evolving from micritic limestones with a slight marly content at thebase, to oolitic–bioclastic–intraclastic grainstones and packstones at thetop. The Tesero Member is overlain by the Mazzin Member, which hasbeen also subdivided into two informal units (Neri et al., 1999): Unit 7is composed of bioturbated to laminated marly lime mudstones, and ofthin biocalcarenite storm layers. The overlying Unit 8 is mainly formedby marly lime mudstones, locally with siliciclastics.

The palaeontological content of the Tesero section is very richand diversified as reported by numerous authors (i.e. Neri and Pasini,1985; Italian IGCP 203 Group, 1986; Broglio Loriga et al., 1988;Posenato, 1988, 1999; Perri, 1991; Nicora and Perri, 1999; Groveset al., 2007). Notably, in the first oolitic bank (subunit 4b; Fig. 2), isthe presence of brachiopods as Ombonia and abundant specimens ofBellerophon vaceki, which is considered a Triassic taxon (Posenato,1988). Nicora and Perri (1999) recorded the first occurrence of theconodont Hindeodus. praeparvus and Hindeodus parvus respectivelyin the Tesero Member at 0.30 cm and in the Mazzin Member,11 m above the litostratigraphical boundary Bellerophon–Werfen for-mations. The overlying conodont Zones are the staeschei Zone and the

la and Tesero sections.

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Fig. 2. Schematic stratigraphic logs of the Bulla and Tesero sections (redrawn from Neriet al., 1999; Farabegoli et al., 2007). Our sampling followed themeasurement of Farabegoliet al. (2007) in order to better correlate palynological and conodont data.

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isarcica Zone respectively at 14.3 m and the at 26 m from the base ofWerfen Fm.

2.2. Bulla section

The Bulla section (Figs. 1, 2), about 80 m thick, crops out near theBulla village (20 km north of Tesero) and exposes the whole sequence

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from the PTB to the supposed Induan/Olenekian boundary (i.e. upper-most Bellerophon Fm and Werfen Fm with Tesero Mb, Mazzin Mb,Andraz Mb, Seis Mb and lower Campill Member). In this study, weinvestigated the first 12 m of the Bulla section including the uppermostpart of the Bellerophon Fm. and the overlying Tesero and MazzinMembers of the Werfen Fm. The section starts with the upper part ofthe Bellerophon Fm. (about 3.5m) composed of grey silty dolomitic car-bonates alternating with grey to reddish shale levels from the Ostracodand peritidal dolomite Unit described by Farabegoli et al. (2007)and followed by the Bulla Member. This latter is, characterized bydark-grey packstones alternating with wackestones and thin clayeysiltstones, and it overlies the Ostracod Unit with an unconformable–paraconformable erosional surface, referred to as “U1” by Farabegoliet al. (2007). Another sharp paraconformable–unconformable surface,referred to as “U2” by Farabegoli et al. (2007), marks the boundarybetween the Bellerophon and Werfen formations. According to Perriand Farabegoli (2003) and Farabegoli et al. (2007), the Werfen Fm iscomposed of the lower Tesero Member overlain by the lower MazzinMember followed by the upper Tesero Member and then by the upperMazzin Member (Fig. 2). The lower Tesero Member (2.30 m thick)contains basal packstones and grainstones intercalated to mudstone,wackestone and oolitic packstone. The lower Mazzin Member (2.9 mthick) consists of mudstone and wackestone and the upper TeseroMember is marked by the oolitic packstone and grainstone bearingbeds (1.35 m thick). The overlying upper Mazzin Member consists ofmudstone intercalated to siltstone and packstone (Perri and Farabegoli,2003; Farabegoli et al., 2007).

The Bulla section yielded important fossil fauna as documented bynumerous contributions (e.g Broglio Loriga et al., 1988; Perri, 1991;Farabegoli and Perri, 1998; Perri and Farabegoli, 2003; Farabegoliet al., 2007; Crasquin et al., 2008). The upper part of the Bulla Memberyielded the most diverse foraminifera assemblage of the BellerophonFm. in the Southern Alps and a very rich ostracod fauna. In the lowerTesero Member the conodonts Hindeodus praeparvus and Hindeodusparvus have been recorded respectively at 0.20 m and 1.30 m abovethe base of the Werfen Fm. In the Mazzin Member, the condonts Is.lobata, Is. staeschei, Is. aequabilis and Is. Isarcica have been recorded(Perri, 1991; Farabegoli and Perri, 1998).

