New Insights on the Red Alga Archaeolithophyllum and Its Preservation from the Pennsylvanian of the Cantabrian Zone (NW

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New insights on the red alga Archaeolithophyllum and its preservation from the Pennsylvanian of the Cantabrian Zone (NW...

Article in Facies · October 2013 DOI: 10.1007/s10347-012-0347-8

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ORIGINAL ARTICLE

New insights on the red alga Archaeolithophyllum and its preservation from the Pennsylvanian of the Cantabrian Zone (NW Spain)

D. Corrochano • D. Vachard • I. Armenteros

Received: 27 June 2012 / Accepted: 7 November 2012 / Published online: 29 November 2012 Ó Springer-Verlag Berlin Heidelberg 2012

Abstract The new species Archaeolithophyllum asym- formed micrite-rich bioherms with abundant shelter metricum nov. sp., from the Bachende Formation (Penn- porosities, which are filled up with radiaxial-fibrous calcite sylvanian, Cantabrian Zone, NW Spain), is described (originally high-Mg calcite) and subsequent blocky spar. herein using cathodoluminescence microscopy. Under They constructed a rigid framework that was basically a plane-polarized light, A. asymmetricum occurs as elongate combination of the foliaceous growth form, crust fusion and arcuate sheets preserved as calcitic mosaics of tiny and division, and synsedimentary marine cementation. anhedral to subhedral crystals. Cathodoluminescence has Paleontological and sedimentological evidence suggests revealed that skeletal walls are composed of dull-bright that A. asymmetricum thrived in an outer platform envi- (locally bright) luminescent calcite that contrasts sharply ronment with relative quiet conditions. The exceptional with the nonluminescent cements filling the intraskeletal preservation of these algae was favored by a rapid pores. Skeletal walls are currently composed of low-Mg cementation of the intraskeletal pores under oxidizing calcite (0.5–2 mol % MgCO3) with low Sr content (aver- conditions in a marine phreatic environment, protecting age 415 ppm). A. asymmetricum shows a strong asymmetry skeletons from early dissolution and recrystallization. of the thallus organization. The internal tissue is well dif- Although the resulting neomorphic microsparite fabric ferentiated into a thick medullar hypothallus and a thin suggests an aragonite precursor, the morphological simi- upper cortical perithallus, the latter being composed of larities (especially reproductive organs) between Archae- nearly rectangular cells arranged in rows perpendicular to olithophyllum and Recent calcitic corallinaceans, and the the external surface. Cell fusions commonly occur in the similar trace element composition of the algal thalli and the perithallial tissue whereas conceptacles exhibit a highly surrounding high-Mg radiaxial-fibrous cements, suggest arched geometry lacking any preserved aperture. A. asym- that originally, Archaeolithophyllum was probably com- metricum accumulations display a growth pattern similar to posed of high-Mg calcite. Based on the morphologic fea- that reported from Late Paleozoic ‘‘phylloid algae’’, and tures, framework strategies (crust fusion and division) and also resemble Miocene frameworks of the corallinacean growth modes, it is suggested that Archaeolithophyllum Mesophyllum. These accumulations of A. asymmetricum might be phylogenetically related to the modern coralline algae.

& D. Corrochano ( ) I. Armenteros Keywords ‘‘Phylloid algae’’ Cathodoluminescence Facultad de Ciencias, Departamento de Geologı´a, Universidad de Salamanca, Plaza de los Caı´dos s/n, 37008 Salamanca, Spain Archaeolithophyllum Pennsylvanian Cantabrian Zone e-mail: [email protected] Spain I. Armenteros e-mail: [email protected] Introduction D. Vachard Universite´ Lille 1, UMR 8217 du CNRS, Ge´osyste`mes, 59655 Villeneuve d’Ascq Cedex, France The ‘‘phylloid alga’’ Archaeolithophyllum appeared in the e-mail: [email protected] Late Mississippian (for a review see Wray 1977a, b; 123 950 Facies (2013) 59:949–967

Vachard and Aretz 2004;Co´zar et al. 2005; Somerville Fig. 1 a Geological map of the Cantabrian Zone showing the c 2008), and is very abundantly represented in Late Penn- location of the Salamoń Gold District, modified from Julivert (1971) and Pe´rez-Estauń et al. (1988). b Stratigraphic correlation of the cores sylvanian and Early Permian shallow-water carbonate SS-14, SS-16, and SS-58 in the Salamoń Gold District, showing the shelves and ramps worldwide. They played an important sample horizon of Archaeolithophyllum asymmetricum sp. nov.; Mr role in mound and reef build-up development, commonly marl, M mudstone, W wackestone, P packstone, G grainstone, forming accumulations with high porosity and gas and oil R rudstone/floatstone, B boundstone, Md mudrock, f, m, c, Vc fine, medium, coarse and very coarse grained sandstone, respectively, Cg reservoir quality (Wray 1964). conglomerate Pray and Wray (1963) defined the term ‘‘phylloid’’ to describe various types of membranous leaf-like calcareous algae, irrespective of taxonomic identity or growth habit, Such cellular structure can hardly be observed under planar found in Late Paleozoic rocks of the Paradox Basin, USA. polarized light microscopy and, thus, most of taxonomic The term ‘‘phylloid’’ is predominantly applied to strongly affinities of neomorphosed ‘‘phylloid algae’’ are uncertain. recrystallized algae, which display a wide spectrum of Based on cathodoluminescence microscopy, this paper growth forms, although cyathiform (cup-shaped) and leaf- documents very well preserved specimens of a new species like (undulating plates) dominate (Enpu et al. 2007a). As of Archaeolithophyllum (A. asymmetricum sp. nov.) and the preservation of their internal cellular structure is illustrates the cellular structure and thallus organization of commonly poor, it is often virtually impossible to distin- this alga. The phylogenetic position of Archaeolithophyl- guish one genus from another (Wray 1977a), and this lum, along with the paleoecology, preservation, and dia- artificial group of algae has been usually used as a ‘‘waste- genesis, is also discussed. paper basket’’ (Schlagintweit 2010), and subsequently considered as unnecessary (Granier 2012). ‘‘Phylloid algae’’ include both Rhodophyta and Chloro- Materials and methods phyta (e.g., Konishi and Wray 1961; Wray 1977a; James et al. 1988; Enpu et al. 2007a, b). The most typical ‘‘phyl- Specimens of Archaeolithophyllum asymmetricum were loid algae’’, or Anchicodiaceae (Shuysky in Chuvashov collected from three cores (SS-14, SS-16, and SS-58) from et al. 1987; Mamet 1991) are represented by Ivanovia the Bachende Formation (late Kashirian/early Myachko- Khvorova 1946, Eugonophyllum Konishi and Wray 1961, vian, Moscovian, Pennsylvanian) in the Salamoń Gold Anchicodium Johnson 1946, and Neoanchicodium Endo in District (Lois-Ciguera sector, Central Asturian Coalfield, Endo and Kanuma 1954 (as well as many junior synonyms Cantabrian Zone, NW Spain; Fig. 1a, b); these cores are of these genera, such as Kansaphyllum Baars 1992; Calci- housed in the core-repository of Castilla y Leoń, Salam- petra Torres et al. 1992; and Iranicodium Senowbari-Dar- anca, Spain. SS-14 is 295 m long, SS-16 is 207 m long, yan and Rashidi 2010). The status of Anchicodium is and SS-58 is 347 m long (uncorrected and apparent controversial. It is included in the ‘‘phylloid algae’’ by e.g., thickness). Description of color in hand samples follows Konishi and Wray (1961) and Senowbari-Daryan and the Rock-Color Chart (Geological Society of America Rashidi (2010), although Baars and Torres (1991) and 1995) terminology. Almost 100 thin-sections were pre- Torres and Baars (1992) suggested that the thallus of pared and studied for microfacies analysis by traditional Anchicodium is cylindrical and branching, and hence the petrographic techniques. Thin-sections and polished slabs organism is not literally a ‘‘phylloid alga’’. Two genera were stained with combined Alizarin red S and potassium previously assigned to the ‘‘phylloid algae’’ differ funda- ferricyanide for distinguishing between ferroan phases in mentally from the anchicodiacean algae, Calcifolium calcites and dolomites (Dickson 1966). Fourteen thin-sec- Maslov 1956 and Archaeolithophyllum Johnson 1956. tions of algal-rich facies were studied in detail, from which Calcifolium is an algospongia (see Co´zar and Vachard two selected thin-sections (SS-14-4CT and SS-16-33.5CT) 2004), a group of incertae sedis algae recently revised by were polished and examined uncovered with a Cold Vachard and Co´zar (2010). Archaeolithophyllum, the Cathode Luminescence 8,200 mk3 equipment microscope questionable Kasimophyllum Mamet and Villa 2004, and (standard working conditions of 12-kV voltage, beam the problematic paleoaplysinaceans have been related to the current 150–180 mA, and chamber pressure 180–200 ancestral corallinaceans (e.g., Johnson 1956; Wray 1977b; mTorr). Microsamples (0.5–0.2 g) of different cement Co´zar and Vachard 2003; Vachard and Kabanov 2007; generations were analyzed for Mg, Sr, Mn, and Fe ele- Anderson and Beauchamp 2010). mental concentrations. They were collected using a high- Cathodoluminescence and epifluorescence microscopy speed dental drill with magnification from a binocular (e.g., Dawson 1992; Kirkland et al. 1993; Moshier and microscope. Analysis was performed using a Perkin-Elmer Kirkland 1993) have been successfully used to identify the ELAN 6000 inductively coupled plasma mass spectrometer cellular structure and nature of some ‘‘phylloid algae’’. (ICP-MS) at the University of Salamanca (Spain), with 123 Facies (2013) 59:949–967 951

