Brasiliporella, a new mid-Cretaceous dasycladacean genus: the
Transcrição
Brasiliporella, a new mid-Cretaceous dasycladacean genus: the
Brasiliporella, a new mid-Cretaceous dasycladacean genus: the earliest record of the Tribe Batophoreae Bruno Granier, Dimas Dias-Brito, Ioan I. Bucur & Paulo Tibana Facies International Journal of Paleontology, Sedimentology and Geology ISSN 0172-9179 Facies DOI 10.1007/s10347-012-0312-6 1 23 Your article is protected by copyright and all rights are held exclusively by SpringerVerlag. This e-offprint is for personal use only and shall not be self-archived in electronic repositories. If you wish to self-archive your work, please use the accepted author’s version for posting to your own website or your institution’s repository. You may further deposit the accepted author’s version on a funder’s repository at a funder’s request, provided it is not made publicly available until 12 months after publication. 1 23 Author's personal copy Facies DOI 10.1007/s10347-012-0312-6 O R I G I N A L A R T I CL E Brasiliporella, a new mid-Cretaceous dasycladacean genus: the earliest record of the Tribe Batophoreae Bruno Granier · Dimas Dias-Brito · Ioan I. Bucur · Paulo Tibana Received: 6 April 2012 / Accepted: 10 May 2012 © Springer-Verlag 2012 Abstract New material collected in Albian and Cenomanian strata from Brazil helps us to better understand the structure of a poorly known dasycladacean alga, Holosporella nkossaensis P. Masse, in Bull Centr Rech Explor-Prod elf aquitaine, 19: 301–317, 1995: each of its fertile laterals, which are regularly arranged in verticils along the cylindrical algal thallus, consists of a distally inXated primary bearing two vesicular ampullae in terminal position. These traits are not known in representatives of the genus Holosporella Pia, 1930 nor in any genera described to date. On this basis, we introduce the new genus Brasiliporella with Brasiliporella nkossaensis emend. nov. comb. as its typespecies. We also discuss the systematic aYnity of the new taxon: it is ascribed to the Tribe Batophoreae, and in a broader manner the current paleontological ‘interpretation of the concept’ (in French: ‘acception’) of the Order Dasycladales, with the creation of two new families and accordingly with the emendation of two other families. Keywords Dasycladales · Dasycladaceae · Bornetellaceae nov. fam. · Triploporellaceae · B. Granier (&) Département des Sciences de la Terre et de l’Univers, UFR des Sciences et Techniques, Université de Bretagne Occidentale (UBO), 6 Avenue Le Gorgeu—CS 93837, 29238 Brest Cedex 3, France e-mail: [email protected] D. Dias-Brito · P. Tibana Departamento de Geologia Aplicada, UNESPetro—Universidade Estadual Paulista—Rio Claro, IGCE, Caixa-Postal 178, av. 24 A 1515, Bela Vista, 13506-900 Rio Claro, SP, Brazil I. I. Bucur Department of Geology, Babeo-Bolyai University, Str. M. Kogalniceanu nr.1, 400084 Cluj-Napoca, Romania Thyrsoporellaceae nov. fam. · Choristosporate · Goniosporate · Cladosporate · Syringosporate · Cretaceous Introduction Masse (1995) described Holosporella nkossaensis, a new species from the Albian of Congo. This species is also found in strata of almost the same age on the opposite western coast of the South Atlantic Ocean in Brazil (Granier et al. 2008). Based on material from the Santonian of Hungary discovered in the early 1970s (Szantner et al. 1969; Sidó 1974), Conrad et al. (2002) introduced two new taxa, the genus Hungariporella and its type-species H. baconica. These authors drew a Wrst comparison of both species—the Congolese and the Hungarian—and came to the conclusion that they were dealing with discrete species, which should also be ascribed to discrete genera. Further to a presentation given by one of us (IIB) at the 10th International Symposium on Fossil Algae (Dragastan et al. 2011) Conrad (pers. comm., 16/09/2011) suggested that the unpublished material presented there has striking similarities with both Hungariporella baconica and Holosporella nkossaensis, which justiWes having here a discussion section