3. Material and methods

Palynological analyses were performed on a total of 52 samples(31 samples from the Tesero section and 21 from the Bulla section).All samples were treated by standard palynological acid maceration,density separation of the organic matter, and filtration of the organic-rich residue at 10 μm. Light microscope observations were performedon palynological slides using a LeicaDM1000microscopewithDifferen-tial Interference Contrast (DIC) technique in transmitted light. Imageswere captured with the digital microscope camera and successivelycorrected for contrast and brightness using the open-source Gimp soft-ware. Palynological slides are stored at the Sedimentary Organic MatterLaboratory of the Physics and Geology Department (Perugia University,Italy). Authors of the species names are given on the species list ofpalynomorphs (Appendix 1) and in the plate descriptions.

4. Palynology

4.1. Tesero section

Diverse and abundant palynological associations occur in theBellerophon Fm. showing a gradual change from bottom to top, whilein the overlying Werfen Fm., the microflora generally lacks diversity.The microflora content includes 45 species of miospores and the abun-dant incertae sedis Reduviasporonites chalastus (Table 1; Pls. 1, 2). Atthe base of the Bellerophon Fm. (AS1 to AS4 levels), the microfloristicassemblage is mainly characterized by several taeniate bisaccates

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Table 1Sporomorphs occurrences in the Tesero section.

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as Protohaploxypinus spp., Lunatisporites spp., Lueckisporites spp.,Striatoabieites spp. and Striatopodocarpites spp., non-taeniate bisaccatesas Klausipollenites spp., Alisporites spp. and Jugasporites delasaucei associ-ated with trilete spores such as Calamospora sp., Leiotriletes adnatus andGondisporites sp. The incertae sedis R. chalastus is abundant in few levels(AS1 and AS2) of the Bellerophon Fm. The Tesero Member (from itsbase to AS21 level), is poor in sporomorphs and R. chalastus characterizesalmost the entire organic content of the palynofacies (Fig. 3). Upwards,from the uppermost Tesero Member to the lower Mazzin Member(AS26), immediately before the first occurrence of Hindeodus parvus,palynological assemblage contains taeniate bisaccates as Lunatisporitesnoviaulensis and Lueckisporites virkkiae, less abundant than inBellerophon Fm., and common Protohaploxypinus sp. Minor non-taeniatebisaccates such as Klausipollenites schaubergeri and Sulcatisporites sp. andacavate long-ranging trilete spores, e.g. Calamospora sp. also occur.The last sampled level in the uppermost part of the MazzinMember yielded Cycadopites follicularis, a few Calamospora sp. andR. chalastus.

4.2. Bulla section

The palynological assemblage from the Bulla section is not diver-sified and well preserved. Only four levels (MC6, MC8, MC10,MC12) of the Bellerophon Fm. yielded palynomorphs (Table 2;Plate III). The microflora includes bisaccate taeniate pollen asLunatisporites pellucidus, Lunatisporites noviaulensis, Lunatisporitessp. and Lueckisporites sp., non-taeniate pollen as Jugasporites delasaucei,Jugasporites sp. and Klausipollenites sp. and laevigate, acavate spores.(See Plates I and II.)

The paucity of sporomorphs mostly represented by bisaccatescan be linked to palaeoenvironmental conditions. As suggested byFarabegoli et al. (2007) the Bulla succession could be deposited inmore distal environment, far from the shoreline, than the Tesero one.Under such conditions, only the most buoyant saccate pollen grainscan be dispersed for long distances by wind and water currents(Tyson, 1995).

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4.3. Discussion

Previous studies reported analogous Late Permian palynologicalassemblages also in other localities of the Southern Alps, alwayscharacterized by Lueckisporites virkkiae, Lunatisporites noviaulensis,Lueckisporites parvus, Jugasporites delasaucei and Klausipollenitesschaubergeri (Massari et al., 1988, 1994; Visscher and Brugman, 1988;Cirilli et al., 1998).

Accordingly, this palynological association shows close similaritieswith those recorded from the Late Permian (Lopingian) of many areasof the world. In the Meishan section (China), ratified as GSSP for basalboundary of Triassic System (Yin et al., 2001), Lueckisporites virkkiaeand Klausipollenites schaubergeri have been found in the Vittatina-Protohaploxypinus Assemblage attributed to the Changhsingian-LowerGriesbachian on the basis of the conodont zones (Ouyang and Utting,1990).