123 952 Facies (2013) 59:949–967 standard calibration and an analytical uncertainty of ±2%. mudrocks (Lena and Sama groups). Distally (to the E), the Trace elements compositions (Ca, Mg, Fe, Sr, and Mn) of carbonate platform overthrusts the siliciclastic succession cements and algae were also determined by 155 electron of the Pisuerga-Carrioń Unit (Fig. 1a), mostly composed of microprobe measurements on two polished thin-sections, basinal mudrocks with local turbidites and olistoliths. The using a Cameca SX-100 electron microprobe by wave- Bachende Formation is sandwiched between two major and length dispersive (WDS) and energy dispersive analysis relatively shallow clastic units. It rests on the Bashkirian (EDS), housed at the University of Oviedo (Spain). A 5-lm Lois Formation, a thick interval (300–700 m thick) com- circular beam was used with an accelerating potential of posed of mudrocks and minor sandstones, and it is overlain 15–20 kV, beam current of 15–20 nA, and counting time of by the Duenãs Formation, a 300-m-thick succession of del- 10 s. Detection limits at the 95 % confidence level were taic siliciclastics with minor limestones interbedded at its 200 ppm for Fe, 210 ppm for Mn, 200 ppm for Mg, and base. The absence of a preserved and continuous deposi- 250 ppm for Sr. tional profile prevents an accurate description of the platform geometry. However, based on stratigraphic correlations and facies relationships, it has been suggested that the Bachende Geological setting and stratigraphy platform developed a relatively flat-topped geometry with slopes fronting the basinal areas (to the E/SE). The Salamoń The Cantabrian Zone constitutes the core of the Ibero- Gold District was situated very close to a probable slope of Armorican Arc and the external part of the Variscan Orogen the carbonate platform (Corrochano 2010). The presence of in the NW Iberian Peninsula. It shows typical features of this slope is supported by abundant carbonate breccia foreland fold-and-thrust belts, and is characterized by a thin- deposits and large amounts of marine cements in the skinned tectonic deformation style with almost absent boundstone facies (e.g., Tucker and Wright 1990). metamorphism and penetrative structures (Pe´rez-Estauń et al. 1988). On the basis of combined stratigraphic and structural Systematic paleontology features, Julivert (1971) and Pe´rez-Estauń et al. (1988) divi- ded the Cantabrian Zone into five geologic units: the Fold and Class Rhodophyta Wettstein 1901 Nappe, the Central Asturian Coalfield, the Ponga Nappe, the Order Archaeolithophyllales Chuvashov in Chuvashov Picos de Europa, and the Pisuerga-Carrioń (Fig. 1a). At least et al. 1987 nomen translat. Vachard and Kabanov 2007 during the Bashkirian-Moscovian, the Cantabrian Zone Family Archaeolithophyllaceae Chuvashov in Chuva- formed a wide (a few 100 km) asymmetric marine foreland shov et al. 1987, emend. Vachard et al. 2001 basin located at an equatorial position on the eastern coast of Pangea (e.g., Colmenero et al. 2002). This basin was pro- Remarks Johnson (1956) originally included Archaeo- gressively narrowed during the forward advance of the lithophyllum in the family Corallinaceae. Wray (1977b) Variscan orogenic front up to its final consumption at the suggested that this genus may belong to the ‘‘ancestral cor- Carboniferous-Permian boundary (e.g., Pe´rez-Estauń et al. allines’’ because of structural differences and the significant 1988; Colmenero et al. 2002). The Carboniferous foreland time hiatus between the last appearance of Archaeolitho- basin was mostly filled by thick, paralic, siliciclastic wedges phyllum in the Permian and the first occurrence of true high- that pass laterally into limestones, deposited on high-rising Mg calcite corallinaceans in the Early Cretaceous (e.g., carbonate platforms in the slowly subsiding distal areas (e.g., Aguirre et al. 2000). On the basis of inferred aragonitic Della Porta et al. 2004; Bahamonde et al. 2007). composition and the recognition of a questionable hypobasal The Salamoń Gold District is located in the southern part and extracellular aragonitic calcification, James et al. (1988) of the Lois-Ciguera sector (Central Asturian Coalfield, Can- and Moshier and Kirkland (1993) highlighted the similarities tabrian Zone, Fig. 1a), NE of Leoń (NW Spain). It is a Carlin- between Archaeolithophyllum and the Recent aragonitic red type deposit formed during the last events of the Variscan algae Peyssonneliaceae (formerly Squamariaceae); however, orogeny (Paniagua et al. 1996; Crespo et al. 2000), and it is a time hiatus also exits between the last Permian occurrence mostly hosted in the limestones and bituminous mudrocks of of Archaeolithophyllum and the first appearance of true the Bachende and Duenãs Formations (Fig. 1b). Peyssonneliaceae in the Early Cretaceous (Wray 1977a; The Bachende Formation (late Kashirian/early James et al. 1988). Myachkovian) consists of a 650-m-thick carbonate succession that represents a delta-top carbonate platform In the present paper, based on the similar reproductive (sensu Bosence 2005) developed in distal shelf areas of the structures (conceptacles), cellular organization and frame- foreland basin (Lois-Ciguera sector, Fig. 1a). To the W work strategies, the tentative hypothesis postulated by (landward), carbonate platform sediments alternate and Johnson (1956) and Co´zar and Vachard (2003) that the interfinger with deltaic sandstones and shallow-marine Archaeolithophyllaceae may represent the stem lineage of 123 Facies (2013) 59:949–967 953 the coralline algae (Corallinaceae, Melobesiaceae) is had little influence on carbonate skeletal mineralogy once reinforced. the mineralized skeletons evolve (Porter 2010). The genera Eugonophyllum, Ivanovia, Anchicodium, The study of Moshier and Kirkland (1993) presents Neoanchicodium, and other junior synonyms, such as several theoretical and analytical problems raised in the Calcipatera, Iranicodium, and Kansaphyllum, are generally discussion paper of Pingitore (1994). Therefore, with the distinguished from the Archaeolithophyllaceae because, aim to clarify the controversial mineralogy of Archaeo- according to cathodoluminescence, these algae display lithophyllum, additional microprobe analyses have been occasionally preserved structure similar to present-day performed herein (see following sections). The new results siphonous green algae, such as Udotea or Halimeda. The (from two different cores) show that the skeletal calcite of ‘‘diagenetic aspect’’ (after aggrading neomorphism and Archaeolithophyllum is currently composed of low-Mg subsequent microborings by endolithic algae) postulated by calcite with low Sr contents. These data do not necessarily Vachard et al. (2001) cannot be confirmed by this study. support an aragonitic precursor, and a high-Mg calcite Nevertheless, the assignment of several genera of ‘‘phyl- skeleton may be also considered. loid algae’’ to siphonous green algae remains questionable, A further argument made by James et al. (1988) and because they differ morphologically and microstructurally Moshier and Kirkland (1993) to support an original ara- from the rare true codiaceans known in the Pennsylvanian- gonitic composition of Archaeolithophyllum was the pres- Permian (e.g., Succodium Konishi 1954, Aphroditicodium ence of oriented aragonite inclusions preserved within Elliott 1970, Tauridium Gu¨venç 1966, and Thaiporella neomorphic calcite mosaics filling the intraskeletal pores Endo 1969). (Sandberg 1985, fig. 3A–D). However, a closer examina- tion of Sandberg’s description suggests that the presence of Discussion In addition to the temporal hiatus of the fossil such aragonite relics does not necessarily imply an original record, the original mineralogical composition of these aragonitic composition of the algae, but just reflects the algae requires discussion. Based on the high elevated Sr precipitation of aragonite cement in the intraskeletal pores. content in the skeletal tissue and aragonite relics growing Considering the exceptionally preserved anatomy of the in the intraskeletal pores (Sandberg 1985; James et al. Bachende Archaeolithophyllum (reproductive organs, cel- 1988; Moshier and Kirkland 1993) suggested that lular structure, cell fusion, and thallus organization), a Archaeolithophyllum was formerly composed of aragonite. strong similarity can be proposed between Archaeolitho- Because modern coralline algae secrete high-Mg calcite, phyllum and Recent corallinaceans (e.g., Johnson 1956; the phylogenetic relationship between Archaeolithophyl- Wray 1977a, b;Co´zar and Vachard 2003). However, it lum and modern corallinaceans resulted controversial. should be noted that morphological differences exist, apart Therefore, these authors postulated that Archaeolithophyl- from the time hiatus until the first occurrence of Corallin- lum was an ancestral aragonitic peyssonnelid alga. aceae in Early Cretaceous times. It has been suggested that secular variations in the Mg/ Reproductive organs are diagnostic criteria in most of Ca of seawater have influenced the polymorphic mineral- taxonomic classifications of modern marine and freshwater ogy of biotic and abiotic carbonates during the Phanerozoic red algae. Reproductive organs in Corallinaceae markedly (e.g., Sandberg 1983; Stanley 2006). Algae have biological differ from those observed in Peyssonneliaceae; in the controls over which calcite polymorphs they precipitate latter, the reproductive structures are confined to nemath- (Stanley et al. 2002; Ries 2006a, b), and cells seem to be ecia, which are external pustules that develop from able to control the crystallography and orientation of cal- simultaneous transverse divisions of surface cells forming cite crystals (Borowitzka 1989, cited in Konhauser 2007). a dome-shaped aggregation of assimilatory filaments Laboratory experiments with artificial seawater have (Denizot 1968). These structures are usually not preserved demonstrated that crustose red algae precipitate only the in fossil forms, because they are located in the outer margin calcite polymorphs even when the chemical conditions of of the plant, are noncalcified, and they flake off when seawater favors the precipitation of aragonite (mMg/ spores are released (Aguirre and Barattolo 2001). In con- Ca = 7) (Ries 2006b). Furthermore, coralline algae pos- trast, no nemathecia are formed in representatives of the sess sulphate galactans and alginates that preferentially Corallinaceae, and as analogous structures, calcified cavi- bind Ca2? over Mg2? resulting in microenvironments that ties grouped into sori or assembled into large cavities favor calcite precipitation over aragonite within their cell (conceptacles), contain the reproductive structures (Silva walls (Konhauser 2007). Consequently, a biomineralization and Johansen 1986). change from aragonite (Archaeolithophyllaceae) to high- Genus Archaeolithophyllum Johnson 1956 Mg calcite (corallinaceans) linked to secular variations in the Mg/Ca ratio of seawater appears unlikely, since oscil- Type species Archaeolithophyllum missouriense Johnson lations between calcite and aragonite seas appear to have 1956 123 954 Facies (2013) 59:949–967

