Journal of Paleontology, 86(1), 2012, p. 718 Copyright 2012, The Paleontological Society 0022-3360/12/0086-0007$03.

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AN EARLY CAMBRIAN SHALLOW-MARINE ICHNOFAUNA FROM THE PUNCOVISCANA FORMATION OF NORTHWEST ARGENTINA: THE INTERPLAY BETWEEN SOPHISTICATED FEEDING BEHAVIORS, MATGROUNDS AND SEA-LEVEL CHANGES
LUIS A. BUATOIS
AND

MARIA GABRIELA MANGANO

Department of Geological Sciences, University of Saskatchewan, 114 Science Place, Saskatoon, SK S7N 5E2, Canada, ,luis.buatois@usask.ca., ,gabriela.mangano@usask.ca.

ABSTRACTAn early Cambrian ichnofauna consisting of Helminthoidichnites tenuis, Helminthopsis tenuis, Multina isp., Oldhamia alata, and Pilichnus cf. dichotomus is documented from shallow-marine deposits ranging from the upper offshore to the offshore transition in the Puncoviscana Formation of northwest Argentina. Although the ichnogenus Oldhamia is more common in Cambrian deep-marine environments, this occurrence provides further evidence that it is also present in shallow-marine environments. The burrow network Multina (senior synonym of Olenichnus) is preserved at the base of tempestites, representing the activity of post-storm colonizers. A drowning surface separating offshore-transition deposits below from upper-offshore deposits above contains widespread evidence of trace fossils in direct association with matgrounds. The undermat miners Oldhamia alata and Pilichnus cf. P. dichotomus occur on this surface, revealing exploitation of organic matter in the biomat. Low sediment rate during drowning and paucity of bioturbation by sediment bulldozers may have promoted the establishment of the matground. In comparison with the simpler animal-matground interactions characteristic of the Ediacaran, the combination of Cambrian evolutionary innovations and the presence of microbial mats promoted more sophisticated interactions. Complex feeding trace fossils revealing that systematic undermat mining, as displayed by Oldamia alata and Pilichnus cf. dichotomus, is a product of the Cambrian explosion.

INTRODUCTION

provide significant information to understand the timing and nature of the Cambrian explosion. In addition to critically assessing ichnodiversity levels during the Cambrian, recent work has focused on understanding feeding strategies recorded by trace fossils, particularly exploring the association between biogenic structures and microbial mats (e.g., Seilacher, 1999; Buatois and Mangano, 2003a). The Ediacaranlower Cambrian Puncoviscana Formation of northwestern Argentina hosts one of the most diverse ichnofaunas for that critical interval (see review by Buatois and Mangano, 2004). Traditionally considered as recording sedimentation in deep-marine environments (Omarini and Baldis, 1984; Jezek, 1990; Acenolaza et al., 1999), this view has changed in recent years with the increasing realization that some Puncoviscana deposits accumulated in shallow-marine areas (Buatois and Mangano, 2003b, 2004). In this paper we document a new trace-fossil locality, referred to as El Mollar, in the Quebrada del Toro, Salta Province, northwest Argentina (Fig. 1.11.3). Shallow-marine deposits at this locality contain a very well-preserved ichnofauna associated with structures indicative of microbial mats, mostly preserved on a drowning surface (i.e., flooding surface). The succession exposed is somewhat atypical of the Puncoviscana Formation because it is comparatively less deformed, allowing measuring of a continuous section. In addition to the trace fossils on the drowning surface, burrow networks are preserved on the base of tempestites. The ichnotaxonomic status of burrow networks is still controversial, and therefore a discussion on network ichnotaxa is presented. The aims of this paper are to: 1) document the trace-fossil content of this new locality; 2) review the ichnotaxonomic status of Cambrian networks and related ichnotaxa (e.g., Multina Orowski, 1968 and Olenichnus Fedonkin, 1985); 3) discuss the role of sea-level changes in the formation of
RACE FOSSILS

an ichnofossil-bearing biomat surface; and 4) evaluate the interactions between sophisticated feeding behaviors and microbial mats.
STRATIGRAPHIC AND DEPOSITIONAL SETTING