Other similarities have been noted with the Lopingian microflorafrom Northern Europe (e.g. Germany: Leschik, 1956; Grebe, 1957;Schaarschmidt, 1963; Great Britain: Clarke, 1965; Ireland: Visscher,1971; Hungary: Haas et al., 1988; Barents Sea: Mangerud, 1994;Greenland: Twitchett et al., 2001) and from the Kazanian type sequenceof Russia (Utting et al., 1997).

Correlation with the Gondwana region shows that the taeniatepollen Lueckisporites virkkiae, Lunatisporites pellucidus, Lunatisporitesnoviaulensis and the non-taeniate Klausipollenites schaubergeri arepresent also in the Late Permian of Pakistan (Balme, 1970; Jan et al.,2009) and L. virkkiae, Protohaploxipinus spp., and Striatopodocarpitesspp. in the Middle East (Stephenson et al., 2003). Similar assemblages,referred to as L. virkkiae Zone, have been documented also in the LatePermian of Israel (Eshet, 1992).

The microfloristic assemblages from the uppermost BellerophonFm. in both Tesero and Bulla sections, confirm an Upper Permian age,also indicated by other fossiliferous assemblages (i.e. brachiopods,Posenato, 1999) documented in the literature. A gymnospermouspollen decrease has been observed within the Tesero and MazzinMembers, after the occurrence of Hindeodus parvus (Figs. 2, 3). Here,

riassic of the Tesero and Bulla sections (Western Dolomites, Italy) and/dx.doi.org/10.1016/j.revpalbo.2014.10.003

Table 2Sporomorph occurrences in the Bulla section.

Fig. 3. Quantitative changes in sporomorph suprageneric groups and in R. chalastus at theTesero section. Due to paucity of sporomorphs no quantitative analysis was performed forthe palynological content of the Bulla succession.

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notwell preserved long-ranging acavate sporeswith bisaccoid pollen asLueckisporites virkkiae were still found. The presence of these “Permiantype” bisaccates within the Lower Triassic strata is still debated asthey are differently considered in literature: a) as reworked from the

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Upper Permian Bellerophon Fm.; b) as ranging into Lower Triassic (i.e.Visscher and Brugman, 1988; Ouyang and Utting, 1990; Mangerud,1994).

5. Previous studies and some considerations about the affinity andthe distribution of Reduviasporonites chalastus (Foster) Elsik, 1999

In the present study, Reduviasporonites chalastus was found invarious abundances at different stratigraphic levels of the Bellerophonand Werfen formations either in the latest Permian and in the earliestTriassic (Fig. 3; Table 1; Plate IV). The origin of the species has beenlargely debated in the literature. Foster (1979), defined Chordecytiachalasta (junior synonym of R. chalastus), as a microfossil incertae sedis.Balme (1980), in his original description of Tympanicysta stoschiana(a junior synonym of C. chalasta in Jansonius and Hills, 1981; cd. 3926),considered it to belong to the plant kingdom, probably representing astage in a fungal or algal lifecycle. Nevertheless the author did not rejectthe possibility that T. stoschiana could be of animal origin. Elsik (1999)formally establishedReduviasporonitesWilson, 1962 as a senior synonymof Chordecystia and Tympanicysta and formally proposed the new combi-nations R. chalastus (Foster, 1979) Elsik, 1999 and Reduviasporonitesstoschianus (Balme, 1970) Elsik, 1999. In this study, we followed thesuggestions of Foster et al. (2002) that considered R. chalastus as seniorsynonym of R. stoschianus.