Diagnosis Phylloid thallus, occasionally bifurcated, above the perithallus and are irregularly distributed over which occurs as isolated blades or foliate and encrusting the upper surface of the thallus; preserved apertures in multilayered masses; internal tissue differentiated into a these chambers are not observed (Fig. 2d). Crust divisions thick hypothallus, with arcuate rows of wide polygonal locally present on the upper surface of the thallus (Fig. 2a). cells, and a thinner perithallus, with small cells; knobby ‘‘Phylloid blades’’ occur attached during growth or devel- and spinose protuberances present on the upper and basal oped apparently free on the substrate. Attachment occurs at surface; conceptacles ovoid to highly arched, irregularly the edge of the blades through perithallus cells (Fig. 2e). distributed over the upper surface of the thallus, with a Cellular walls currently composed of low-Mg calcite single atypical aperture; cell fusions occasionally present. (0.5–2 mol % MgCO3) with low Sr concentrations (aver- Cellular walls currently composed of low-Mg calcite with age 412 ppm, range from below detection to 1,160 ppm, low Sr concentrations. n = 37). Remarks and comparisons The specimens described here Archaeolithophyllum asymmetricum sp. nov. closely resemble the type material of A. missouriense Figs. 2, 3, 4, 5, 6, 7 reported by Johnson (1956) in the Midcontinent, USA. 1993 Archaeolithophyllum sp. - Moshier and Kirkland: However, A. asymmetricum is strongly asymmetric, and the figs. 7, 9 perithallus layer is absent or poorly developed on one side. Some A. asymmetricum seem to have been illustrated as Etymology From the latinized Greek asymmetricus: Archaeolithophyllum sp. in Moshier and Kirkland (1993), devoid of symmetry. who also noted the strong asymmetry of many specimens in the Providence Limestone, USA (p. 1035). The Holotype Fig. 2a-e. Thin-section SS-14-4-CT. arrangement of the perithallial cells in vertical rows per- Repository Department of Geology, University of pendicular to the thallus surface is another important dis- Salamanca (Spain). tinctive feature of A. asymmetricum, because in previously described species these cells are arranged in rows parallel Type locality Bachende Formation in the Salamoń Gold to the surface of the thallus. Finally, the presence of cell District (42°5604700N and 5°705200W), in the Lois-Ciguera fusion, the overall small diameter of either the hypothallic sector of the Cantabrian Zone (NW Spain). and perithallic cells, and the thallus and the perithallus Type level Late Kashirian/early Myachkovian time thickness, distinguish A. asymmetricum from other Ar- interval of the Bachende Formation. chaeolithophyllum (see review table in Co´zar et al. 2005). Diagnosis A species of Archaeolithophyllum characterized by (a) the strong asymmetry of thallus (i.e., a peri- Lithofacies and petrographic description thallus unilaterally developed on the lower side), (b) the arrangement of the perithallial cells, (c) the cell fusions, Archaeolithophyllum asymmetricum specimens were col- and (d) highly arched conceptacles without any preserved lected from a pale yellowish brown (10YR6/2) micritic aperture. limestone (16–19.5 m thick) located in the uppermost part Description ‘‘Phylloid algal’’ thalli up to 3.2 cm long and of the studied cores (Fig. 1b). The micritic interval shows a up to 680 lm thick (averaging 280 lm). Calcified thallus lenticular morphology that is evident from stratigraphic with an internal tissue well differentiated into a thick correlations between different cores in the Salamoń Gold medullary hypothallus and a thin upper perithallus. District, and from field observations in surrounding areas Hypothallus coaxial and multilayered, consisting of wide (facies D2 of Corrochano et al. 2012). polygonal cells (up to 30–40 lm wide and 80–90 lm long) The A. asymmetricum accumulations have a dominantly that form arched rows in longitudinal sections. The peri- muddy matrix and contain common to abundant (20–40 % thallus is up to 45 lm thick and it is made up of smaller of the rock volume) remains of ‘‘phylloid algal’’ plates. cells (up to 6 lm wide and 14 lm long), nearly rectangu- Algal boundstones and wackestones (subordinate pack- lar, which are arranged in rows perpendicular to the surface stones and floatstones), and algal cementstones are the of the thallus (Figs. 2b, c, 3a, b). Cell fusions present as dominant microfabrics. Archaeolithophyllum dominated on voids extending from a cell from one filament to a cell or the substrate, creating an almost monospecific community cells of contiguous filaments in the perithallus (Figs. 2c, e, formed by undulate algal blades that grew attached, closely 3b). Thallus locally swelled keeping similar size of skeletal packed and juxtaposed near and above one another, pro- elements (Fig. 2a). Highly arched and protruding concep- ducing a dense and open framework (Figs. 3e, f, 4, 5a). tacles (up to 150 lm wide and 100 lm high) are preserved Most of the framework voids are filled with early marine