The Puncoviscana Formation is the metasedimentary basement of northwest Argentina (Turner, 1960; Acenolaza and Toselli, 1981; Ramos, 2008). It consists of a thick, folded succession of wackes and mudstone, with subordinate presence of conglomerate, limestone and volcanic rocks affected by very low grade regional metamorphism, ranging from slates to schists (Do Campo and Nieto, 2003; Do Campo and Guevara, 2005). As noted in many studies (e.g., Mon and Hongn, 1988, 1991; Hongn, 1996; Moya, 1998; Becchio et al., 1999; Mangano and Buatois, 2004), rocks of different type, degrees of metamorphism and tectonic deformation have been included under the name Puncoviscana Formation, suggesting the possibility of further subdivision. Unfortunately, the intense deformation of the Puncoviscana Formation complicates establishing a sound stratigraphic subdivision and for proposing detailed correlations. Although the Puncoviscana Formation was considered originally as Precambrian (Turner, 1960, 1972), the discovery of the trace fossil Oldhamia provided uncontroversial evidence that this unit includes Cambrian strata also (Mirre and Acenolaza, 1972; Acenolaza and Durand, 1973). Recent geochronologic studies suggested that sedimentation in the Puncoviscana basin comprised the terminal Proterozoic and the early Cambrian (Ramos, 2000, 2008; Hongn et al., 2010). TIMS and SHRIMP U-Pb zircon geochronology data indicate that deposition may have started during the latest Ediacaran, but took place mainly during the Fortunian, coeval with 540535 Ma calc-alkaline arc volcanism, with probable slightly younger deposits towards the southwest of the basin (Escayola et al., 2011). This age is consistent with recent ichnologic analysis, which suggested an 7

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FIGURE 1Location and geological maps of the study area: 1, general map showing distribution of outcrops of the Puncoviscana Formation and overall extension of the Puncoviscana Basin (after Ramos, 2008); 2, general location map of Salta Province, Salta city, and the El Mollar; 3, geologic map of the Quebrada del Toro showing location of the El Mollar (modified from Durand and Acenolaza, 1990).

earliest Cambrian age for the trace fossil-bearing strata (Buatois and Mangano, 2004; Mangano and Buatois, 2004). In contrast to the proposal of Acenolaza (2003), there is no evidence of an Ediacaran age for any of the trace-fossil localities (Buatois and Mangano, 2005). The Puncoviscana Basin is represented by a north-south trending outcrop belt of more than 800 km length and 150 km wide, extending from southern Bolivia to the surroundings of the city of Tucuman, northwestern Argentina (Ramos, 2008) (Fig. 1.1). Historically, the Puncoviscana Formation has been interpreted as entirely deposited in deep-marine submarine fans (Omarini and Baldis, 1984; Jezek, 1990; Acenolaza et al., 1999). However, a more recent re-evaluation of its ichnologic content and sedimentary facies indicated that shallow-marine deposits are represented also (Buatois and Mangano, 2002, 2003b, 2004; van Staden and Zimmermann, 2003; Lopez de Azarevich et al., 2010). A more complex paleoenvironmental framework consisting of deep-marine turbiditic deposits along a western belt and shallow-marine environments affected by wave action along an eastern belt has been proposed (Buatois and Mangano, 2002, 2003a, 2003b, 2004). In particular, Buatois and Mangano (2004) described lower-offshore to middle/lower-shoreface facies, forming coarsening-upward parasequences and displaying evidence of oscillatory flows. Identical shallow-marine facies have been recently identified by Lopez de Azarevich et al. (2010). In addition, van Staden and Zimmermann (2003) recognized abundant glauconite layers interbedded with conglomerate in the Rio Corralito outcrops, suggesting shallow-marine deposition.

SEDIMENTARY FACIES

A bed-by-bed 21 m thick section (Fig. 2) was measured on one of the flanks of a fold near the El Mollar locality along the Old National Road 51 in the Quebrada del Toro, Salta Province, northwest Argentina (Fig. 1.3). Two sedimentary facies have been recognized in this section. Facies A consists of mudstone and parallel-laminated siltstone interbedded with discrete layers of erosionally based, laterally extensive to rarely lenticular, thin, very fine-grained sandstone with micro-hummocky cross-stratification (Fig. 3.1) and symmetrical to near-symmetrical ripples (Fig. 3.2). Hummocky cross-stratification occurs locally. Mudstone and siltstone intervals are 560 cm thick, and sandstone beds are 320 cm thick. Sandstone to mudstone ratios range from 1:3 to 1:5. Sandstone bases typically contain small tool marks, flute marks and load casts. Sandstone beds may show patchily distributed wrinkle marks. Multina isp. occurs at the base of thin sandstone layers, while Helminthoidichnites tenuis and Helminthopsis tenuis are relatively common at the top. The symmetrical to near-symmetrical ripples are interpreted as wave ripples and combined-flow ripples, respectively. The presence of these structures together with micro-hummocky and hummocky cross-stratification suggests that the sandstone layers are distal tempestites (Dott and Bourgeois, 1982; Myrow, 1992; Cheel and Leckie, 1993). The interbedded siltstone and mudstone intervals mostly record sediment fallout. Facies A is interpreted to record alternating background suspension fall-out and distal storm deposition above the