Several authors (e.g. Elsik, 1999; Visscher et al., 2011) interpretedReduviasporonites as fungal remains, representing hyphae and conidiaof ascomycetes similar to those of themodern Rhizoctonia. The prolifer-ation of Reduviasporonites chalastus recorded at the PTB in variouslocalities (e.g. Israel, Southern Alps, Australia, South Africa) has beenclose associated with the Permian–Triassic mass extinction event(Eshet et al., 1995; Visscher et al., 1996; Steiner et al., 2003; Sandleret al., 2006). This close relationship led several palynologists to supposethat saprophytic fungi may have metabolised the dead vegetationformed during the extinction event. They considered the acme of theresulting “fungal spores” as a proxy bio-event for recognising thePermian–Triassic transition throughout the world. Foster et al (2002)

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demonstrated, by radio-isotopic studyon the cellwall ofReduviasporonitesfrom theMazzinMember (Austria), that it is isotopically lighter (in termsof δ13Corg) than the plant remains associatedwith it. Hence, itwas unlikelyto have consumed thismaterial as food, as would be expected for a fungalsaprophyte and, for this reason, the authors suggested an algal origin forReduviasporonites. This result has been strongly contrasted by Sephtonet al. (2009) sustaining that Reduviasporonites has not any distinctivebiomarkers of either fungi or algae.

Afonin et al. (2001) also proposed an algal affinity and demonstratedthat the spores assigned to Reduviasporonites are more closely alliedto the Zygnematalean algae Spirogyra Link and/or Mougeotia Agardhof the Division Chlorophyta (green algae), a genus having a widedistribution.

In this study, we noted several common traits between themorpho-logical characteristics of Reduviasporonites chalastus and those ofSpirogyra and/or Mougeotia (Plate IV). The dark coloured inner bodiesare similar to the “accumulation bodies” of dinoflagellate cysts. How-ever, differently from Spirogyra, the inner bodies of R. chalastus do notseem spirally organised, but they appear as a granular mass lining thecell walls. Anyway, lingering doubts indicate that more detailed studiesare still needed, for example, for understanding the reason of the lack ofzygospores in several levels rich in R. chalastus. Commonly, in Spirogyra,the zygospores are characterized by a rounded shape and very thick,robust wall that made them more resistant than the filaments. Hence,they should be better preserved and easier to find than the filaments.In assemblages with abundant R. chalastus, no zygnematacean sporessuch as Brazilea (supposedly a fossil spore of the genus Spirogyra), norTetraporina (supposedly a fossil spore form of the genus Mougeotia),norMaculatasporites (considered by Grenfell, 1995, to represent sporesof any of the recent genera of Zygnema, Zygnemopsis, Zygogonium) havebeen found. Finding the affinity of R. chalastus to Spirogyra and/orMougeotia not very convincing, we tentatively compared it to othergenera of chlorophycean algae. Thus, we found R. chalastus showingclose morphological similarities to some algae belonging to the OrderTrentepohliales (Plate IV). The trentepohliacean algae are not aquatic,but rather subaerial, growing on humid soil, rocks, tree bark, leaves,stems and fruits. The presence of sporopollenin-like substances in thecell walls, as well as special structures of carbohydrates and alcohols(Patterson and Van Valkenburg, 1991), probably is an adaptive featureagainst desiccation in the subaerial habitat. Some species of theTrentepohliales are endophytic or parasitic, whereas others grow inclose association with lichen forming fungi (Chapman and Waters,2002). According to López-Bautista et al. (2002), the sporopollenin-

Plate I. Bisaccate pollen from Tesero section. Scale bar indicates 10 μm.

1. Platysaccus sp. (slide AS4, 97.7/45.5)2. Jugasporites delasaucei (Potonié and Klaus) Leschik, 1956 (slide AS4, 110/38)3, 8. Klausipollenites shaubergeri (Potonié and Klaus) Jansonius, 1962 (3: slide AS1, 374. Lueckisporites virkkiae Potonié and Klaus, 1954 (slide AS4, 38.1/119.2)5. Lueckisporites sp. (slide AS4, 35.5/103.5)6. Klausipollenites sp. (slide AS25, 41.2/108.9)7, 12, 13, 15. Alisporites spp. (7: slide AS1, 29.9/111.1; 12: slide AS4, 32.2/100.3; 13: slide AS49. Protohaploxypinus limpidus (Balme and Hennelly) Balme and Playford, 1967 (sli10. Striatopodocarpites sp. (slide AS4, 32.2/107.2)11. Striatopodocarpites fusus (Balme and Hennelly) Potonie 1958 (slide AS4, 30.7/1014. Pinuspollenites sp. (slide AS4, 25.4/97.7).

