C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56Fig. 1. Devonian succession (Ponta Grossa and Furnas Formations) of the Paraná Basin, Paraná State, Brazil. Numbers represent the outcrops studied, organized in the table, and their respective GPS data. 45
46 C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56 occurrence of these lingulid associations through the Palaeozoic and until Recent times, there is some discrepancy between Palaeozoic and post-Palaeozoic records, especially Cenozoic to Recent, which may represent a taphonomical megabias (Kowalewski, 1996a). Recently, some authors proposed models to explain the lingulid shell-beds formation and requirements worldwide (e.g. Emig, 1986; Kowalewski, 1996a,b; Kowalewski and Flessa, 1996; Emig, 1997; Emig and Gutiérrez-Marco, 1997; Aceñolaza et al., 2003; Zonneveld et al., 2007). These models have proven useful for interpreting con- centrations composed mostly of lingulids (although their utility may be limited in case of some obolids which were likely semi-infaunal, see Zhang et al., 2007, 2009). In this study, some taphonomical and palaeoenvironmental aspects on Devonian lingulids are addressed. These include: ﬁdelity of the fossil record, their susceptibility to transport and reworking, degree of time averaging, environmental conditions required for their preservation, and their ecology. Under- standing the genesis of the fossil occurrences of an animal, which represents ecology, biology, and shell mineralogy that all differ dramatically from those of the co-eval fauna (and consequently should result in different taphonomic responses to post-mortem agents),Fig. 2. Stratigraphic column of Tibagi 1 (0–2 m) and Tibagi 2 (2–25 m) outcrops. All should provide insights into taphonomic and environmental processesoutcrops that show the abundant presence of lingulids have very similar lithologies. that led to the formation of the studied fossiliferous units.Fig. 3. Identiﬁcation and age of the studied outcrops according to the sequence stratigraphic setting of the Devonian succession of the Paraná Basin in central Paraná State, Brazil(Bergamaschi, 1999), and correlation with Members of the Ponta Grossa Formation.
C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56 472. Material and methods mation, d) bioclast position on the bedding plan, e) fossilization type, f) degree of fragmentation, and g) miscellaneous characteristics (e.g. Some samples for this study (MPI 0001–0599, MPI 1839–1902, borings, drill holes, sedimentary structures, ﬂattening or deformation).MPI 2535–2659, MPI 3000–3197, MPI 4036–4438, MPI 4451–4522, Percentage graphs were constructed using the high resolution dataand MPI 4556–4582) are stored at Universidade Estadual de Ponta collected from the Tibagi 2 outcrop. These were generated usingGrossa (UEPG, Departamento de Geociências, Laboratório de Paleon- Microsoft® Ofﬁce Excel 2003, and denote: a) the percentage oftologia, Ponta Grossa, PR, Brazil), being catalogued as Bosetti-2004 occurring taxa (Fig. 6A), and b) lingulid valve positions in relation toﬁeld collection, Zabini-2007 ﬁeld collection, and as Laboratory the bedding plan (Fig. 6B).collection. The latter includes samples collected recently in the city Lingulid abundance is expressed using a qualitative terminology —of Tibagi (Paraná State, outcrops Tibagi 1 and 2 — Figs. 1, 2). Other rare, present and abundant. These terms are deﬁned as follows: rarespecimens (MPI 6567–6680) are part of the Bosetti-1989 ﬁeld (∼1–2 specimens observed in a 10 m lateral bed exposure); presentcollection at Universidade Federal do Rio Grande do Sul (UFRGS,Departamento de Paleontologia e Estratigraﬁa, Porto Alegre, RS,Brazil). Additionally, specimens described by Petri (1948) were alsoused. The samples with lingulids were taken from 12 mid-east regionoutcrops in Paraná State (Fig. 1). The localities Campus, Casa de Pedra,Curva I, Curva II, Fazenda Rivadávia, Jaguariaíva (railroad), Type-section of the Tibagi Member, Vila Francelina and Cescage were alsoanalysed, but did not contain a representative number of lingulids.The outcrops are correlated to sequences B, C, D and E of Bergamaschi(1999) (Fig. 3). Approximately 7000 valves were analysed, withindividual samples consisting of one or more fossil specimens. Mostoutcrops were sampled following the methodology proposed bySimões and Ghilardi (2000), Bosetti (2004) and Ghilardi (2004),which involves the demarcation of grid squares (Fig. 4) and ﬁeldcharts, with the aim of acquiring high resolution data (Fig. 5). During the initial stage of the research, a taphonomical databasewas compiled including the following variables: a) number and typeof taxa in the sample, b) lithology of the host rock, c) biometric infor- Fig. 5. Demarcation of the grid square at Tibagi 2 outcrop (A and B). At this outcrop three grid squares were marked, the ﬁgure represents grid square number one (see its location on Fig. 3). A) Demarcation of the grid square before collect. B) The grid square after collecting of the fossils. Note that the area within the space demarcated by theFig. 4. A) Tibagi 2 outcrop, in perspective. B) Schematic drawing of the same outcrop, lines was combed in the search for fossils. C) Representation of the distribution of thewith demarcation of grid squares 1, 2 and 3. fossils found in the ﬁeld work, within grid square one.