 BUATOIS AND MANGANOEARLY CAMBRIAN SHALLOW-MARINE TRACE FOSSILS

FIGURE 2Stratigraphic section of the Puncoviscana Formation at El Mollar locality, showing the development of two parasequences separated by a drowning surface (DS).

storm wave base, but well below the fair-weather wave base in an upper-offshore environment. This is the most abundant facies in the El Mollar section, forming up to 10 m thick intervals. Facies B consists of regularly interbedded, erosionally based, laterally extensive very fine-grained sandstone, parallel-laminated siltstone, and mudstone. Sandstone beds display hummocky cross-stratification, micro-hummocky cross-stratification, and symmetrical to near-symmetrical ripples. Mudstone and siltstone intervals are 113 cm thick, and sandstone beds are 517 cm thick. Sandstone to mudstone ratios are typically 1:1. Sandstone bases may contain flute marks. Multina isp. locally occurs at the base of thin sandstone layers. Abundant, though patchily distributed, wrinkle marks and elephant-skin textures occur on the planar tops of sandstone and siltstone beds. Oldhamia alata, Pilichnus cf. P. dichotomus, Helminthoidichnites tenuis, and Helminthopsis tenuis occur on one of these surfaces, marking the top of facies B package.

In thin section across this surface, microsequences of finingupward siltstone capped by microbial mat are observed (Fig. 4.1). Locally, the mat is deformed (Fig. 4.1). Under magnification, the typical texture of endobenthic matgrounds is present, consisting of filaments intertwined in angles ranging from 045u, is present (Fig. 4.2) (see Noffke, 2010). Framboidal pyrite is common. Floating quartz grains are enveloped within the biomat (Fig. 4.3). Burrows are observed deforming the matground above and below; burrow fill is rich in pyrite (Fig. 4.4). As in the case of facies A, the symmetrical to nearsymmetrical ripples represent wave ripples and combined-flow ripples, respectively. The presence of these structures, as well as of hummocky and micro-hummocky cross-stratification, indicates storm deposition, while the interbedded siltstone and mudstone mostly record sediment fall-out (Cheel and Leckie, 1993; Dumas and Arnott, 2006). The increased sandstonemudstone ratios indicates a slightly more proximal position than facies A. Facies B is interpreted to record alternating

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FIGURE 3Upper-offshore tempestites in the Mollar section showing evidence of oscillatory flows: 1, sharp-based sandstone bed with microhummocky cross-stratification; 2, sharp-based sandstone bed with symmetrical to near-symmetrical ripples. Lens cap is 5.5 cm wide.

background suspension fall-out and distal storm deposition right below the fair-weather wave base in an offshoretransition environment. This facies forms a single discrete two-meter thick interval. The facies described are stacked forming two parasequences separated by a drowning surface. This low-energy surface clearly indicates a rapid upward deepening without any associated erosion. The lower parasequence consists of a thick upperoffshore interval capped by a thin offshore-transition interval. The upper parasequence consists entirely of upper-offshore deposits.
SYSTEMATIC PALEONTOLOGY

the host rock. Overlapping among different individuals is common. Diameter is 0.82.1 mm and may slightly vary along the course of individual trails. Maximum observed length is 17.0 mm. Preserved in full relief on sandstone tops. Discussion.Helminthoidichnites tenuis is one of the most common components of Ediacaran and lower Cambrian ichnofaunas (e.g., Hofmann et al., 1994; MacNaughton et al., 2000; Jensen et al., 2006). This ichnotaxon is interpreted as a grazing trace (pascichnion) most likely produced by vermiform animals (Buatois et al., 1998). Ichnogenus HELMINTHOPSIS Heer, 1877 _kiewicz 1968 HELMINTHOPSIS TENUIS Ksiaz Specimens.Approximately ten specimens studied in the field. Description.Simple, unbranched, meandering trails, oriented parallel to the bedding plane. Fill is identical to the host rock. Diameter is 0.71.1 mm. Maximum observed length is 14.0 mm. Preserved in full relief on sandstone tops. Discussion.Helminthopsis tenuis is particularly abundant in EdiacaranEarly Cambrian ichnofaunas (e.g., Buatois and Mangano, 2003a; Jensen et al., 2006). It is interpreted as a grazing trace (pascichnion) probably produced by vermiform animals (Ksiaz _kiewicz, 1977; Buatois et al., 1998).