123 Facies (2013) 59:949–967 955

Fig. 2 Cathodoluminescence images of Archaeolithophyllum asym- Fig. 2a showing the strong asymmetry of the phylloid thallus with the metricum sp. nov. a Section through the holotype specimen (thin- perithallus layer developed only on the upper surface of the alga. section SS-14-4CT) showing the exceptional preservation of the c Enlargement of Fig. 2b showing the perithallial cells arranged in thallus organization. Note the conceptacles, the hypothallial coaxial rows perpendicular to the surface of the thallus. Note cell fusions of organization, the swelling of the thallus (black arrow) at the right side adjacent filaments (arrows). d Highly arched conceptacles (arrows) of the photograph, a crust fusion at the left side (white arrow), and the developed on the upper surface of the thallus. Note absence of development of shelter porosities beneath the algal thalli; these voids preserved apertures and the coaxial organization of the hypothallus. are usually filled by radiaxial fibrous calcite and subsequent zoned e Magnification of a crust fusion through perithallial cells. Arrows blocky spar (left part of the image). For magnification of point to cell fusions encased zones, see Fig. 2b–e. b Magnification of quadrangle in radiaxial-fibrous cements (RFC) (Fig. 4). Subordinate fau- and Claracrusta catenoides. Other bioclasts, such as bry- nal elements include organisms that mostly lived attached to ozoans, solitary rugose corals, brachiopods, ostracods, the phylloid thalli, such as Tuberitina, Palaeonubecularia, echinoderms, gastropods, algosponges, Thartharella,