Plate II. Bisaccate pollen from Tesero section. Scale bar indicates 10 μm. (see on page 8)

1–4. Sulcatisporites spp. (1: slide AS4, 30.6/97.2; 2: slide AS5, 26.6/120.4; 3: slide AS25. Alisporites landianus Balme, 1970 (slide AS1, 26/115.2)6. Alisporites sp. (slide AS2, 34.1/119.9)7, 15. Lueckisporites virkkiae Potonié and Klaus, 1954 (7: slide AS23, 30/101; 15: slide8, 9. Klausipollenites shaubergeri (Potonié and Klaus) Jansonius, 1962 (8: slide AS3, 210. Lunatisporites noviaulensis (Leschik) Foster, 1979 (slide AS4, 37.1/117)11. Klausipollenites decipiens Jansonius, 1962 (slide AS3, 46.6/118.1)12. Lunatisporites pellucidus (Goubin) Balme, 1970 (slide AS4, 65.5/132)13. Striatopodocarpites fusus (Balme and Hennelly) Potonie 1958 (slide AS4, 49.9/1114. Protohaploxypinus microcorpus (Schaarschmidt) Clarke, 1965 (slide AS4, 42.2/10

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like substances in the cell wall of trentepohliacean algae would havefacilitated their preservation as fossils.

The trentepohliacean alga Trentepohlia Martius is organised inbranching filaments and, as in Reduviasporonites chalastus, the cell-shape is spherical to ellipsoidal; although its chloroplasts can assumealso a spiral shape, in general they are arranged as a globular massin the centre of each cell like often the cell-infill inner bodies ofR. chalastus. Different to that in Spirogyra, the zoosporangia of Trentepohliaare mainly diffused by the wind. This feature could explain the reason oftheir absence in the association with R. chalastus in the Tesero levels.

Although Reduviasporonites chalastus shares some morphologicalfeatures with the Trentepohliales algae, and shows similarity toZygnematales algae, it displays additional traits typical of other greenalgae. For this reason, we think that it is a chlorophycean alga probablyliving in brackish waters.

Wood (1995), Eshet et al. (1995), Visscher et al. (1996), Steineret al. (2003), and Sandler et al. (2006) stressed the importance ofthe so called “fungal event” as a reflection of environmental changeacross the Permian–Triassic boundary and thus considered as a chrono-stratigraphic marker for worldwide correlation. On the contrary, Fosteret al. (2002) observed that Reduviasporonites chalastus appears to havethrived outside the postulated time of mass extinction, consequently itcannot be used as “practical” chronostratigraphic marker. We agreewith this consideration because we often find R. chalastus or similarorganic remains in a stressed palaeoenvironment of different age.

In the Tesero Member the proliferation of Reduviasporoniteschalastus could be related to the presence of hypersaline ponds in flatcoastal areas and its bloom strictly linked to the ecological parametersof the depositional palaeoenvironment. Consequently it should be notused as practical biomarker for worldwide correlation of the PTB.

6. Conclusions

The present study was carried out in order to correlate the palyno-logical assemblage across the Permian–Triassic boundary ratified byconodont zones in two of the better known Permian–Triassic sectionsof the Southern Alps. Additionally the origin and the palaeoecologicalpreferences of Reduviasporonites chalastus and the inference on itsworldwide distribution across the PTB are discussed.

The Tesero section yielded well preserved and diversifiedmicrofloristic assemblages in respect to the Bulla sectionwhere the pal-ynological content is less abundant. In the Bulla section, indicated as theparastratotype of the PTB for the western Tethyan domain (Farabegoli

.8/48.5; 8: slide AS21, 42/110)

, 30/100; 15: slide AS4, 27.7/99.1)de AS4, 25.5/97.8)

4.1)

5, 28/99; 4: slide AS30, 34.4/118)

AS1, 28.8/109)6/95.1; 9: 28.6/106)

7.1)0).

riassic of the Tesero and Bulla sections (Western Dolomites, Italy) and/dx.doi.org/10.1016/j.revpalbo.2014.10.003

Plate I

7A. Spina et al. / Review of Palaeobotany and Palynology xxx (2014) xxx–xxx

Please cite this article as: Spina, A., et al., Palynology of the Permian and Triassic of the Tesero and Bulla sections (Western Dolomites, Italy) andconsideration about the enigmatic ..., Rev. Palaeobot. Palynol. (2014), http://dx.doi.org/10.1016/j.revpalbo.2014.10.003

Plate II (caption on page 6).