Ichnotaxa are listed alphabetically. Specimens are housed at the Paleontological Museum Egidio Feruglio, Trelew, Argentina (MPEF-IC). Ichnogenus HELMINTHOIDICHNITES Fitch, 1850 HELMINTHOIDICHNITES TENUIS Fitch, 1850 Figure 5 Specimens.Approximately 30 specimens studied in the field and seven slabs with twelve specimens collected (MPEFIC 424, 426, 427, 428, 430, 432, 433). Description.Simple, unbranched, straight to gently curved trails, oriented parallel to the bedding plane. Fill is identical to

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FIGURE 4Microbial-mat textures in thin-section across the drowning surface: 1, microsequences of fining-upward siltstone capped by microbial mat; note mat deformation (arrows) and enrichment in organic matter; scale bar51 mm; 2, texture of endobenthic matground, consisting of filaments intertwined in angles ranging from 045u; scale bar51 mm; 3, floating quartz grains (arrows) enveloped within the mat; scale bar50.5 mm; 4, burrow in cross section, displaying spreite and visibly deforming the matground above and below; note abundant pyrite within the burrow; scale bar51 mm.

Ichnogenus MULTINA Orowski, 1968 MULTINA isp. Figure 6.16.3 Specimens.Approximately 50 specimens studied in the field and one slab with one specimen collected (MPEF-IC 419). Description.Irregular overlapping networks having straight, meandering to winding strings. Network size is 28.1190.4 mm and string diameter is 2.04.0 mm. Preserved in full relief on sandstone bases. Discussion.The ichnogenus Multina was introduced by Orowski (1968) based on specimens from upper Cambrian shallow-marine deposits of the Holy Cross Mountains in Poland. These specimens, included in the type ichnospecies M. magna, were subsequently redescribed by Orowski and Zylinska (1996). This ichnotaxon consists of approximately 1 mm wide curved epichnial grooves that display common overlap and bifurcate forming 25 cm wide irregular polygons. Orowski and Zylinska (1996) noted indistinct transverse furrows in some of the specimens, suggestive of backfill and peristaltic movements. Another occurrence of M. magna was documented from Lower Ordovician turbidites of northwest Argentina (Buatois et al., 2009). A second ichnospecies, M. minima, was introduced by Uchman (2001) based on specimens from Eocene deep-marine deposits of Spain. Multina minima is

characterized by overlapping strings, containing abundant swellings and forming highly irregular and very small (less than 5 mm wide) networks. It shares with M. magna the presence of common overlap. The type specimen of M. minima and a second occurrence in Lower Cretaceous relatively deepwater deposits of Bulgaria (Uchman and Tchoumatchenco, 2003) are preserved as positive hyporeliefs. However, recently described specimens from the Eocene of Spain are preserved as full reliefs on the base of turbidites (Rodrguez-Tovar et al., 2010). Specimens from the Puncoviscana Formation have strings that are morphologically highly variable, being locally more meandering than those of M. magna, which are typically curved. In contrast, networks are more irregular than in M. magna. Also, Multina from the Puncoviscana Formation lacks the abundant swellings that typify M. minima, and is remarkably larger. Partially preserved networks consisting of isolated strings may be confused with grazing traces, such as Helminthopsis or Helminthorhaphe Seilacher, 1977 (fig. 5-3). As noted by Orowski and Zylinska (1996), Multina display morphologic similarities with some graphoglyptid ichnotaxa, most notably Paleodictyon Meneghini, 1850, Protopaleodictyon Ksiaz _kiewicz, 1958, and Megagrapton Ksiaz _kiewicz, 1968. However, although the morphology of the networks may be superficially similar, these three ichnotaxa are open galleries that are passively filled (i.e., they are agrichnial structures of farmers and trappers rather than fodinichnial

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FIGURE 5Helminthoidichnites tenuis preserved as full relief at the top of a sandstone tempestite; field photo; scale bar52 cm.