dedicated to this speciWc issue. However, it appears that this new material originating from Upper Cretaceous strata of Morocco corresponds to a third discrete form (that will be described in a separate forthcoming paper). Actually, the main (and possibly unique) similarity could be the fact that in all cases calciWcation was rather weak, probably mostly because it was extracellular sensu stricto (Granier 2012): the extracellular sensu lato category, which is typical for the ‘calcareous green algae’ (CGA), comprises another sub-category, i.e., the “intercel- 123 Author's personal copy Facies Fig. 1 Map of the Brazilian localities lular” type known from taxa in which the distal part of the laterals forms a cortex. In most CGA, precipitation of aragonite needles takes place in the mucilage (see the recent review paper by Granier 2012), but in at least two of the studied species it did not occur behind a cortical protection as it is, for instance, the case in Bornetella, Cymopolia, or Neomeris. The needle network was then denser along the cell wall, i.e., the main axis and the laterals, than it was in the remaining space between these laterals. Later micritization (Bathurst 1966) altered and masked the original pattern. As a consequence of this poor preservation, also documented from Upper Albian-Cenomanian strata in Brazil by Granier et al. (2008), the algal body plan is subject to interpretation. 123 Specimens from Congo (“Holosporella”) According to Masse (1995: excerpt from the original diagnosis): “the arrangement of: ‘closely packed verticils of fertile spherical to subspherical ampullae’ around an axial central siphon and the occurrence of a peduncle of length lower or equal to those of the related ampulla allow us to ascribe the species to the genus Holosporella Pia, 1930. (…) Our material proves the variability of the characteristic within the same species: peduncles are sometimes shorter (…), of equal length (…) and very rarely longer (…) than the ampulla.” To summarize: due to the poor preservation of most specimens, the author could not observe more than one segment within the laterals. However, branching, i.e., Author's personal copy Facies Fig. 2 The early marine Wbrous cement (arrows) around a micritized algal bioclast precedes a later micritic inWlling of part of the formerly openpore space. Sample 490, 648-m depth, Petrobras Well 2-AD-1, State of Maranhão, Brazil the occurrence of secondary segments or ampullae, might have been detected on his pl.2, Wg. 8, bottom left corner or in his pl. 2, Wg. 12, left side (Masse 1995). In that paper (Masse 1995), representatives of the genus Holosporella are erroneously assigned to the Family Dasycladaceae though they should have been referred to the Family Triploporellaceae. Specimens from Hungary (Hungariporella) According to Conrad et al. (2002: excerpt from the original diagnosis of their species): “Thallus simple, cylindrical, here and there swelled, with a cylindrical main axis developing whorls of up to three orders of laterals. Primary segments of laterals horizontal or slightly tilted, Wrst quite large, forming a holdfast, then either globose, or elongated, bulgy, swollen at the tip, indicative of containing one or two conspicuous gametangia. Secondary segments of laterals, whenever present, around four in number, dustered at the rip of the primaries, clavate with a proximal peduncle. Slender third-order twigs may arise from the tip of the secondaries.” In their Wgure legend of the holotype section (Conrad et al. 2002, pl. II, Wg. 1, also illustrated in their text-Wg. 3, specimen 17), the authors describe laterals consisting of an “elongated primary segment” with “two conspicuous bulges” (looking like a dumbbell, i.e., two vesicles connected by a medial narrowing), vesicular secondary segments, and “slender” tertiary segments. However due to the obliquity of the section itself and because there are many examples in “calcareous green algae” where the lack of calciWcation at the contact surface of adjacent laterals (in Dasycladales) or Wlaments (in Bryopsidales) led to interconnections of the pores left within the calcareous coatings after the decay of the algal organic material, one can question the interpretation of these authors. 