123 956 Facies (2013) 59:949–967 lasiodiscids, and Ozawainella, are sporadically associated Fig. 3 Archaeolithophyllum asymmetricum sp. nov. a, b, d, f c (Fig. 5a). Cathodoluminescence images. c, e Planar polarized light images. a Specimen embedded in a clotted and peloidal micrite that shows a Under plane-polarized light, many A. asymmetricum mottled cathodoluminescence (CL). Note the well-developed peri- specimens resemble recrystallized ‘‘phylloid algae’’, in thallus at the right side of the thallus and the arrowed gastropod. which internal structure was dissolved and replaced. b Detail of the same specimen, showing perithallial cells arranged in Cathodoluminescence (CL), however, has revealed the rows perpendicular to the surface of the thallus. Note the polygonal morphology of the hypothallial cells. Arrows point to cell fusions. skeletal features of these algae (Fig. 3c, d). Skeletal walls c Cathodoluminescence and d planar polarized light photomicro- of A. asymmetricum are composed of dull-bright (locally graphs of the same specimen, showing a shelter porosity developed bright-yellow) luminescent calcite, which contrasts sharply beneath the algal thallus, which is filled with radiaxial fibrous with nonluminescent calcite cements filling the intraskel- cements (RFC) and subsequent zoned blocky spar (BS). Note in Fig. 3c the cloudy appearance of the RFC, and in Fig. 3d, the etal voids. Staining shows that both phases, skeletal walls irregular contact between both cement phases; also note the lower and cell-filling cements, are nonferroan calcite. A. asym- irregular contact between RFC and peloidal micrite. e Archaeolitho- metricum occurs as elongate and arcuate sheets that under phyllum framework showing the typical ‘‘phylloid appearance’’ of transmitted plane-polarized light are preserved as calcitic these algae under planar polarized light. RFC radiaxial fibrous calcite; BS blocky spar. White arrows point to crust fusions and black arrow mosaics of tiny anhedral to subhedral crystals averaging indicates late diagenetic calcite veins. Note the clotted and peloidal 30 lm that include both skeletal walls and cell-filling micrite surrounding the algal thalli. f Cathodoluminescence detail cements (Fig. 3c, d). Crystals are generally wider towards view of a crust fusion through perithallial cells. Note the occurrence the center of the thallus (Fig. 5b), where they can reach of radiaxial fibrous calcite (RFC) and the presence of diagenetic coarse-grained crystals with dull and bright luminescence (white 145 lm, and generally are finer than those observed in arrow) other ‘‘phylloid algae’’ (e.g., Wray 1977a, fig. 86; Toomey 1980, fig. 11; Mamet and Villa 2004, figs. 12, 13). Many crystals show micron-sized dark impurities grouped in length. They show a cloudy appearance, caused by abun- botroydal masses 20–30 lm wide that usually occupy the dant inclusions of tiny crystals (Figs. 3c, 5b). nucleus of the crystals. In some cases, the cement intercalates thin micritic films The matrix surrounding the Archaeolithophyllum thalli that displays dull-bright CL. RFC show a discontinuous is dominantly micritic, and it might be subdivided into strip mottled CL parallel to the fibers and are nonlumi- homogeneous or clotted and peloidal micrite (Fig. 3e). The nescent with local dull-bright (locally bright-yellow) clotted and peloidal micrite surrounding the algal thalli elongated patches (Fig. 3d, f). This pattern is similar to display a mottled CL (Fig. 3a), and consists of peloids and other RFC observed in Carboniferous limestones of the aggregate grains (dark-dull CL) rimmed by a discontinuous Cantabrian Zone (e.g., Van Der Kooij et al. 2007). bright rind, followed by an interpeloidal nonluminescent RFC are followed by coarse, blocky and nonferroan fine sparite cement. Peloids consist of subspherical to calcite cements, as evidenced by pink staining. This tran- irregular aggregates of micrite with poorly defined outlines, sition in the void-filling cements is marked by an irregular averaging 80–100 lm in diameter. They could be closely contact that also records the change from a ‘‘cloudy’’ to packed or largely spaced, resulting in close or open fabrics. ‘‘clear’’ appearance of the calcite. In the blocky spar Ellipsoidal pellets (up to 520 lm wide) are also present crystals, there are several zoned generations with distinct floating in the matrix. luminescence patterns (Figs. 2a, 3d). The first generation is A. asymmetricum accumulations are characterized by composed of dull-bright calcite, with a thin bright inter- the formation of growth cavities that exhibit abundant calated phase. It is followed by dull calcite that displays fabric-selective primary porosity, which is usually lined by thin alternating phases of bright and nonluminescent calcite nonferroan RFC (sensu Kendall and Tucker 1973), finally separated by sharp boundaries. Finally, the last cementa- occluded by a blocky sparite (Figs. 2a, 3c–e); locally, tion stage consists of nonluminescent calcite. primary pores are filled-up by laminated geopetal reddish The ‘‘phylloid algal’’ blades, the micritic matrix and the micrite. These cavities constitute shelter porosities located different cement phases described, are cross-cut by a ran- below the algal thalli, although some pores are surrounded dom network of irregular veins filled by late diagenetic dull and supported by the peloidal micritic sediment. Geopetal or nonluminescent ferroan and nonferroan calcites (Fig. 3a, inner sediments always postdate fibrous cementation and b). only occlude the lower two-thirds of the void space; the Although under standard microscopy A. asymmetricum upper one-third is occluded by blocky spar crystals. The from the Bachende Formation is very similar to those RFC exhibits undulating extinction and forms isopachous specimens reported in the Providence Limestone by Mos- crusts with an average thickness of 1.6 mm. It represents hier and Kirkland (1993), the ‘‘chemical preservation’’ of 25–30 %, locally over 50 %, of the rock volume, and is these algae is quite different, suggesting that exceptional formed by elongate fibers that usually are 0.5–1 mm in preservation of ‘‘phylloid algae’’ could occur under 123 Facies (2013) 59:949–967 957

different diagenetic conditions and pore-water composi- the Bachende Formation it is not revealed. In the former, tion. Chemical staining in the Providence Archaeolitho- ferroan calcite ﬁll cell voids, whereas in the Bachende phyllum shows the cellular structure of the algae whereas in Formation these cell-void cements are nonferroan calcite

123 958 Facies (2013) 59:949–967

Fig. 4 Core photograph (a) and diagram drawing (b)ofArchaeo- cements. A A. asymmetricum, RFC radiaxial ﬁbrous calcite, B blocky lithophyllum asymmetricum nov. sp. framework. Note that primary spar, F late diagenetic fractures and stylolites pore space is commonly ﬁlled by pervasive early diagenetic marine

Fig. 5 a Framework of Archaeolithophyllum asymmetricum nov. sp. view of a thallus of A. asymmetricum preserved as a calcitic mosaic of Note the foraminiferal encrustations (F) in upper left. A A. asym- tiny anhedral to subhedral crystals that become wider toward the metricum, RFC radiaxial ﬁbrous calcite, OzOzawainella. b Close-up center of the thallus

(85 % of the samples have Fe concentrations below Geochemical data 700 ppm). In addition, in the Providence Limestone, these algae display a zoned cell-ﬁlling calcite (bright-yel- Trace element concentrations of Archaeolithophyllum low ? dull-red ? dull-yellow luminescence) with rela- asymmetricum and associated framework cements are tive high concentrations of Mn and Fe, whereas in the presented in Table 1 and Fig. 6. In general, concentrations Bachende Formation they show a nonluminescent calcite. measured on the microprobe slightly differ from those Furthermore, CL preservation of skeletal walls is also obtained using ICP-MS, although they show a good cor- different at both localities, and in contrast to the Bachende relation; these differences are probably due to the mixing specimens, Providence Archaeolithophyllum skeletons of different cement phases during sampling for ICP-MS. display nonluminescent calcite walls (Moshier and The location of the four transverses (two per thin-sec- Kirkland 1993). tion) for microprobe is shown in Figs. 3c, d and 7.