structures of deposit feeders). According to Uchman (1998), many lower Paleozoic occurrences of Megagrapton may in fact represent Multina, noting that these specimens display common overcrossing of the strings, which is not common, if present at all, in graphoglyptids. The same may be true of many lower Paleozoic recordings of Protopaleodictyon. The ichnogenus Olenichnus, with O. irregularis as its type ichnospecies, was proposed by Fedonkin (1985) based on earliest Cambrian specimens from shallow-marine deposits. Olenichnus comprises networks consisting of sinuous strings forming irregular networks preserved as negative hyporeliefs. The type specimen and additional material from the lower Cambrian of the Mickwitzia Sandstone in Sweden, have been revised by Jensen (1997), who noted the presence of knobs indicative of vertical components. Olenichnus has been regarded as a junior synonym of Multina by Uchman and Alvaro (2000). There are slight differences in the morphology of the networks that warrant keeping Olenichnus irregularis as a valid ichnospecies of Multina, M. irregularis. This ichnospecies is characterized by irregular networks of very variable size that do not form polygons as in M. magna. The absence of swellings distinguishes M. irregularis from M. minima. Uchman (1998) noted similarities between Multina and Pseudopaleodictyon Pfeiffer (1968), which was proposed based on Palaeophycus hartungi Geinitz (1867), becoming Pseudopaleodictyon hartungi. Based on the original description and illustration, the true nature of Pseudopaleodictyon is hard to evaluate. Although the exact date of publication of Pseudopaleodictyon is unclear (varying between 1966, 1968 and 1969, depending on source), the use of Pseudopaleodictyon is not recommended, and Multina is preferred instead. Two other network ichnogenera have been described from continental deposits. These are Vagorichnus described by Buatois et al. (1995) from Jurassic lacustrine turbidites of

FIGURE 6Multina isp. preserved as full relief at the base of sandstone tempestites: 1, general view of networks consisting of meandering and winding strings, field photo, note branching (arrow); 2, close-up of networks, field photo; 3, individual string resembling a meandering grazing trail rather than a burrow network, MPEF-IC 419, scale bar51 cm long. Lens cap is 5.5 cm wide.

China and Labyrintichnus described by Uchman and Alvaro (2000) from Miocene marginal-lacustrine deposits of Spain. The former was first regarded as a junior synonym of Multina by Uchman and Alvaro (2000) and Schlirf et al. (2001), but

 BUATOIS AND MANGANOEARLY CAMBRIAN SHALLOW-MARINE TRACE FOSSILS subsequently retained as a valid ichnotaxon by Uchman et al. (2007). As in Multina, Vagorichnus is a feeding structure involving active fill. However, in contrast to Multina, Vagorichnus is characterized by a wider morphologic variability, a complex branching pattern (including also secondary successive branching), and abundant aligned knobs that indicate vertical meandering movements (Buatois et al., 1995). Labyrintichnus is morphologically very similar to Multina but it is passively filled (Uchman and Alvaro). Both Vagorichnus and Labyrintichnus are regarded here as valid ichnotaxa. Ichnogenus OLDHAMIA Forbes, 1849 OLDHAMIA ALATA Seilacher, Buatois and Mangano, 2005 Figure 7.17.6 Specimens.Thirteen slabs with forty five specimens collected (MPEF-IC 422 to 430 and 432 to 435). Description.Horizontal complex burrow systems consisting of closely spaced tunnels (i.e., probings) that form up to eight wing-like units resembling small Lophoctenium structures. Individual tunnels are 0.11.6 mm wide. Wing-like units are 4.420.1 mm wide. As noted by Seilacher et al. (2005), probings proceed to the convex side commonly starting at the center and protruding outward, but a centripetal pattern has been detected in some lobes. Although opposite, 180u symmetrical wings, 29.532.5 mm wide, locally occur (Fig. 7.3), more complex asymmetrical patterns defined by successive wings alternating along an imaginary axis are more common. These cumulative structures are 17.335.0 mm. Successive wings are adjacent to each other, and do not overlap (Fig. 7.4). Abrupt changes in the length of individual tunnels reveal lobation within wings, having two to four, 3.58.5 mm wide lobes in each wing. Discussion.This ichnospecies was originally referred to as a new ichnospecies of Oldhamia by Buatois and Mangano (2004), and subsequently formally proposed by Seilacher et al. (2005), who described and interpreted this ichnotaxon in detail. Poorly preserved and isolated tunnels may be confused with grazing trails (e.g., Helminthoidichnites). In contrast to other ichnospecies of Oldhamia, probings in O. alata not only bend back on previous ones, but follow them so closely resembling the spreite of a tiny Lophoctenium Richter, 1850 (Seilacher et al., 2005). Oldhamia alata is reminiscent of the asymmetrical morphotypes of O. curvata (Fig. 7.5) but probes in O. alata stay in closer contact with previous ones and the wing-like units, in cases displaying lobation, are far more complex and commonly alternate (Seilacher et al., 2005). Oldhamia is a feeding trace (fodinichnion) produced by vermiform animals (Seilacher, 1999). Oldhamia is essentially a Cambrian ichnogenus, being particularly abundant during the early to middle Cambrian (Lindholm and Casey, 1990; Herbosch and Verniers, 2011). Although it is more common in deep-marine environments (e.g., Hofmann et al., 1994), it also occurs in shallow-marine deposits (e.g., Goldring and Jensen, 1996) (see Paleoenvironmental significance of Oldhamia). Ichnogenus PILICHNUS Uchman, 1999 PILICHNUS cf. P. DICHOTOMUS Uchman, 1999 Figure 8.1, 8.2 Specimens.One slab with three specimens collected (MPEFIC 426). Description.Tunnel systems consisting of horizontal, straight to locally winding strings commonly displaying dichotomous branches. T-shaped branchings are very rare. Tunnel fill is the same than the host rock. One of the specimens locally shows sinusoidal strings (Fig. 8.2). Strings