123 Author's personal copy Facies Fig. 3 “Outcrops” of the Riachuelo Formation, UPFA SE-11, sample 18, Fazenda Massapé, 25 km NW of Aracaju, State of Sergipe. a Large piece of pisolitic limestone; b Detail of the pisoids Xoating in an oolitic matrix New material from Brazil Barreirinhas There are few discontinuous outcrops with shallow-water marine Cretaceous carbonates in Brazil. They occur in some sedimentary basins located on the eastern South Atlantic continental margin of Brazil (e.g., the Potiguar Basin or the Sergipe Basin). They give us a foretaste of the thick successions—attaining locally hundreds of meters— which are identiWed in the subsurface thanks to borehole and seismic data (e.g., the Barreirinhas, Sergipe or Potiguar basins). Granier et al. (2008) Wrst reported the occurrence of Holosporella nkossaensis from core samples of an oVshore borehole which cut through strata of the Ponta do Mel Formation (Upper Albian—Cenomanian) in the Potiguar Basin (Fig. 1). They stated, however, that “as in the type-specimens of this poorly known form, calciWcation in our material is rather weak and the structures are in part masked by micritization”; therefore this material was not favorable for a reinterpretation. New material was collected from two basins on the eastern South Atlantic margin of Brazil (Fig. 1): Barreirinhas and Sergipe. These new specimens allow a better interpretation of the body plan features. Locality: Petrobras Well 2-AD-1, 200 km East of São Luís, State of Maranhão, SAD 69 (Brazil): UTM 23S 829578.70E 9694525.22N as indicated by ANP-BDEP (latitude 2° 45’ 31.984” S, longitude 42° 2’ 9.855” W; see Google Maps: –2.758885 –42.036072). 123 Stratigraphic level: nian in age. BonWm Formation, Albian-Cenoma- Material: Three core samples labeled CB-490 (at 648 m depth), CB-505 (at 652-m depth), and CB-520 (at 674-m depth). Facies, assemblage, and paleoenvironment: Microfacies are poorly sorted oolitic and bioclastic pack- to grainstones. Ooids are of the superWcially coated type; their diameters vary with that of their nuclei, commonly a micritized grain, from 200 to 500 m. Nubeculariid foraminifers contribute to grain encrustation. Larger grains consisting mostly of gastropods, mollusc shells, and “solenoporacean” nodules (i.e., Elianella, Marinella, and Pycnoporidium) Xoat in the grainy matrix. Among the remaining bioclasts there are a number of broken thalli of the alga studied. Author's personal copy Facies Fig. 4 Oblique section of Brasiliporella nkossaensis, emend., nov. comb. Sample 505, 652 m depth, Petrobras Well 2-AD-1, State of Maranhão, Brazil. d: ?dehiscence, MA main axis, R1 primary segment of a lateral, R2 gametophores In sample CB-490, the occurrence of micritic inWllings following an early phase of cementation (thin Wbrous sparitic calcite) provides evidence for a hardground (Fig. 2). Sergipe dense vegetation, only a few large pieces of rock crop out here and there (Fig. 3). Therefore it is not possible to elaborate a detailed lithological succession. Stratigraphic level: Material: Locality: Fazenda Massapé (top of a hill), 25 km NW of Aracaju, State of Sergipe, SAD 69 (Brazil): UTM 24S 0700470E 8810980N (latitude 10° 44’ 47.28⬙ S, longitude 37° 10⬘ 0.39⬙ W; see Google Maps: –10.746466 –37.166774). Due to local climate, soil development, and Riachuelo Formation, Albian in age. Almost 20 samples labeled UPFASE-11. Facies, assemblage, and paleoenvironment: Microfacies are oolitic, slightly bioclastic, locally pelletoidal (averaging 75 m in diameter), grainstones. Ooids are well-”sorted” (because this wording is charged with connotations, the use 123 Author's personal copy Facies 123 Author's personal copy Facies 䉳 Fig. 5 Brasiliporella nkossaensis, emend., nov. comb. All specimens from sample 505, 652 m depth, Petrobras Well 2-AD-1, State of Maranhão, Brazil. Scale bar is 500 m (a–f), 200 m (g–h). a Shallow tangential section illustrating the proximal part of the gametophores, arranged in pairs in each lateral. b Deep oblique tangential section. The primary segments of the laterals are arranged in quincunxes (see the cross pattern visible in the lower third of the picture). c Deep oblique tangential section. The primary segments of the laterals are arranged in quincunxes, i.e., they alternate from one row to the one above and the one below. The proximal peduncular parts of some gametophores are visible here on the side (actually on the top) of some primary segments (center of the picture). Detail of Fig. 7a. d Deep tangential section of a curved portion of thallus. e Very shallow tangential section cutting through the gametophores. f Oblique sections, a low-angle (sub-transverse) one and a higher-angle (sub-axial) one. g Oblique (sub-transverse) section illustrating the verticillate arrangement of the laterals. Detail of f. h Tangential section at the level of the gametophores. Detail of e of “calibrated” which is neutral might be more appropriated), averaging 200 m in diameter. They are of radial to concentric types and include few hemi-ooids. Their nuclei may consist of sand-sized quartz and feldspar grains. Another typical facies is represented by Xoatstones of coated grains (commonly pisoids) and grain aggregates with the oolitic grainstone matrix. Bioclasts are mostly thalli of the alga studied, some balanid plates, and a few echinoderm remains, and small agglutinated foraminifers (? textulariids or ? trochamminids). Occurrence of both radial ooids and hemiooids as well as of rather long and very little fragmented cylinders, which represent the weak external calciWcation of our fragile algal thalli, plead for a rather low-energy environment (though the texture is grain-supported). Calcareous green algae (more precisely the occurrence of the Dasycladales, see Granier 2012: Fig. 2) suggest the shallow euphotic zone. In addition, some features, such as the very low phycological and zoological diversity, argue for brackish waters. Description and discussion of the new material Description The thallus is cylindrical, more or less straight, and its calcareous coating forms a continuous tubular structure, often broken into shorter fragments after the decay of the alga. In the Sergipe samples the longest tube measures 5.8 mm in length. External diameters (D) range from 275 to 825 m (450 on average), internal diameters (d) range from 75 to 225 m (125 on average); the ratio d/D varies from 18 to 42 % (27 on average). In the Barreirinhas samples, external diameters (D) range from 375 to 925 m (625 on average), internal diameters (d) from 100 to 325 m (175 on average); the ratio d/D varies from 14 to 42 % (27 on average). These measurements are matching those given for Holosporella nkossaensis by Masse (1995): external diameters (D) range from 333 to 887 m (512 on average) and internal diameters (d) from 82 to 161 m (113 on average). In contrast, we found huge discrepancies when comparing these values with those given by Conrad et al. (2002) for Hungariporella baconica: external diameters (D) range from 762 to 3428 m and internal diameters (d) from 333 to 1286 m, the ratio d/D varying from 20 to 59 %. The same discrepancies can be found with the yet undescribed Moroccan form with D ranging from 1.25 to 2.10 mm (1.57 on average), d from 0.32 to 0.80 mm (0.54 on average), and the ratio d/D varying from 19 to 44 % (34 on average). Objectively, on the sole basis of these size parameters, there are three discrete species. There are also three discrete body plans, as it is documented hereafter, and consequently three discrete genera. Because we can hardly compare homologous portions of the algae we do not take into consideration other measurements for comparisons. The Barreirinhas samples provide the best specimens to understand the algal structure. The cylindrical main axis bears at regular intervals (up to 100 m in h = spacing + thickness) verticils of 8 (Fig. 6f) to ?10 laterals alternating from one row