48 C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56(∼ 5–10 specimens observed in a 10 m lateral bed exposure);and abundant (∼ 10–50 specimens observed in a 10 m lateral bedexposure). The lingulid shells are preserved as internal, external or compoundmoulds and impressions. The original shell material has not beennaturally preserved. Moulds usually display the characteristic orna-mentation of lingulids, such as growth lines and, more rarely pre-served muscle scars. Traditionally, ever since they were ﬁrst found in 1876 by Orville A.Derby, the brachiopods studied here have been referred to Lingula(Bosetti et al., 2008). This terminology dates back to Darwins work,which coined the term “living fossils”, and includes the genus Lingula asone of its examples (Emig, 2008). The last speciﬁc revision of the PontaGrossa Formation lingulids was performed by Bosetti (1989), whoretained the traditional taxonomy and used the genus name Lingula. Nomajor revision of these fossils has been attempted after Bosettis work Fig. 7. Lingulid oblique to the sediment, showing its anterior part faced up, and the(Zabini et al., 2007). As proposed by Biernat and Emig (1993) and Emig “linguliform” shape of the shell, somehow deformed by the compression of the sediment.(2003) the genus Lingula appeared only in the later Cretaceous or theearliest Tertiary. C.C. Emig (personal communication; 2008) suggests Dignomia. Seemingly, Lingulidae and Obolidae families are preservedthat the lingulid fossils here studied, mostly likely, belong to the genus together, with no apparent differences in their ecological requirements and/or mode of life, once they are found together in the same bedding planes. A comprehensive investigation, including a taxonomic revision, based on the new data from Emig (2003), is being prepared by the authors and will be published elsewhere. At present, it is preferable to use the term “lingulid” to refer to these fossils. As described by Emig (2003), these lingulid fossils show a typical “linguliform” shell shape, and therefore are considered to have been infaunal burrow dwellers (Cherns, 1979; Emig, 1997, 2003). The fact that these fossils are found in small numbers, perpendicular to the sediment, with its anterior part oriented up (Fig. 7), supports this interpretation.Fig. 6. Data charts. A) Relative abundance of major taxonomic groups through sampledgrid squares (1, 2 and 3) of Tibagi 2 outcrop. n = 2001; B) Percentages of lingulids in life Fig. 8. The chronostratigraphic chart of the Paraná Basin, showing the second-orderposition, oblique and concordant to the sediment, through sampled grid squares (1, 2 depositional sequences (simpliﬁed after Milani et al., 1994). The dashed rectangleand 3) of Tibagi 2 outcrop. n = 1783. indicates the area studied in this paper.
C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56 49Fig. 9. Specimens showing exceptional shell preservation. A, B, D and F are photographs of concretions containing lingulids at various angles, collected from outcrop km 217Transbrasiliana. Figures C, E and G correspond to the photographs above and portray the thicknesses of the valves. Figures H, I, J and K represent both axes of shell orientation. Considering the fact that the Paraná Basin was a shallow intracratonic into six third-order sequences, labeled A to F. Based on that work andsea in the Devonian, it is more convenient to organize palaeoenviron- the data of Bosetti (2004) and Soares (2007), the outcrops used in thisments using a general bathymetric nomenclature proposed by Reading paper can be correlated to the sequences B, C, D and E (Fig. 3).and Collinson (1996). The facies delineation outlined by Walker and Plint(1992) is used here, as coarse (shoreface or lowermost intertidal) to ﬁne 4. Acquired data and results(offshore or deeper subtidal) sedimentary facies pattern; as shown in theﬁgures, and explained in the discussion part. Lingulids were dominant fossils within twelve out of twenty-one analysed outcrops (Fig. 6A). Following Kidwell (1991) and Kidwell et al. (1986) the lingulid assemblages of the twelve outcrops bearing abundant3. Geological setting lingulids can be described as: concordant valves, pauciespeciﬁc pave- ments and sometimes stringers, matrix-supported, with simple internal The Paraná Basin is a huge intracratonic basin on the South- structure. Regarding sampled fossils, lithologies, and depositionalAmerican platform, located in southernmost Brazil and north/north- systems, the following observations can be made:western Uruguay, with its portions extending also into Paraguay andArgentina (Fig. 1). The basin covers an area of about 1,700,000 km2, has – Most lingulid valves (approximately 90%) are concordant to thea NE–SW elongated shape, and is approximately 1750 km long and bedding plane (Fig. 6B);900 km wide (Petri and Fúlfaro, 1983). – The valves are mainly ﬂat, as their convexity was not noticeable; The sedimentary inﬁll of the basin from Ordovician to Cretaceous – The degree of valve disarticulation was difﬁcult to evaluate due towas controlled by tectonic–eustatic cycles (i.e., rise and fall of the base their extreme ﬂattening; it is assumed that all valves parallel to thelevel) linked to Late Palaeozoic orogenic events caused by subduction bedding plane represent disarticulated specimens, as in almost alland terrain accretion along the southwestern margin of the lingulid beds (Emig and Gutiérrez-Marco, 1997; Zonneveld et al.,Gondwana continent. The process of the sedimentary inﬁlling was 2007, among others).largely arrested by the onset of the Mesozoic rifting of the South – Dorsal and ventral valves could not be differentiated in oneAtlantic (e.g. Zalán et al., 1990; Milani et al., 2007). The prevalence of another in the studied material;tectono–eustatic cycles (Milani et al., 2007), has generated a – The degree of fragmentation is variable, ranging from entire valvesstratigraphic record that is marked by numerous interruptions to fragments smaller than 0.5 cm;brought about by erosion and non-deposition. Milani et al. (1994, – Some lingulid shells exhibit exceptional preservation, obtaining1998) argued that the inﬁll of the basin consisted of six second-order the thickness of the valves was possible in certain cases (∼ 0.1–depositional sequences, ranging in age from the Late Ordovician to the 0.2 mm, Fig. 9).Late Cretaceous (Fig. 8). The stratigraphic interval studied herein – Some samples display distinguishable patterns of distribution,occurs in the second sequence of Milani et al. (1994); the so-called showing rosette and bimodal stringer spatial arrangementsParaná Sequence, ranging from the Late Silurian to the Late Devonian (Plate I);(Fig. 8). This sequence is represented, lithostratigraphically, by the – The studied layers represent silty shales, and very ﬁne-grainedFurnas and Ponta Grossa formations. Lingulid brachiopods occur in siltstones. Except for Vila Placidina outcrop, lingulids