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are 0.10.2 mm wide. Preserved in full relief on sandstone tops. Discussion.Pilichnus was proposed based on specimens from Cretaceous turbidites in Germany (Uchman, 1999). The stratigraphic and paleoenvironmental range of Pilichnus have been recently expanded to include Cambrian (e.g., Zhang et al., 2007; Mangano, 2011) and Ordovician shallow-marine deposits (e.g., Mikulas, 2003), and Carboniferous turbidites (e.g., Mikulas et al., 2004). It is most likely a feeding trace (fodinichnion). Structures similar to Pilichnus are produced in modern shallow-marine environments by the polychaetes Heteromastus filiformis and Capitella cf. aciculata (Hertweck et al., 2007). The main difference between the specimens studied and P. dichotomus is the absence of a pyritized fill, which is characteristic of the latter ichnospecies (see Uchman, 1999) and the local presence of sinusoidal strings in the Puncoviscana material. Apparently branching sinusoidal specimens illustrated by Seilacher et al. (2005, their fig. 8B) are either overlapping Cochlichnus Hitchcock, 1858 or fragmentary preserved Pilichnus cf. P. dichotomus.
DISCUSSION

Paleoenvironmental significance of Oldhamia.The ichnogenus Oldhamia is a common component of Cambrian deepmarine environments worldwide (see Herbosch and Verniers, 2011 and references therein). As a result, it has been assumed in some studies that Oldhamia is an indicator of deep-marine environments (e.g., Tacker et al., 2010). However, this ichnogenus has also been recorded in shallow-marine facies (Kowalski, 1987; Goldring and Jensen, 1996; Buatois and Mangano, 2004). While Oldhamia is invariably the dominant ichnotaxon in deep-marine deposits where it occurs in relatively low-diversity assemblages, in shallow-marine settings it is typically an accessory element in more diverse assemblages (Goldring and Jensen, 1996; Buatois and Mangano, 2003b, 2004; Seilacher et al., 2005). At ichnospecific level, Oldhamia alata and O. geniculata are only known from shallow-marine deposits whereas O. flabellata and O. curvata have only been recorded in deep-marine environments; O. antiqua and O. radiata have been recorded in both shallow- and deep-marine settings (Seilacher et al., 2005). Microbial mats and sea-level changes.In recent years there has been increased interest on the interplay between sequence stratigraphy and matground development. A number of studies documented that in Precambrian rocks, microbial mat textures are preferentially preserved in the lower portion of highstand systems tracts (Banerjee and Jeevankumar, 2005; Sarkar et al., 2005; Catuneanu, 2007). It has been argued that Precambrian sequences are characterized by very thin transgressive systems tracts and well-developed highstand systems tracts; even in some cases, transgressive systems tracts are absent and highstand systems tracts are stacked (Sarkar et al., 2005). Within this framework, the absence of transgressive packages is related to the presence of gently dipping shelves, which facilitated rapid transgressions in combination with a generally low sediment supply. Sarkar et al. (2005) also proposed that binding of clastic particles resulting from the prolific growth of microbial mats reduced the effects of wave and current reworking, allowing sediment aggradation in spite of the low sediment supply. On the other hand, Noffke (2010) suggested that microbially induced sedimentary structures are particularly widespread during transgressive phases. She based her view on actualistic grounds, namely the widespread development of coastal areas during the Holocene transgression, leading to extensive