to the next (Fig. 5a–d). Each lateral consists of a primary segment, thinning from the proximal part to the medial part before bulging in its distal part (it is suggested that it is homologous to the primary segment of a cladosporate form), bearing two rounded vesicles—interpreted as gametophores (up to 85 m in diameter)—in terminal position, which corresponds to a choristosporate form (Figs. 4, 5g, 6c, g, j, 7d, f–g). The bulge is compressed vertically, a pattern which is visible from some deep tangential sections (Fig. 5b–d). In some oblique or axial sections the bulge (R1) is sometimes misinterpreted as a gametophore (R2F) (Figs. 2, 5e, 6c and material illustrated by Masse (1995), pl. 2, Wgs. 1–2, 6–8, 11). The pairs of gametophores are lying in the whorl plane deWned by their respective laterals, a pattern which is visible from transverse or low-angle sub-transverse sections (Figs. 4, 5a, c, 6f, j, 7d, g), as well as in some deep tangential sections (Fig. 5a, c). There is no evidence for secondary sterile segments of laterals, nor for tertiary segments as reported for Hungariporella baconica by Conrad et al. (2002). But there are very few breaks in the gametophores (Fig. 4) that might be marks of dehiscence preceding the release of the gametes. Discussion The combination of supposedly cladosporate and choristosporate traits found in some representatives of the Dasycladaceae, e.g., in Genotella pfenderae, Cymopolia satyavanti, 123 Author's personal copy Facies 123 Author's personal copy Facies 䉳 Fig. 6 Brasiliporella nkossaensis, emend., nov. comb. All specimens from sample 505, 652 m depth, Petrobras Well 2-AD-1, State of Maranhão, Brazil. Scale bar is 500 m (a–f, h–i), 200 m (g, j). a Tangential section, partly encrusted by Marinella lugeoni (upper left hand side). b Oblique section with micritic intercellular inWll. c Oblique section with drusy sparitic intercellular inWll. d Transverse section with micritic intercellular inWll, partly encrusted by Nubeculariid foraminifers. e Oblique section with drusy sparitic intercellular inWll. f Transverse section with a micritized verticil composed of eight laterals. h Sub-transverse section with drusy sparitic intercellular inWll. Detail of Fig. 7b. h Oblique section with drusy sparitic intercellular inWll. i The microfacies is a packstone of superWcial ooids and bioclasts. j Sub-transverse section with drusy sparitic intercellular inWll. Detail of i and Cymopolia tibetica, can be considered either as a transitional stage from cladospory to choristospory, or eventually as a teratologic (atavistic) case, but never as a character to be used in the systematics at the genus level. Cymopolia satyavanti, still treated by some authors as Indopolia satyavanti, and related forms are probably closely related to Holosporella nkossaensis because they have two gametophores per lateral but they also have a number of secondary sterile segments that are lacking in the studied alga. Choristospory justiWes its ascription to the Family Dasycladaceae. We assume that the number of gametophores, i.e., a pair per lateral, is not an anomaly (as in Indopolia) but a generic character. Accordingly we ascribe it to the Tribe Batophoreae, which was up to date restricted to a single modern genus: consequently the tribe would date back to the Albian (mid-Cretaceous) times. Systematics Phylum Chlorophyta Class Dasycladophyceae Hoek et al., 1995 Order Dasycladales Pascher, 1931 Family Dasycladaceae (Kützing, 1841), emend. Tribe Batophoreae Valet, 1969 Genus Brasiliporella nov. gen. Origin of the name: From the country name, Brasil, which is linked with the term Terra Brasilis utilized in Portuguese documents and maps from the beginning of the XVIth century. The term has traditionally been associated with the red wood of Caesalpinia echinata, popularly known in Portuguese as pau-brasil, which was very abundant in coastal areas and the basis of the Wrst economic activities in colonial Brazil. Type species: Holosporella nkosaensis P. Masse, 1995. Diagnosis: Cylindrical thallus with simple verticils (euspondyle) arranged in more or less regular alternation. Fertile laterals consisting of a primary segment bearing two vesicular ampullae in terminal position (choristosporate). No record of any secondary sterile segment of laterals. Brasiliporella nkossaensis P. Masse, 1995 emend. nov. comb. (Figs. 2, 4–7) 1995 2008 Holosporella nkossaensis n.sp.