8 A. Spina et al. / Review of Palaeobotany and Palynology xxx (2014) xxx–xxx

Please cite this article as: Spina, A., et al., Palynology of the Permian and Triassic of the Tesero and Bulla sections (Western Dolomites, Italy) andconsideration about the enigmatic ..., Rev. Palaeobot. Palynol. (2014), http://dx.doi.org/10.1016/j.revpalbo.2014.10.003

Plate III. Bisaccate pollen from Bulla section (Scale bar indicates 10 μm).

1. Alisporites sp. cf. splendens (Leschik) Foster, 1979 (slide MC10, 95.2/30.4)2. Jugasporites sp. (slide MC8, 33/102)3, 4. Alisporites sp. (3: slide MC 12, 37/110.3; 4: slide MC10, 102/25)5. Lueckisporites sp. (slide MC6, 49/98)6. Jugasporites delasaucei (Potonié and Klaus) Klaus, 1963 (slide MC8, 44.5/109)7. Klausipollenites sp. (slide MC10, 102/17.5)8, 9. Lunatisporites noviaulensis (Leschik) Foster, 1979 (8: slide MC12b, 25.6/108.9; 9: slide MC12, 29.9/101.1)10, 11. Lunatisporites pellucidus (Goubin) Helby, 1972 (10: slide MC12, 35.4/100; 11: slide MC12, 33.3/102)12 Lunatisporites sp. (slide MC12, 31/100).

9A. Spina et al. / Review of Palaeobotany and Palynology xxx (2014) xxx–xxx

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Plate IV.

1–7, 13–17. Reduviasporonites chalastus (Foster) Elsik, 1999 from the Tesero section.8, 9, 12. Some examples of Spirogyra (from Bourelly, 1990; 8: filament with aplanospore; 9: filament with spirally arranged chloroplasts; 12: zygote).10, 11. Some examples of Trentepohlia (from Bourelly, 1990).

10 A. Spina et al. / Review of Palaeobotany and Palynology xxx (2014) xxx–xxx

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et al., 2007), the putative PTBhas been placed in the TeseroMember andcorrelated to theupper TeseroMember in the Tesero section (Farabegoliet al., 2007).

In this stratigraphic framework and extending the above consider-ations also to the Tesero section, the features of the palynologicalassemblages can be summarized as follows:

– The microflora content from the Bellerophon Fm. is dominated bynon-taeniate and taeniate bisaccates that in the Tesero section,where the assemblage is more diversified, are mostly representedby bisaccate taeniate (e.g. Protohaploxypinus spp., Lunatisporitesspp., Lueckisporites spp., Striatoabieites spp. and Striatopodocarpitesspp.) and non-taeniate (e.g. Klausipollenites spp., Alisporites spp.and Jugasporites delasaucei) and, more rarely, trilete spores associat-ed with few Reduviasporonites chalastus.

– In the Bulla section, the Bellerophon Fm. yields a microflora less di-versified and only containing bisaccates such as Lueckisporites sp.,Lunatisporites pellucidus and Jugasporites delasaucei. No remains ofReduviasporonites chalastus were found. The different amount ofthe total organic matter (palynomorphs including) could be relatedto a more distal position from the land masses of the Bulla deposi-tional environment. Nevertheless the blooming of taeniate andnon-taeniate sporomorphs, among which Lueckisporites sp. andJugasporites delasaucei is indicative of a late Permian age.

– The palynogical content of the Tesero Member, in both the investi-gated section is generally rather scarce or absent due to the lack ofa lithology that might preserve palynomorphs such as oolitic lime-stone which is the characteristic facies of this member.

– The lower-middle portion of the Tesero Member, in the Teserosection, yielded palynofacies dominated by Reduviasporoniteschalastus associated with rare sporomorphs (Fig. 3). The amountof R. chalastus drastically decreases in the uppermost part ofthe Tesero Mb, (above the supposed PTB) and in the lowerMazzin Member. Here few pollen as Lueckisporites virkkiae andKlausipollenites schaubergeri are associated with long rangingspores. The occurrence of few “Permian type” pollen within theselower Triassic strata can be explained in two ways: a) these speciescan range into Early Triassic as suggested by several authors (i.e.Ouyang and Utting, 1990; Mangerud, 1994); b) they are reworkedfrom the underlying strata.