were rarelythe Ponta Grossa Formation. found in black shales; Many outcrops of the Ponta Grossa Formation are located in the – The main sedimentary structures are plane bedding, micro-mid-east region of the Paraná State (Fig. 1). The unit is represented by hummocky bedding, and trace fossils (e.g. Zoophycos);siliciclastic rocks that formed in a shallow epicontinental sea that was – In general, the studied units reﬂect conditions of a more distallocated at 50° to 60° palaeolatitude, with a predominantly temperate deposition in relation to the shore; usually they represent theclimate. The Ponta Grossa Formation is characterized by shoreface, lower shoreface to transitional offshore zones above average stormtransitional offshore and offshore deposits. It is mainly formed by grey wave-base (Fig. 10).and locally dark, bituminous and silty shales, with minor ﬁne-grainedsandstone beds. Lithostratigraphically, the Ponta Grossa Formation is In outcrops that reﬂect shallow water environmental conditions,subdivided into three members: Jaguariaíva, Tibagi, and São Dom- the occurrence of lingulid valves is extremely rare. Such is the case iningos (Lange and Petri, 1967). Bergamaschi (1999) and Bergamaschi sequence C (Tibagi Member), represented by the basal portion ofand Pereira (2001) divided the Paraná Sequence of Milani et al. (1994) Lambedor outcrop and the type-section of Tibagi Member where,
50 C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56until this study, lingulid valves had not been found. It is also The results demonstrate that the studied pavements include anoteworthy that brachiopods belonging to the rhynchonelliform larger number of valves parallel to the bedding planes, followingbrachiopods (e.g. Australocoelia tourteloti, Derbyina whitiorum and taphonomical pathway 2 of Kowalewski (1996a). According toAustralospirifer iheringi) are not usually found in association with large actualistic data the lingulid shells are rapidly degraded and mechan-numbers of lingulid valves, and vice-versa. ically abraded (Kowalewski, 1996a; Emig and Gutiérrez-Marco, 1997; Aceñolaza et al., 2003). Thus, their record must be understood as a consequence of event-concentration of shell assemblages accumulat-5. Discussion ed in a very short time period or transported for only short distances (Emig, 1986; Kowalewski, 1996a; Emig, 1997; Emig and Gutiérrez-5.1. Palaeoecology and taphonomy Marco, 1997). Investigations of Mesozoic and Recent lingulids (Kowalewski, 1996a) demonstrate that it is very rare to have lingulid5.1.1. Paleoecological approach valves preserved in the post-Paleozoic fossil record. Hence, why Despite the probable occurrence of two families among the Palaeozoic, and in this case, the Devonian lingulids studied here, werelinguliform taxa studied here, we consider all as having the same – preserved, including even fragmented valves? Do these resultsinfaunal – life habit and similar palaeoecological requirements. These undermine the validity of actualistic data?arguments are based on our current knowledge on their biology, Kowalewski (1996a) proposed two explanations for the betterecology, and taphonomic signatures. The coarse-grained to clay preserved older fossils (1) an increase in the intensity of the externalsiltsones and the black clay shale are typically found in the marine taphonomical agents as time passes (bioturbation, bioerosion, breakingparts of the analysed Devonian succession, and their associated fauna predation, sediment chemistry and frequency of burial events), which is(trilobites, molluscs and other typical marine invertebrates) also difﬁcult to demonstrate or (2) a decrease of fossilization potential ofsupports the marine origin of the deposits (see Simões et al., 2000; more recent lingulids, the most likely explanation for this would be theRodrigues et al., 2003, for similar interpretations). Moreover, decreasing thickness or less intense biomineralization of lingulid valves.fossilization of lingulid valves in marine conditions has been already The measurements taken on some of the best preserved specimensdemonstrated by several authors (Cherns, 1979; Pickerill et al., 1984; available to us indicate that the valve thicknesses of the DevonianPashin and Ettensohn, 1992; Zonneveld et al., 2007). lingulids was around 0.1–0.2 mm (Fig. 9). Their valves were probably According to the results (especially, lithologies, sedimentary organophosphatic and their mode of life was likely infaunal. However,structures, and taphonomic data), it is reasonable to assume that the we do not know the real proportions of the organic and the mineralizedtwelve outcrops that produced an abundant record of lingulid valves contents of the original shell. Assuming that their external morphologyrepresented environmental conditions varying from shoreface (lower- and life habit were the same of the extant ones, we can say that probablymost intertidal), with coarse to medium-grained siltstones, to transi- the intrinsic factors involved in generating these shell accumulationstional offshore zones (shallow subtidal) typiﬁed by clay silstones were similar than the modern ones. Yet it is clear that more data(Fig. 11), where lingulids in life position are occasionally present. about the shell geochemistry could improve our knowledge as to the taphonomy of the studied associations.5.1.2. Taphonomical approach Therefore, we propose that there are similarities between actualistic The death of lingulids can be linked to a number of different factors data and those from the Devonian in the Paraná Basin, in the context ofaccording to Ferguson (1963), Paine (1963), Emig (1981a,b, 1986), the intrinsic factors involved in generating these shell accumulations.Hammond (1983) and Kowalewski (1996a), including biological The possibility that the extrinsic factors may have been unusual, and notfactors (e.g. diseases, senility and predation), chemical factors (e.g. comparable to typical conditions known from modern marine settings,salinity and anoxic water) and physical factors (e.g. temperature and should be taken into consideration. This point has already been raised bysedimentation rate). After death, lingulid valves can follow three Speyer and Brett (1988), who highlighted the limited circulation ofdifferent taphonomical pathways that are dependent upon whether water and the oxygen-stratiﬁed epeiric sea basins. Such epeiric depositsthe organism remains or is removed from its burrow (lingulids are dominate the Palaeozoic fossil record of many regions, including theinfaunal, the burrow of a lingulid is considered to be where in the Paraná Basin.sediment the animal actually lived). According to Kowalewski How can it be explained that the taphocoenoses studied here are(1996a), who studied Glottidia palmeri in the Colorado River Delta not only formed by organisms in life position preserved by obrution(Mexico), the three taphonomical pathways are possible: (1) the (Simões