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Table 1 Mean value (and range) of minor element content (ppm) in the skeleton of Archaeolithophyllum and marine cements Trace element (ppm) Mg Mn Fe Sr Population Measuring device

SS-14-4-RFC 2,436 61 715 133 ICP-MS Radiaxial-fibrous cement 3,688 bdl bdl 446 n = 35 Microprobe (1,516–5,596) (bdl-1674) (bdl-1557) (bdl-723) SS-14-4-B 1,476 41 740 187 ICP-MS Blocky calcite 2,965 bdl bdl 433 n = 20 Microprobe (1,342–12,310) (bdl-340) (bdl-555) (bdl-706) SS-14-4-V 1,802 121 2,672 211 ICP-MS Calcite vein 1,793 285 3,754 398 n = 7 Microprobe (1,030–3,048) (bdl-454) (959-5737) (bdl-727) Micrite 2,967 bdl bdl 347 n = 5 Microprobe (2,300–4,016) (bdl-278) (bdl-2068) (bdl-436) Archaeolithophyllum 3,213 217 276 412 n = 37 Microprobe (1,168–5,003) (bdl-540) (bdl-1469) (bdl-1160) Cell-filling cement 3,438 bdl 275 417 n = 47 Microprobe (1,901–6,685) (bdl-1840) (bdl-729) RFC radiaxial fibrous calcite, B blocky calcite, V burial calcite veins, bdl below detection limit

Fig. 6 Microprobe transverses illustrated in Figs. 3c, d and 7. Note that Archaeolithophyllum skeletal tissue (bright-dull CL pattern) is enriched in Mn with respect to the cell-ﬁlling cements (nonluminescent)

Transverses B–B0 and D–D0 show the trace element vari- concentrations of these cations or their ratio), with Mn2? as ations between the algal skeleton (dull-bright lumines- the main activator and Fe2? as the main quencher of lumi- cence) and the cell-filling cements (non-luminescence), and nescence (e.g., Hemming et al. 1989; Machel et al. 1991; the other two (A–A0 and C–C0) were carried out on the Boggs and Krinsley 2006). The quencher concentrations algal skeletons and radiaxial-fibrous calcites. necessary for extinction of Mn-activated luminescence The cathodoluminescence intensity (brightness) results (15–20 ppm are sufficient to produce detectable lumines- from compositional changes in calcite cements, especially cence) have not been determined with sufficient accuracy, from Mn2? and Fe2? concentrations (either the absolute and probably depend on the Fe/Mn ratio (Machel et al. 1991), 123 960 Facies (2013) 59:949–967

Fig. 7 Planar polarized light (a) and cathodoluminescence (b) pho- fibrous calite, B blocky spar. Note a shallow burial fracture filled with tomicrographs showing the microprobe transverses C–C0 and D–D0 dull calcite (arrows) cutting the nonluminescent RFC and the algal illustrated in Fig. 6 (thin-section SS-14-16). A Archaeolithophyllum skeleton, all of them cut by a late burial vein asymmetricum sp. nov., V late burial calcite vein; RFC radiaxial- although variable Fe/Mn ratios and absolute proportions of Discussion either cations have been suggested in the literature (Boggs and Krinsley 2006; and references therein). Paleoecology of Archaeolithophyllum asymmetricum Therefore, due to the prior uncertain identification (or differentiation) of skeletons and cell-filling cements, the New models for the habit and growth mode of ‘‘phylloid Mn contents in transverses B–B0 and D–D0 are interpreted algal’’ communities have been put forward in the last years to represent the luminescent skeletal walls of the algae, (e.g., Samankassou and West 2002, 2003; Forsythe 2003). whereas the nonluminescent areas are identified as the Rigid cement-rich ‘‘phylloid algal’’ frameworks with a cements filling the intraskeletal voids, which in turn are positive relief might be usually assigned to erect and characterized by lower Mn concentrations (Table 1 and cyathiform ‘‘anchicodiacean’’ algae, such as Ivanovia or Fig. 6). ICP-MS analyses have not been realized in the Eugonophyllum (e.g., Samankassou and West 2002; algal blades (either the skeletal walls or the cell-filling Forsythe 2003; Enpu et al. 2007a). Nevertheless, (a) this cements), because it was technically impossible to isolate reconstruction differs sensibly from many previous ones the algal plates from the adjacent matrix and RFC. (e.g., Konishi and Wray 1961; Heckel and Cocke 1969; Based on microprobe analysis, the skeleton of Archae- Frost 1975; Toomey and Cys 1979; Toomey 1983; Morin olithophyllum is currently composed of neomorphosed et al. 1994; Krainer 1995; Krainer et al. 2003), and low-Mg calcite (0.5–2 mol % MgCO3). It shows Sr values (b) these algae have been preferentially assigned to that range between below detection limit to 1,160 ppm Anchicodium sensu lato (Baars and Torres 1991; Krainer (average 412 ppm). Fe contents average 276 ppm (range et al. 2007 with references therein). Conversely, according between below detection to 1,469 ppm), and Mn values to Samankassou and West (2002), Forsythe (2003), and range between below detection to 540 ppm (average Enpu et al. (2007a), the ‘‘phylloid’’ rhodophycean 217 ppm). Cell-filling cements have a similar trace element Archaeolithophyllum might be usually associated with low composition, although the Mn concentration is significantly energy and muddy shelf environments, playing a passive lower (below detection limit). and secondary role as a constructor in build-up develop- RFC in the Archaeolithophyllum framework are cur- ment. In fact, due to its prostrate growth form, this red alga rently composed of low-Mg calcite (0.6–5 mol % might usually form biostromes without any positive

MgCO3), and show trace element concentrations similar to topography (Forsythe 2003). Nevertheless, the encrusting the skeleton of Archaeolithophyllum and to other marine- form A. lamellosum locally formed semi-rigid crusts phreatic RFC reported in the literature (Richter et al. 2011, capable of providing a self-supporting skeletal framework table 3). RFC in the Bachende samples show Sr values that with a remarkable sediment-binding function in the depo- range between below detection limit to 723 ppm (average sitional environment (Wray 1964; Toomey 1980). The 446 ppm), and Fe and Mn concentrations that average membranous form A. missouriense is generally related in below detection. the literature with open shelf banks formed by fragmented