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FIGURE 7Oldhamia alata preserved as full relief at the top of a sandstone tempestite: 1, general view showing overall high density of specimens covering the bedding plane; corrugations on the left represent evidence of microbial mats; 2, close-up of specimens displaying typical branching pattern; note associated corrugations on the bedding surface; MPEF-IC 423; 3, detailed view of specimens showing symmetrical arrangement of wing-like lobes; upper specimen is the holotype; MPEF-IC 424; 4, close-up of specimens showing the non-overlapping nature of wing-like lobes; isolated tunnels, such as those on the right, may be confused with grazing trails; MPEF-IC 425; 5, detailed view of specimens that are somewhat reminiscent of O. curvata; nonoverlapping nature of the lobes, however, indicates affiliation with O. alata; MPEF-IC 430; 6, close-up view of specimens in close association with microbial mat textures. MPEF-IC 431. Scale bars51 cm.

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FIGURE 8Pilichnus cf. P. dichotomus preserved as full relief at the top of a sandstone tempestite; corrugations represent evidence of microbial mats; MPEF-IC 426: 1, general view showing overall branching pattern; scale bar51 cm; 2, various branching systems; note local development of sinusoidal morphology of elements in the system on the lower right; scale bar51 cm.

microbial mat ecosystems flourishing in tidal-flat, lagoon and shelf settings. Faults affecting the succession preclude placing the El Mollar section within a broad sequence-stratigraphic framework. Despite this, the association of an ichnofossil-bearing matground and the drowning surface is clear. In El Mollar section, the best-preserved trace-fossil suite is in direct connection with wrinkle marks and elephant-skin textures and it occurs at the siltstone mantling the planar top of a sandstone tempestite. This layer is capped by a low-energy drowning surface, and the top of the tempestite therefore represents an omission surface. Buatois and Mangano (2004) noted that low sediment rate during drowning and paucity of bioturbation by sediment bulldozers may have promoted the establishment of the matground. Interestingly, the ichnofauna of this bed resembles that of the deep-marine deposits, particularly with respect to the dominance of Oldhamia. Tiering structure.The tiering structure of the 5 cm thick offshore-transition bed right below the drowning surface is very simple (Buatois and Mangano, 2004). Three ichnoguilds have been recognized, from shallow- to deep-tier, Helminthopsis, Oldhamia and Multina. The Helminthopsis ichnoguild consists of Helminthopsis tenuis and Helminthoidichnites tenuis. This ichnoguild represents the activity of transitory, surface to near-surface, mat-grazer structures produced by

vagile vermiform animals. The Helminthopsis ichnoguild is ubiquitous in both Ediacaran and early Cambrian shallowand deep-marine deposits, revealing one of the simplest feeding strategies associated with biomats (Buatois and Mangano, 2003a). The Oldhamia ichnoguild consists of Oldhamia alata and Pilichnus cf. P. dichotomus. This ichnoguild includes semi-permanent, very shallow-tier, undermat-miner structures produced by stationary vermiform organisms. The Oldhamia ichnoguild is particularly common in lower to middle Cambrian deep-marine deposits but it has been recorded also in shallow-marine deposits, albeit rarely (see Paleoenvironmental significance of Oldhamia). The presence of Oldhamia in Ediacaran rocks has been questioned (Tacker et al., 2010) and accordingly this ichnoguild is most likely a product of the Cambrian explosion. The Multina ichnoguild consists of semi-permanent, shallowto middle-tier, deposit feeder structures produced by vagile vermiform organisms. It is represented by Multina isp., which is preserved at the base of the tempestite. Burrow networks invariably cross-cut tool marks, indicating that Multina is a post-event burrow emplaced after storm deposition. The producers burrowed into the tempestite and moved along the sand-mud interface. This ichnoguild serves as a proxy to evaluate burrowing depth in offshore settings by the early Cambrian, pointing to incipient colonization of the infaunal ecospace. The tracemakers moved along the lithologic interface, therefore producing negligible bioturbation and little or no disturbance. As in the case of the Oldhamia ichnoguild, the Multina ichnoguild seems to have been a product of the Cambrian explosion with no equivalents in the Ediacaran. Possible networks compared with Olenichnus have been recorded in the Huns Member of the Urusis Formation in Namibia by Jensen and Runnegar (2005). However, these are very shallow-tier structures. The Multina ichnoguild occurs in Tremadocian deep-marine deposits of the Puna area in northwest Argentina, suggesting a possible onshore-offshore pattern for this ichnotaxon (Buatois et al., 2009). The ichnofauna of the offshore-transition beds unrelated to the drowning surface (i.e., those offshore-transition layers below the uppermost bed) is very simple, and consists only of Multina isp. preserved at the base of sandstone tempestites, ranging from 5 to 8 cm thick. The tiering structure of the upper-offshore deposits is very similar to that of the offshoretransition bed below the drowning surface. However, the ichnoguild represented by undermat miners (Oldhamia alata and Pilichnus cf. P. dichotomus) is not present, and only two ichnoguilds have been recorded: the Helminthopsis and Multina ichnoguild. This provides further evidence that the conditions associated with the drowning event were instrumental in promoting development and preservation of matgrounds and associated trace fossils. The onset of sophisticated feeding strategies and the persistence of matgrounds.Interactions between organisms and matgrounds were widespread during Ediacaran times, but remain relatively simple due to the absence of organisms capable of developing complex feeding structures (Buatois and Mangano, in press). These were dominated by mat grazer structures of the Helminthopsis ichnoguild and mat scratcher structures of the Radulichnus ichnoguild, the latter in direct association with the mollusk-like Kimberella (Fedonkin, 2003; Gehling et al., 2005; Seilacher et al., 2005; Fedonkin et al., 2007; Seilacher and Hagadorn, 2010). Interactions are also evidenced by the presence of serially repeated resting trace fossils of Dickinsonia and the related genus Yorgia preserved