et al., 1998a,b; Nunes, 1999), but also by accumulation ofshells remain in the burrow, and the organic parts of the animal abundant fragments, including fragments preserved together withdecompose. Such shells can remain in the sediment for more than a disarticulated, but complete valves?year but no longer than a decade. (2) The shell is removed from its According to Emig (1990) and Kowalewski (1996a) thin valves ofburrow, before or immediately after death. Its disarticulated valves G. palmeri are easily fragmented even in low energy environments.can be deposited at the surface and will not remain for long at the The degree of fragmentation in Devonian lingulids is variable: withinsediment-water interface. Mechanical disintegration occurs in a matter the same hand sample, specimens can range from entire valves toof days and can be postponed for no longer than a few weeks (see also fragments less than 0.5 cm in their maximum dimension. The mostEmig and Gutiérrez-Marco, 1997; Aceñolaza et al., 2003; Zonneveld likely explanation is that fragmentation must have occurred byet al., 2007). (3) Storms deposit the recently dead lingulids into the reworking, through orbital wave movement above fair weather wave-supratidal zone and the soft parts undergo a dehydration process, base, and also by the action of more episodic storms (Emig, 1997;mummifying them. This type of preservation has not been found in the Emig and Gutiérrez-Marco, 1997; Aceñolaza et al., 2003) that could befossil record. responsible for mixing and burying the assemblages. The degree ofPlate I. Samples showing lingulid valves in rosette and stringer arrangements.A). A typical rosette arrangement (sample MPI 2647);B–F. Variations of the rosette shape; photos are from MPI 2630, 2637, 2628, 2643 and 2628 samples, respectively.G–H). Stringer arrangements;G). A bimodal stringer arrangement and, on the right side, a rosette arrangement (sample MPI 2631); and H) two almost parallel stringer arrangements of valve fragments (sample MPI 2623).
C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56 51
52 C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56Fig. 10. A–K) The twelve “lingulid bearing” outcrops. The column to the left, which is not in scale, shows the relative sea level changes related to the outcrops. All outcrops record very similar environmental conditions, which together with thesedimentary structures and the taphonomical analyses of the fossils, represent shoreface (lowermost intertidal) to offshore (deeper subtidal) zones.
C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56 53fragmentation of the valves is also increased with the increase in clay (Plate I). As described by Parsons et al. (1988), stringer pavements arecontent. At the distal part of the transitional offshore zone the degree probably the result of reworking of the valves in deep environments.of fragmentation increases. However, disarticulated non-fragmented Such dispositions of the valves were possibly a distal response/reworkingvalves are still present (Fig. 11). of a major storm event in the shoreface zone. Rodrigues et al. (2003) and The occurrence of different valve sizes and associated fragments Simões et al. (2000), who examined conulariid taphonomy in the sameindicates that if there was transport, it was probably minimal. This is basin, noted that this was probably an epeiric muddy sea stronglybecause hydraulic size-sorting is not evident in lingulid associations inﬂuenced by episodic storms. Therefore, major erosion of the sea ﬂoor(i.e., shell fragments of various size together with complete shells by severe storm events may have thrown shell fragments below theoccur in the same pavements). Hence, the fossils represent a storm wave-base. Here, these became associated with entire valvesparautochthonous (sensu: Kidwell et al., 1986) assemblage — the recently removed from burrows, as indicated by the record of valvesincipient transport prevented the bioclasts from being removed from and valve fragments in black mudstone from the offshore zone.their original area of distribution or habitat. Despite the evidence of transport and reworking of the bioclastic Time averaging, at least to some extent, may have occurred. This is material, this transport was probably never energetic enough tobecause variably sized fragments (probably representing valves that hydraulically select the different-sized lingulid valves, and it onlypersisted for days or weeks around the sediment–water interface; see reoriented the valves and the valve fragments in stringers or rosetteKowalewski, 1996a) co-occur with disarticulated, complete valves shapes, along the pavements (Fig. 11A–D). Furthermore, the thin valves(likely removed from their burrows just before burial). of the studied lingulids, present from the shoreface to the distal part of Based on the taphonomical analysis of the material here collected we the transitional offshore zone, must have been preserved through thecan say that the original habitat of the lingulids ranged from shoreface action of fast, episodic sedimentary accumulations (Fig. 12). Similarsettings down to the transitional offshore, with preference for the cases were studied by Emig (1997) and Aceñolaza et al. (2003), whoshoreface zone (Fig. 11). The high fragility of these bioclasts reinforces worked on the genera Lingula spp. and Lingulepis sp., respectively.this hypothesis, because the valves could not be transported for longdistances (Emig, 1986; Kowalewski, 1996a; Emig, 1997; Emig and 5.2. Taphofacies modelGutiérrez-Marco, 1997). Therefore, they probably were preserved closeto their original habitat. Parallel valves per se do not indicate the original Based on the current data, a lingulid taphofacies for the Devonianhabitat (e.g. Zonneveld et al., 2007), but in this case, these are associated sequences B, C, D and E (Bergamaschi, 1999) is here proposed.with low numbers of lingulids in perpendicular or oblique orientation, According to Speyer and Brett (1986) a taphofacies is deﬁned wheni.e., in situ lingulids (see Kowalewski, 1996a). suites of sedimentary rocks are characterized by particular combina- The rosette and stringer pavements are probably the result of tions of preservational features of the contained fossils. As mentionedreworking, and have not been described for lingulid valves previously above, the lingulids studied in this paper are ubiquitous in someFig. 11. Distribution of the fossil record of lingulids. The bathymetric proﬁle was constructed based on sedimentological and taphonomical data. A) Lingulids preserved in lifeposition; B) entire valves preserved in a rosette arrangement; C) lingulid valves arranged in bimodal stringers; D) the arrows show the presence of ostracodes deposited above thestringer, indicating bioclast deposition, which occurred by the deposition of the heavier elements followed by successively lighter elements; and E) valve fragments preserved in astringer.