123 Facies (2013) 59:949–967 961 and undulating algal thalli floating in a muddy matrix with processes, it is reasonable to infer that the ‘‘phylloid algal’’ variable amounts of shelter porosities (constructional facies were most extensively developed in relatively shal- mounds of Samankassou and West 2002;Type3of low waters within the photic zone, but below fair-weather Samankassou and West 2003; Archaeolithophyllum com- wave-base. Most models suggest that Late Paleozoic munity of Forsythe 2003; and para-autochthonous occur- phylloid accumulations occurred in widely distributed belts rences of Vera et al. 1984). along open and shallow carbonate shelves or ramps, at Toomey (1976) suggested that dense meadows of water depths of approximately 15–30 m, where red algae ‘‘phylloid algae’’ induced community restrictions on the usually inhabit lower light environments. Nevertheless, seafloor, because the algae dominated the available living because modern coralline algae inhabit a wide depth range, space; the only organisms that could survive in such ranging from intertidal settings to 270 m water depth environments were epiphytic organisms that lived attached (Littler et al. 1986), with an acme between 60 and 120 m on the phylloid blades, such as encrusting foraminifers, the (Aponte and Ballantine 2001), to provide an approximate grazing animals that could browse and feed on them (e.g., depth for the studied accumulations appears to be specu- gastropods), and those confined to the framework cavities. lative. Davies et al. (2004) concluded that the Late The matrix surrounding Archaeolithophyllum asym- Paleozoic Archaeolithophyllum mounds were similar to the metricum is composed of homogeneous and/or clotted and Holocene Mesophyllum buildups that were observed in peloidal micrite. Comparable peloidal fabrics have been deep-water (80–120 m) and outer platform environments described by several authors from Paleozoic build-ups as on the Eastern Australian Shelf. microbially precipitated (e.g., Pratt 1995). These deposits It is accepted in the literature that development of Late result from the trapping, binding, calcifying, and metabolic Paleozoic phylloid algal buildups was mostly controlled by activities of elements of benthic communities, such as high-frequency glacioeustatic sea-level fluctuations; these cyanobacteria, bacteria, algae, and fungi (Riding 2000). deposits are commonly integrated in shallowing-upward Based on the elevated content of the peloidal fabrics, it is sequences capped by subaerial exposures or oolitic shoals inferred that microbial activity played an important role, (e.g., Toomey 1980; Dawson and Carozzi 1986; Gold- although secondary, in the mound growth. hammer et al. 1991; Soreghan and Giles 1999). Because There are several genetic models that have been pro- A. asymmetricum mounds lack any evidence of glacioeu- posed to explain internal fabrics of Pennsylvanian ‘‘phyl- static falls despite the moderate amplitude sea-level chan- loid algal’’ mounds (for review see Samankassou and West ges occurring during Moscovian times (at least 40 m, Rygel 2002). The genetic model suggested in the present paper is et al. 2008), a water depth of several tens of meters could quite similar to the ‘‘cup-model’’ proposed for assumed be inferred during the growth of these communities. ‘‘udoteacean’’ algae by Samankassou and West (2002)in However, this interpretation must be regarded with caution, the Frisbie Limestone of Kansas, USA, or the Eugono- because the upper part of the Bachende Formation displays phyllum community reported by Forsythe (2003) in the high subsidence rates that could preclude exposure Hueco Mountains outliers (La Colmena and the Butterfield (Corrochano et al. 2012) and thus, the absence of subaerial Trail Mounds), Texas, USA, where several processes, exposure features might not be indicative of a deep-water acting simultaneously, induced mound-building. Baffling environment. and trapping of lime mud and bioclastic carbonate particles The presence of brachiopods, bryozoans, and solitary by Archaeolithophyllum, physical accumulation of phylloid corals also suggests open-marine and well-oxygenated blades, the dense framework formed by attached algal waters (Wilson 1975). Furthermore, the foraminifers thalli, pervasive early marine cementation, and micritic Ozawainella of the Moscovian of the Cantabrian Zone, as precipitation, were the most important processes on mound the Fusulinida in general, are consistent with open marine, growth. The dense and rigid framework resulted from the outer platform environments (Della Porta et al. 2005). The foliaceous nature of the algae and various methods of crust occurrence of in situ algal thalli surrounded by clotted and division and fusion, developing a comparable structure to peloidal micrite, and the absence of tractive structures, that reported by Bosence (1983, 1985) for the corallinacean suggests relatively quiet areas below storm wave-base. Mesophyllum in the Miocene of North Malta and in modern This depositional environment is consistent with the open sediments from the southern Gulf of Lion. The presence of Mesophyllum frameworks reported by Bosence (1983, geopetal structures supports the inferred rigidity of the 1985), who suggested ‘‘quiet water settings of moderate algal framework. depth’’. The depositional environment assumed in this paper for Primary framework voids were rapidly cemented by A. asymmetricum agrees well with those most commonly synsedimentary RFC followed by a diagenetic blocky spar. accepted for ‘‘phylloid algal’’ accumulations worldwide. This cementation occurred after biogenic encrustations of Because algae depend on sunlight for important metabolic phylloid blades, and would have provided a rigid 123 962 Facies (2013) 59:949–967 framework for the mounds to maintain their aggradational precipitation from sea water without significant diagenetic growth (Roylance 1990). The shelter voids are usually alteration. These data also support a mineralogical stabil- found beneath the Archaeolithophyllum thalli, although ization under slow and/or restricted interchange rock-fluid locally, some blades are completely coated, suggesting that after early cementation and progressive burial compaction. cements grew both from the floor and the roof of the The burial increase would result in a progressive reduction cavities (upward and downward growth). in the Sr/Ca ratio of the diagenetic calcites (Baker et al. 1982), and hence, the Sr loss is consistent with a high-Mg calcite predecessor. Diagenesis Cathodoluminescence observations and trace element compositions (Table 1, Fig. 6) show that the nonlumines- Cement phases cent RFC were probably synchronous with the nonluminescent calcite filling the inter-peloidal voids of the matrix It is proposed that the overall cementation stages in the and the nonluminescent cell-filling cements of the algal studied samples occurred under progressive burial marine skeleton. The nonluminescent pattern confirms a cemen- conditions during early diagenesis; meteoric influence can tation stage under oxidizing conditions in a marine phreatic be ruled out by the lack of petrographic, sedimentologic or environment (e.g., Moore 1989). These conditions are isotopic (d18O and d13C; Corrochano and Barba 2007) consistent with their low Mn (below detection limit) and Fe evidence of subaerial exposure. compositions (Table 1), and in brachiopod shells, have Radiaxial-fibrous marine calcites are common cements been interpreted as indicative of a marine signal (Popp filling primary voids in ancient limestones. They have been et al. 1986; Bruckschen et al. 1999). The early cementation interpreted as (a) neomorphic replacement products of resulted in a rapid lithification of the sediment, and oxi- aragonite cements (Kendall and Tucker 1973; Bathurst dizing conditions favored the Mn presence in marine 1977) or high-Mg calcite cements (Wilson and Dickson cements mainly as Mn4?, and the Fe presence as Fe3?, 1996), and (b) primary marine-phreatic high-Mg calcite rather than the divalent forms Mn2? and Fe2?. cements (e.g., Kendall 1985; Saller 1986; Flu¨gel and Koch Although most of the RFC samples lack conspicuous 1995; Nelson and James 2000; Richter et al. 2011; among imprint of diagenesis alteration, some of them (27 %) have others). The present consensus favors a marine-phreatic Mn values of more than 250 ppm, which are considered high-Mg calcite precursor (Flu¨gel 2004). One of the diagenetically overprinted. During stabilization process, strongest lines of evidence for high-Mg calcite composition RFC could have been affected by diagenetic alteration as is the presence of abundant dolomite inclusions that give a revealed by the mottled and patchy dull-bright CL (sup- turbid appearance to the RFC (Lohmann and Meyers 1977; ported by the sporadic high Mn content, up to 1,674 ppm, Davies 1977; Kendall 1985; Saller 1986). However, as Fig. 6) usually observed in the intermediate part of the pointed out by Saller (1986), this interpretation must be fibers (Figs. 1a–c, 2d). This alteration is probably coeval regarded with caution, because radiaxial calcites in the with the stabilization process of the algal skeleton, which Enewetak Atoll are variable in composition and range from shows a similar bright-dull CL pattern (Figs. 2b, c, 3f) and low-Mg calcite (1.6 mol % MgCO3) to moderately high- trace element composition (Table 1). Mg calcite (11.1 mol % MgCO3). Saller (1986) interpreted The precipitation of blocky and well-concentric-zoned these variations as the result of original heterogeneity in the calcite cement occluding remaining void space occurred calcite precipitation. under deeper burial conditions, and probably reflects fur- In the Bachende Formation, based on morphology, ther changes in Eh or progressive changes in pore-fluid cathodoluminescence, occurrence, good preservation, chemistry. Figure 7 illustrates cross-cutting relationships inclusion pattern, and similarity to other study cases (see between a calcite vein associated with the blocky calcite, references above), a primary high-Mg calcite cement revealing that the blocky calcite postdates the nonlumi- formed in the marine-phreatic realm is suggested. More- nescent cements and the chemical stabilization of the algal over, the isotopic values of fibrous cements reported by skeleton. Corrochano and Barba (2007) in the Salamoń Gold District Deep burial diagenesis is represented by a random net- that exhibit a mean value of -2.1 % for d18O and 5.14 % work of at least two generations of irregular and thin veins, for d13C, are in good agreement with other published data filled up by dull and nonluminescent ferroan to nonferroan of marine-phreatic cements in the Cantabrian Zone (e.g., low-Mg calcite, which postdates and, in some instances, is Van der Kooij et al. 2007, 2009), and with the predicted coeval with fracturing and chemical compaction. The composition of the Carboniferous Paleo-Tethyan seawater depleted Mg content is consistent with a burial diagenetic (e.g., Bruckschen et al. 1999). This suggests that the RFC environment, where pore waters have low Mg/Ca ratios in this study have an isotopic composition consistent with and cements are precipitated as low-Mg calcite. The high 123 Facies (2013) 59:949–967 963