– The good preservation of the green alga Reduviasporonites chalastusis probably due to the organic composition of the wall, highlyresistant to processes such as bacterial degradation and chemicalbreakdown (i.e. oxidation; Tappan, 1980) that characterized the de-positional environment of the Tesero Member. Its palaeoecologicalpreference should be related to anomalous water salinity rangingfrom brackish to hyperhaline. Being the distribution and abundanceof R. chalastus controlled by ecological parameters at local and largescale it cannot be further used as a chronostratigraphic marker. Theabundance of R. chalastus in the Tesero section and its absence in theBulla section should confirm the distal position as depositional envi-ronment of the latter. Due to its probable ecological preference,R. chalastus could easier colonize and grow in coastal ecosystems.

Acknowledgements

This paper is dedicated to Jan Jansonius who passed away after along illness on January 25, 2013. He was 84 years old. A.S. met himwith J.U. in Calgary in December 2001 during her PhD. Jan with Johnhelped Amalia to grow not only from the scientific point of view butalso as a human being.

We are indebted to Prof. Mike Stephenson (British GeologicalSurvey) who reviewed this manuscript. Many thanks are due toDr. Giovanna Giorgetti (Siena University) for the SEM images andDr. Gianluca Cornamusini (Siena University) for the optical microscopephotos. We are grateful to Prof. Bruno Granetti (Perugia University) for

Please cite this article as: Spina, A., et al., Palynology of the Permian and Tconsideration about the enigmatic ..., Rev. Palaeobot. Palynol. (2014), http:/

the interesting and helpful discussion about themorphological featuresof Spirogyra and Trentepohlia.

Appendix 1. List of taxa

– Pollen:Bisaccate taeniate:

Hamiapollenites sp.Lueckisporites parvus Klaus, 1963Lueckisporites virkkiae Potonié and Klaus, 1954Lueckisporites sp.Lunatisporites albertae Jansonius, 1962Lunatisporites hexagonalis Jansonius, 1962Lunatisporites interruptus Jansonius, 1962Lunatisporites noviaulensis (Leschik) Foster, 1979Lunatisporites pellucidus (Goubin) Balme, 1970Lunatisporites sp. cf. transversundatus Jansonius, 1962Lunatisporites sp.Protohaploxypinus limpidus (Balme and Hennelly) Balme and

Playford, 1967Protohaploxypinus microcorpus (Schaarschmidt) Clarke, 1965Protohaploxypinus samoilovichiae (Jansonius) Hart, 1964Protohaploxypinus sp.Striatoabieites sp.Striatoabieites richteri (Klaus) Hart, 1964Striatopodocarpites communis (Wilson) Hart, 1964Striatopodocarpites fusus (Balme and Hennelly) Potonie, 1958Striatopodocarpites sp.

Bisaccate non-taeniate:Alisporites landianus Balme, 1970Alisporites sp. cf. splendens (Leschik) Foster, 1979Alisporites splendens (Leschik) Foster, 1979Alisporites tenuicorpus Balme, 1970Alisporites spp.Jugasporites delasaucei (Potonié and Klaus) Leschik, 1956Jugasporites sp.Klausipollenites decipiens Jansonius, 1962Klausipollenites forma Jansonius, 1962Klausipollenites schaubergeri (Potonié and Klaus) Jansonius, 1962Klausipollenites vestitus Jansonius, 1962Klausipollenites sp.Pinuspollenites sp.Sulcatisporites kraeuseliiMädler, 1964Sulcatisporites spp.

Monosaccate:Cordaitina sp. cf. vulgaris (Zauer) Varyukhina, 1971

Monocolpate:Cycadopites follicularis Wilson and Webster, 1946Sverdrupollenites agluatus Utting, 1994

– Trilete spores:Acavate:

Calamospora sp.Dictyotriletes sp.Leiotriletes adnatus (Kosanke) Potonié and Kremp, 1955Punctatisporites fungosus Balme, 1970Simeonospora sp. cf. minuta (Jansonius) Utting, 1994

Cavate:Densoisporites sp.Proprisporites pocockii Jansonius, 1962

– Incertae sedis:Reduviasporonites chalastus (Foster) Elsik, 1999

riassic of the Tesero and Bulla sections (Western Dolomites, Italy) and/dx.doi.org/10.1016/j.revpalbo.2014.10.003

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