54 C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56Fig. 12. Distribution of lingulid patches and fragments. The frame shows the reworking of the bioclastic material and the processes that generated the fossil record of the group. A)Reworking of bioclasts above the average storm wave-base and lingulid distribution along the shoreface and transitional offshore. B) The storm event removed entire or fragmentedvalves and suspended the bioclasts; it removed the bioclastic materials that were in the sediment-water interface or in the ﬁrst levels of the sediments from the inferior shorefaceuntil reaching average storm-wave-base. C) After the storm, the bioclasts were deposited at the sea bottom. Note that some lingulids that are still in life position can be preserved assuch. Some currents can select and throw valves (entire and fragmented) to the distal portion of the transitional offshore.outcrops and make up the most common fossils in some parts of the sequence C (Bergamaschi, 1999), which mostly consists of shorefacestudied succession. units (Fig. 3). However, the occurrence of lingulid valves in such This taphofacies is composed by entire lingulid valves, parallel to the sandstone layers is extremely rare, and, when present, they are poorlybedding plane, and fragments of valves occurring in association with preserved and in small numbers (b5% in comparison with otherlower shoreface to offshore sedimentation, as indicated by fossil outcrops). In addition, lingulid occurrences conﬁned spatially toburrows (e.g. Zoophycos) and depositional sedimentary structures distinct patches (a patchy distribution means that the lingulids aresuch as micro-hummocky bedding. The majority of entire or fragmen- commonly restricted to particular sedimentary and biological faciested, disarticulated valves, occurs in ﬁne-grained host rock such as light and are not normally found outside them) (Ager, 1967) weregrey siltstones or dark grey siltstones and rare black shales. Commonly observed in siltstones by Nunes (1999). Nunes report representedsuch siltstones bear intercalations of centimetric lenses of very ﬁne- the ﬁrst record of lingulid patches in the studied Devonian succes-grained sandstone and in which lingulid fossils are not preserved. This sions. Nunes (1999), who worked in the outcrops along the PR-092taphofacies is not comparable to any of those proposed by Speyer and Highway between Jaguariaíva and Arapoti cities (Paraná State)Brett (1988) mainly because: (1) we do not notice the occurrence of (Fig. 1), identiﬁed lingulid patches in bioturbated siltstones contain-corrosion; (2) the degree of disarticulation and fragmentation in ing approximately 218 shells perpendicularly or obliquely preserved.our material is always high (it increases downslope); and (3) the Field work by Bosetti (2004) and Zabini (2007), and more recentreorientation/sorting is considered together in the Speyer and Brett surveys on several localities, failed to identify these patches. Considering(1988) model, but reorientation in our samples occurred mostly in the the amount of samples and outcrops surveyed in the present study, thedistal part of the transitional offshore, and sorting was not detected data reported by Nunes (1999) is considered unusual for this succession.among the lingulid valves. That is, lingulid patches are not a common feature in the Devonian successions analysed here. Also, the data provided by Nunes (1999)5.3. Did the Devonian lingulids live in the same substrate as extant support the hypothesis that these Devonian lingulids also lived in thespecimens? transitional offshore zone, namely, the clay siltstones. According to Emig (1984; 1997) Lingula spp. currently lives in 6. Summarysubstrates of ﬁne-grained sands, in infralittoral settings within depthsdown to 160 m. The Devonian succession studied here could The taphonomical signatures of the fossil lingulids are explainedpotentially preserve lingulids in sandstone layers in life position, in by reworking (most likely, mechanically via orbital wave movement),