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JOURNAL OF PALEONTOLOGY, V. 86, NO. 1, 2012 Feruglio) curated the specimens and assisted us with photography.
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on top of microbial mats (Ivantsov and Malakhovskaya, 2002; Fedonkin, 2003; Gehling et al., 2005; Sperling and Vinther, 2010). The combination of persistent matgrounds, albeit more patchily distributed, and the evolutionary innovations of the Cambrian explosion allowed for more complex interactions to develop (Buatois and Mangano, in press). The sophisticated behavioral programs exhibited by Oldhamia alata and, to a lesser extent, Pilichnus cf. P. dichotomus in the Puncoviscana Formation clearly illustrate an increase in the complexity of organism-matground interactions with respect to those displayed by their Ediacaran counterparts. The closely guided probings in O. alata, leaving no unexplored sediment between successive elements within a lobe, evince an improved feeding program even if compared to other Oldhamia ichnospecies more common in deep-sea deposits (Seilacher et al., 2005). Pilichnus cf. P. dichotomus, with its systematic dichotomous branching pattern, reflects efficient exploration of the organic matter preserved below the microbial mat. In fact, it may be argued that systematic undermat mining is an evolutionary innovation which resulted from the Cambrian explosion. The best candidates for Ediacaran undermat miners are the structures illustrated in the Vingerbreek Member of the Nudaus Formation of Namibia by Bouougri and Porada (2007), which await further description.
CONCLUSIONS

An early Cambrian shallow-marine (upper offshore to offshore transition) ichnofauna is documented from the Puncoviscana Formation of northwest Argentina. The ichnofauna consists of Helminthoidichnites tenuis, Helminthopsis tenuis, Multina isp., Oldhamia alata and Pilichnus cf. P. dichotomus. This occurrence provides further evidence that the ichnogenus Oldhamia, although much more common in Cambrian deep-marine environments, is also present in shallow-marine environments. A drowning surface separating offshore-transition below from upper-offshore deposits above contains widespread evidence of trace fossils in direct association with matgrounds. The undermat miners Oldhamia alata and Pilichnus cf. P. dichotomus are particularly well preserved on this surface. Low sediment rate during drowning and paucity of bioturbation by sediment bulldozers may have promoted the establishment of the matground. While the trace-fossil record of animal-matground interactions during the Ediacaran is relatively simple, the combination of Cambrian evolutionary innovations and the presence of microbial mats were conducive to more complex interactions. Systematic undermat mining, as displayed by Oldamia alata and Pilichnus cf. P. dichotomus, is a product of the Cambrian explosion.
ACKNOWLEDGMENTS

Financial support for this study was provided by the Antorchas Foundation, and Natural Sciences and Engineering Research Council (NSERC) Discovery Grants 311727-05/08 and 311726-05/08 awarded to Mangano and Buatois, respec tively. I. Sabino assisted during the field work. N. Carmona provided detailed comments on the microbial mat structures. M. Bertling, D. Seilacher, A. Uchman and A. Zylinska provided feedback on various ichnologic aspects. B. Novakovski did the thin-sections. D. Dawson helped us to track the elusive publication date of the Pseudopaleodictyon paper. Reviewers S. Jensen and D. McIlroy, and Editor B. Pratt provided very useful comments. E. Ruigomez (Museo Egidio

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ACCEPTED 18 AUGUST 2011