Fe and Mn values, which are probably derived from the chemical reactions occur along a microscopic diage- associated clay minerals or organic matter (e.g., Jenkyns netic front (Bathurst 1975). In the present study, chemical et al. 2002), are consistent with a burial environment and imprints of A. asymmetricum specimens allow the excep- also support the quenching of Mn-activated luminescence. tional recognition of the skeletal features of the algae under Finally, all of these cementation phases appear to have CL. This suggests that the stabilization process affecting undergone further neomorphic alterations that are revealed the algal skeleton was the result of in situ replacement, and by anomalous coarse-grained crystals, randomly distrib- not a solution removal followed by a later void-fill cement uted in all the phases, with a bright to dull zonation precipitation. As proposed by Moshier and Kirkland (1993) (Fig. 3f). in the Providence Limestone, and because cell walls show no clear evidences of corrosion, it is inferred that the Archaeolithophyllum asymmetricum preservation preservation of the algal structure was favored by a rapid marine cementation of the cell pores. This cementation Because poor preservation of internal microstructure would have protected skeletons from dissolution and would affects most ‘‘phylloid algae’’, Dawson and Carozzi (1986) have promoted slow rates of neomorphism by reducing the suggested that the susceptibility of these algae to dissolu- microporosity and fluid exchange (Moshier and Kirkland tion and recrystallization is probably a function of its ori- 1993). ginal metastable mineralogy and the highly porous nature Recent high-Mg calcitic coralline algae contain of their original skeletal microstructure. In that sense, 1,500–5,600 ppm Sr (e.g., Milliman 1974; Veizer 1983), although it has been demonstrated in laboratory that the and Carboniferous high-Mg calcite in exceptionally pre- presence of metastable carbonate polymorphs is not served echinoderm skeletons contains 620–3,710 ppm Sr essential for the development of algal moldic porosity (Dickson 1995). Strontium values of Archaeolithophyllum (Dawson and Carozzi 1993), this type of neomorphic fabric in the Bachende Formation range between below detection is mainly related to preferential dissolution of aragonite, limit to 1,160 ppm (average 412 ppm). These concentra- and to a lesser extent, of high-Mg calcite (e.g., Moore tions are consistent with the expected loss associated with 1989). diagenesis, because most diagenetic waters have Sr/Ca The microprobe results presented herein (Table 1) indi- ratios less than that of seawater and the distribution coef- cate that the skeleton of Archaeolithophyllum is currently ficient of Sr in calcite is much below the unity. As the Mg composed of neomorphosed low-Mg calcite (0.5–2 mol % content of marine carbonates fluctuates within a wide

MgCO3) with low Sr content. Because modern red algae range, it is therefore on their own not necessarily helpful in secrete metastable carbonate like high-Mg calcite (coralli- determining the diagenetic history (Bruckschen et al. naceans) or aragonite (peyssonneliaceans), both are possi- 1999), but a reduction due to diagenesis in biogenic car- ble as the mineralogy of Archaeolithophyllum. Following bonates is also expected (e.g., Dickson 1995). the criteria summarized by Sandberg (1983) in determining Recent coralline algae have mean Mn concentrations of original aragonite mineralogy (e.g., presence of aragonite 45 ppm (Milliman 1974) and hence, the values reported in relics, elevated Sr content), in the Bachende samples there this paper (40.5 % of the samples have Mn content of more is not enough evidence to suggest an aragonitic precursor. than 250 ppm) may indicate a diagenetic phase with In contrast, based on the (a) morphological similarity influence of reducing conditions that probably took place in (especially reproductive organs) between Archaeolitho- a shallow burial environment. During burial of carbonate phyllum and modern calcitic corallinaceans and (b) similar sediments, exchange of pore waters with normal marine CL properties and trace element composition of the algal water may be restricted, and the consequent reduction in thalli and the surrounding high-Mg RFC, a high-Mg calcite Eh will increase the presence of Mn2? in the cements, and precursor is suggested. The different modes of preservation thus, a bright luminescent pattern could be expected. of Archaeolithophyllum elsewhere, which range from good The differential Mn incorporation (and the subsequent to completely obliterated, was interpreted by Wray (1977a) different CL) of single crystals that under plane polarized as the result of the variable Mg contents of the high-Mg light include both cell-filling cements and skeletal walls calcites. remains uncertain, although it is probably related to the It is generally agreed in the literature that high-Mg relative retention of the original trace element composition calcite generally transforms to low-Mg calcite with little of the algae, or probably reflects differences in the Mn loss of textural detail (e.g., Bathurst 1975; James and distribution coefficients between the two different phases. Choquette 1984), although biomoulds of high-Mg calcite Machel and Burton (1991) suggested that distribution skeletal grains have also been reported (e.g., Budd 1992; coefficients change due to variations in activity coeffi- Melim et al. 2001; and references therein). Such stabil- cients, temperature, crystal growth ratios, and microstruc- ization usually retains the original microstructure because ture. In that sense, because skeletal microstructures 123 964 Facies (2013) 59:949–967 strongly influence the mechanism of biogenic high-Mg represent the phylogenetic lineage of modern coralline calcite stabilization (e.g., Henrich and Wefer 1986; Budd algae. 1992), the shape and size of the individual skeletal crystals and their arrangement could also have played a major role Acknowledgments Thanks are due to P. Barba for logging assis- in the differential neomorphism of the cell-filling cements tance. SIEMCALSA S.A. is thanked for all the facilities providing and the algal skeleton. access to the core material of the Salamoń Gold District. The con- structive reviews and helpful suggestions of Brenda Kirkland and Elias Samankassou, strongly improved the original manuscript. This work was supported by the Spanish MICINN project CGL2004- Conclusions 02645/BTE.

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