C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56 55either above the fair weather wave-base or during rare storm events. Bosetti, E.P., 1989. Paleontologia dos Lingulida (Brachiopoda: Inarticulata) da Formação Ponta Grossa, Devoniano – Paraná, Brasil. MSc dissertation, Federal University ofSuch storms could have resulted in the distal deposition of shell Rio Grande do Sul, Porto Alegre, 119 pp.fragments below the average storm wave-base, mixing the fragments Bosetti, E.P., 2004. Tafonomia de alta resolução das fácies de offshore da sucessãowith intact valves recently removed from lingulid burrows. In addition, devoniana da região de Ponta Grossa – Paraná, Brasil. PhD thesis, Federal University of Rio Grande do Sul, Porto Alegre, 137 pp.concordant lingulid valves are associated with rare, articulated, lingulid Bosetti, E.P., Peyerl, D., Horodyski, R.S., Zabini, C., 2008. Formação Ponta Grossa: História,valves in life position. These lines of evidence suggest that the studied Fácies e Fósseis. I Simpósio de Pesquisa em Ensino e História de Ciências da Terra/ IIIlingulid associations represent autochthonous to parautochthonous Simpósio Nacional sobre Ensino de Geologia no Brasil. Unicamp, Campinas, Brasil, pp. 353–360.assemblages. Boucot, A.J., 1971. Malvinokaffric Devonian marine community distribution and Taphonomical data do not provide any conclusive evidence that implications for Gondwana. Academia Brasileira de Ciências 43, 23–49 Suppl.Palaeozoic lingulid valves had a different taphonomic response to Bretsky, P.W., 1969. Evolution of Paleozoic benthic marine invertebrate communities. Palaeogeography, Palaeoclimatology, Palaeoecology 6, 45–59.environmental factors than that would be expected for modern Campbell, J.D., 1987. Triassic records of the genus Lingula (Brachiopoda: Inarticulata) inlingulids. This suggests that the principles of actuopalaeontology may New Zealand. Journal of the Royal Society of New Zealand 17, 9–16.be applicable for comparing the intrinsic factors involved in shell Cherns, L., 1979. The environmental signiﬁcance of Lingula in the Ludlow series of thepreservation. However, more studies on the chemical content of the Welsh Borderland and Wales. Lethaia 12, 35–46. Emig, C.C., 1981a. Implications de données récentes sur les Lingules actuelles dans lesshell are still necessary to fully understand the role of intrinsic shell interpretátions paleoecologiques. Lethaia 14, 151–156.characteristics on the formation of these lingulid concentrations. Emig, C.C., 1981b. Observations sur lecology de Lingula reevei Davidson (Brachiopoda, The environmental conditions of the studied Palaeozoic sea were Inarticulata). Journal of Experimental Marine Biology and Ecology 52, 47–61. Emig, C.C., 1984. Importance du sédiment dans la distribution des Lingules. Lethaia 17,probably different than the ones that typify modern seas. Most likely, 115–123.limited water circulation and oxygen-stratiﬁcation represent the two Emig, C.C., 1986. Conditions de fossilisation du genre Lingula (Brachiopoda) etmain differences between Palaeozoic and post-Palaeozoic deposition- implications paléoécologiques. Palaeogeography, Palaeoclimatology, Palaeoecol- ogy 53, 245–253.al environments, in which lingulids occur. Emig, C.C. 1990. Examples of post-mortality alteration in Recent brachiopod shells and High resolution taphonomical analyses, integrated with an actualis- (paleo)ecological consequences. Marine Biology 104, 233–238.tic approach regarding lingulid habits and habitats, conclusively Emig, C.C., 1997. Ecology of the inarticulated brachiopods. In: Kaesler, R.L. (Ed.), Treatise on Invertebrate Paleontology. Part H — Brachiopoda. : The Geologicaldemonstrate that the preservation mode described for the Devonian Society of America and The University of Kansas. Colorado and Lawrence, pp.Paraná Basin represents a typical taphofacies. The lingulid valves 473–495.occurred in proximal settings ranging from the shoreface (lowermost Emig, C.C., 2003. Proof that Lingula is not a living fossil, and emended diagnosis of the Family Lingulidae. Cárnets de Geologie/Notebooks on Geology 1, 1–8.intertidal) to the transitional offshore (shallow subtidal) zones. In situ, Emig, C.C., 2008. On the history of the names Lingula, anatina, and on the confusion ofautochthonous assemblages formed preferentially in the shoreface the forms assigned them among the Brachiopoda. Cárnets de Geologie/Notebookssettings, whereas the transitional offshore deposits contained mostly on Geology 8, 1–13.paraautochthonous associations representing a mixture of complete, Emig, C.C., Gutiérrez-Marco, J.C., 1997. Signiﬁcation des niveaux à Lingulidés à la limite Supérieure du grès Armoricain (Ordovicien, Arenig, Sud-ouest de LEurope).disarticulated, and fragmented valves. The Paraná State Devonian Geobios 30, 481–495.sequences B, C, D, E, all include lingulid occurrences that can be Ferguson, L., 1963. The paleoecology of Lingula squamiformis Philips during a Scottishclassiﬁed into the proposed taphofacies. In contrast, lingulid patches, are Mississipian marine transgression. Journal of Paleontology 37, 669–681. Ghilardi, R.P., 2004. Tafonomia comparada e paleoecologia dos macroinvertebradosrarely observed in those sequences. (ênfase em trilobites), da Formação Ponta Grossa (Devoniano, Sub-bacia Apucarana), Estado do Paraná, Brasil. PhD thesis, University of São Paulo, São Paulo, 113 pp.Acknowledgements Goujet, D., Emig, C.C., 1985. Des Lingula fossiles, indicateurs de modiﬁcations de lenvironnement dans un gisement du Dévonien inferieur du Spitsberg. Comptes Rendus de lAcadémie des Sciences de Paris 301, 945–948. Financial support to this research was provided by Coordenação de Hammond, L.S., 1983. Experimental studies of salinity tolerance, burrowing behaviorAperfeiçoamento de Pessoal de Nível Superior (CAPES) and Conselho and pedicle regeneration in Lingula anatina. Journal of Paleontology 57, 1311–1316.Nacional de Desenvolvimento Cientíﬁco e Tecnológico (CNPq). M.H. Havlíček, V., Röhlich, P., 1984. Devonian and Carboniferous brachiopods from the Northern ﬂank of the Murzuq Basin (Libya). Sborník geologických věd Paleontologie 28, 117–177.acknowledges CNPq for a study grant (process 302666/04-4). E.P.B. Hiller, N., 1990. Benthic communities and sedimentary facies in the Lower Wittebergacknowledges CNPq for a study grant (process 480427/07-0). Dr. Michal Group (Devonian, South Africa). Annals of the South African Museum 99, 215–230.Kowalewski (Virginia Tech) read the manuscript and his comments Hiller, N., 1993. A modern analogue for the Lower Ordovician Obolus conglomerate of Estonia. Geological Magazine 130, 265–267.greatly improved the content and the language. We also thank Dr. Troy Holmer, L.E., Popov, L.E., Streng, M., Miller, J.F., 2005. Lower Ordovician (Tremadocian)Dexter and Dr. Juan C. Cisneros for help with the language. Two lingulate brachiopods from the House and Fillmore Formations, Ibex area, westernanonymous referees are recognized for worthy suggestions and great Utah, USA. Journal of Paleontology 79, 884–906. Hori, R.S., Campbell, H.J., 2004. Lingularia sp. (Brachiopoda) from Middle Triassicimprovement to the manuscript. bedded chert in Shikoku, Japan. Journal of the Geological Society of Japan 110, 758–764. Kidwell, S.M., 1991. The stratigraphy of shell concentrations. In: Allison, P.A., Briggs, D.E.References G. (Eds.), Taphonomy: Releasing the Data Locked in the Fossil Record. Plenum Press, NewYork, pp. 211–290.Aceñolaza, F., Emig, C.C., Gutiérrez-Marco, J.C., 2003. Lingulid shell beds from the Kidwell, S.M., Fürsich, F.T., Aigner, T., 1986. Conceptual framework for the analysis and Ordovician of Argentina, with notes on other peri-Gondwanan occurrences. In: classiﬁcation of fóssil concentrations. Palaios 1, 228–238. Albanesi, G.L., Beresi, M.S., Peralta, S.H. (Eds.), Ordovician from the Andes. INSUGEO, Kowalewski, M., 1996a. Taphonomy of a living fossil: the lingulide brachiopod Glottidia Serie Correlación Geológica, vol. 17. San Miguel de Tucumán, pp. 237–244. palmeri Dall from Baja California, Mexico. Palaios 11, 244–265.Ager, D.V., 1967. Brachiopod palaeoecology. Earth-Science Reviews 3, 157–179. Kowalewski, M., 1996b. Time-averaging, overcompleteness, and the geological record.Archbold, N.W., 1981. Lingula (Lingulidae, Brachiopoda) from the late Artinskian The Journal of Geology 104, 317–326. (Permian), Carnarvon Basin, Western Australia. Proceedings of the Royal Society of Kowalewski, M., Flessa, K.W., 1996. Improving with age: the fossil record of lingulide Victoria 92, 183–191. brachiopods and the nature of taphonomic megabiases. Geology 24, 977–980.Archbold, N.W., Cisterna, G.A., Sterren, A.F., 2005. Lingulida (Brachiopoda) from the Lange, F.W., Petri, S., 1967. The Devonian of the Paraná Basin. Boletim Paranaense de Early Permian of Argentina. Proceedings of the Royal Society of Victoria 117, Geociências 5–55 21 and 22. 307–317. Mergl, M., 2008. Lingulate brachiopods from the Acanthopyge Limestone (Eifelian) ofBergamaschi, S., 1999. Análise estratigráﬁca do Siluro-Devoniano (Formações Furnas e the Barrandian, Czech Republic. Bulletin of Geosciences 83, 281–298. Ponta Grossa) da sub-bacia de Apucarana, Bacia do Paraná, Brasil. PhD thesis, Mergl, M., Kordule, V., 2008. New Middle Cambrian lingulate brachiopods from the Skryje- University of São Paulo, São Paulo, 167 pp. Týřovice área (Central Bohemia, Czec Republic). Bulletin of Geosciences 8, 11–22.Bergamaschi, S., Pereira, E., 2001. Caracterização de sequências deposicionais de 3° Milani, E.J., França, A.B., Schneider, R.L., 1994. Bacia do Paraná. Boletim Geociências da ordem para o Siluro-Devoniano na sub-bacia de Apucarana, Bacia do Paraná, Brasil. Petrobrás 8, 69–82. Ciência-Técnica-Petróleo 20, 63–72. Milani, E.J., Faccini, U.F., Scherer, C.M.S., Araújo, L.M., Cupertino, J.A., 1998. SequencesBiernat, G., Emig, C.C., 1993. Anatomical distinctions of the Mesozoic lingulide and stratigraphy hierarchy of the Paraná Basin (Ordovician to Cretaceous), brachiopods. Acta Palaentologica Polonica 38, 1–20. southern Brazil. Boletim do Instituto de Geociências/USP 29, 125–173.
56 C. Zabini et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 292 (2010) 44–56Milani, E.J., Melo, J.H.G., Souza, P.A., Fernandes, L.A., França, A.B., 2007. Bacia do Paraná. stratigraphy? Some Paleozoic examples from the Paraná Basin, Brazil: XL Congresso Boletim de Geociências da Petrobrás 15 (2), 265–287. Brasileiro de Geologia, Belo Horizonte, p. 444.Nunes, J.R.S., 1999. Análise tafonômica dos braquiópodes inarticulados (Lingulida) do Simões, M.G., Mello, L.H.C., Rodrigues, S.C., Leme, J.M., Marques, A.C., 2000. Conulariid Membro Jaguariaíva, Formação Ponta Grossa (Devoniano) e suas implicações taphonomy as a tool in paleoenvironmental analysis. Revista Brasileira de Geociências paleoambientais. BSc honours dissertation, State University of São Paulo, Botucatu, 60 pp. 30, 757–762.Paine, R.T., 1963. Ecology of the brachiopod Glottidia pyramidata. Ecological Mono- Soares, S.P., 2007. Sistemática, tafonomia e paleoecologia de Trilobita, Phacopida (Homalo- graphs 33, 187–213. notidae, Calmonidae), Formação Ponta Grossa (Devoniano), Sub-bacia Apucarana,Parsons, K.M., Brett, C.E., Miller, K.B., 1988. Taphonomy and depositional dynamics of Estado do Paraná, Brasil. MSc dissertation, São Paulo University, São Paulo, 174 pp. Devonian shell-rich mudstones. Palaeogeography, Palaeoclimatology, Palaeoecol- Speyer, S.E., Brett, C.E., 1986. Trilobite taphonomy and Middle Devonian taphofacies. ogy 63, 109–139. Palaios 1, 312–327.Pashin, J.C., Ettensohn, F.R., 1992. Paleoecology and sedimentology of the dysaerobic Speyer, S.E., Brett, C.E., 1988. Taphofacies models for epeiric sea environments: Middle Bedford fauna (Late Devonian), Ohio and Kenthucky (USA). Palaeogeography, Paleozoic examples. Palaeogeography, Palaeoclimatology, Palaeoecology 63, 225–262. Palaeoclimatology, Palaeoecology 91, 21–34. Walker, R.G., Plint, A.G., 1992. Wave and storm dominated shallow marine systems. In:Petri, S., 1948. Contribuição ao estudo do Devoniano Paranaense. Departamento Walker, R.G., James, N.P. (Eds.), Facies Models – Response to sea level change: Nacional de Produção Mineral, Boletim 129. . 125 pp. Geological Association of Canada, pp. 219–238.Petri, S., Fúlfaro, V.J., 1983. Geologia do Brasil. EDUSP, São Paulo. Zabini, C., 2007. Lingulídeos da Sucessão Devoniana da Bacia do Paraná, região dosPickerill, R.K., Harland, T.L., Fillion, D., 1984. In situ lingulids from deep-water Campos Gerais, Brasil: revisão de conceitos biológicos-ecológicos e análise carbonates of the Middle Ordovician Table Head Group of Newfoundland and the tafonômica básica. MSc dissertation, Federal University of Rio Grande do Sul, Trenton Group of Quebec. Canadian Journal of Earth Sciences 21, 194–199. Porto Alegre, 130 pp.Popov, L.E., Ebbestad, J.O.R., Mambetov, A., Apayarov, F.Kh., 2007. A low diversity Zabini, C., Bosetti, E.P., Horodyski, R.S., Matsumura, W.M.K., 2007. Lingulídeos: revisão shallow water lingulid brachiopod–gastropod association from the Upper Ordovi- dos conceitos morfo-anatômicos, ﬁsioloógicos, reprodutivos, paleo-ecológicos e a cian of Kyrgyz Range. Acta Palaeontologica Polonica 52 (1), 27–40. importância do grupo no Devoniano da região dos Campos Gerais do Paraná, Brasil.Reading, H.G., Collinson, J.D., 1996. Clastic coasts. In: Reading, H.G. (Ed.), Sedimentary Terra Plural 1 (1), 123–141. Environment. Blackwell Science, pp. 154–231. Zalán, P.V., Wolff, S., Conceição, J.C., Marques, A., Astolﬁ, M.A., Vieira, I.S., Appi, V.T.,Rodrigues, S.C., Simões, M.G., Leme, J.M., 2003. Tafonomia comparada dos Conulatae Zanotto, O.A., 1990. Bacia do Paraná. In: Raja Gabaglia, G.P., Milani, E.J. (Eds.), (Cnidaria), Formação Ponta grossa (Devoniano), Bacia do Paraná, Estado do Paraná. Origem e Evolução das Bacias Sedimentares, pp. 135–168. Petrobras. Revista Brasileira de Geociências 33 (4), 381–390. Zhang, Z., Shu, D., Han, J., Liu, J., 2007. A gregarios lingulid brachiopod LongtancunellaSimões, M.G., Ghilardi, R.P., 2000. Protocolo tafonômico/paleoautoecológico como ferra- chengjiangensis from the Lower Cambrian, South China. Lethaia 40, 11–18. menta nas análises paleossinecológicas de invertebrados: exemplos de aplicação em Zhang, Z., Han, J., Emig, C.C., Shu, D., 2009. Epibiontes on the lingulate brachiopod concentrações fossilíferas do Paleozóico da Bacia do Paraná, Brasil. Pesquisas em Diandongia from the Early Cambrian Chengjiang Lagerstatte, South China. Geociências 27, 3–13. Proceedings. Biological Sciences/ Royal Society 277 (1679), 175–181.Simões, M.G., Kowalewski, M., Torello, F.F., Anelli, L.E., 1998a. Long term time- Zonneveld, J.-P., Beatty, T.W., Pemberton, S.G., 2007. Lingulide brachiopods and the averanging despite abrupt burial: Paleozoic obrution deposits from epeiric settings trace fossil Lingulichnus from the Triassic of Western Canada: implications for of Paraná Basin, Brazil. Geological Society of American Annual Meeting, Abstracts faunal recovery after the end-Permian mass extinction. Palaios 22, 74–97. with Programs, Toronto, p. 384.Simões, M.G., Torello, F.F., Kowalewski, M., Klein, C., Mello, L.H.C., Ghilardi, R.P., 1998b. Are the obrution deposits the most precise and the best resolved beds inevent