—P. Masse: 304, pl. 2, Wgs. 1–12 (holotype: pl. 2, Wg. 6) Holosporella nkossaensis ¡ Granier et al.: 312, pl. 2, Wg. F Type material: The holotype consists of an oblique section from a thin-section cut in a core sample at 3479.63 m, well N’Kossa NKF 202 (oVshore Congo), housed in the collection of TOTAL Exploration & Production, CSTJF (Centre ScientiWque et Technique Jean Féger), Pau, France (formerly “Elf Aquitaine Coll.”). Emended diagnosis: Representative of the genus Brasiliporella characterized by the typical pattern of its laterals, more speciWcally its distally inXated primary segments of laterals and the arrangement of the pairs of gametophores that are lying in the whorl planes (Fig. 8), as well as by its general measurements (listed below). Measurements: (L: maximum length; D: external diameter; d: internal diameter; l⬘: length of the primary segment of the laterals; l⬙: length of the secondary segment of the laterals; w: number of laterals per verticil; h: height of a “whorl” (verticil) plus one interverticillar spacing; p⬘: width of the primary segment of the laterals; p⬙: width of the secondary segment of the laterals) L = 5.8 mm; D = »275 (min) to »925 (max) m; d = »75 (min) to »325 (max) m; e = (D–d)/2 = l⬘ + l⬙ = »100 (min) to »325 (max) m; d/D = 14 % (min) to 42 % (max); w = 8 to ?10; h = »75 (min) to »95 (max) m; l⬘ up to 160 m; p⬘ narrowing down to »20 m; p⬘ bulging up to »105 m; l⬙ (peduncle + gametophore) up to 140 m; p⬙ (gametophore) up to »85 m. ClassiWcation of the Dasycladales (by B. Granier & I.I. Bucur) The monograph of Berger and Kaever (1992) gives an overview of the Order Dasycladales which comprises two families with modern representatives: (1) the Dasycladaceae and (2) the Polyphysaceae (formerly known as Acetabulariaceae), as well as three families which are known only from the fossil record: (3) the Seletonellaceae, a fam- 123 Author's personal copy Facies 123 Author's personal copy Facies 䉳 Fig. 7 Brasiliporella nkossaensis, emend., nov. comb. Specimens from sample 505, 652 m depth, Petrobras Well 2-AD-1, State of Maranhão, Brazil, except for specimen in d, which is from sample 520, 674 m depth, same locality. Scale bar is 500 m (a–e), 200 m (f–g). a–c, e The microfacies is a packstone of superWcial ooids and bioclasts (mollusc shells and calcareous algae). a Deep oblique tangential section with drusy sparitic intercellular inWll on the left-hand side (for detail see Fig. 5c) and oblique section of a micritized specimen on the right-hand side. b Sub-transverse sections (for details see c and Fig. 6g). c Oblique section (for detail see f). d Oblique section of a micritized specimen. e Oblique section (for detail see h). f Oblique section. Detail of c. g Sub-transverse section. Detail of b. h Oblique section. Detail of e ily that includes all aspondyle (i.e., non verticillated) forms, (4) the Triploporellaceae, and (5) the Diploporaceae. The Family Dasycladaceae is, in turn, subdivided into four tribes: (1) Tribe Bornetelleae, which comprises the genera Bornetella, Dactylopora, Digitella, and Zittelina, and (2) Tribe Parkerelleae with the genera Carpenterella, Jodotella, and Parkerella. The gametophores are in lateral position on the primary segment of fertile laterals, i.e., of the goniosporate type (Granier 2010). The Wrst goniosporate was possibly Granieria iberica or Uragiella ? liasica, which date back to the Liassic (Barattolo and Parente 2000). (3) Tribe Dasycladaceae: Fertile laterals have one gametophore (exceptionally two) in terminal position on their primary segment (choristosporate type). The number of secondary segments of laterals varies from one genus to the other: – one in the fossil Montiella, – two in Neomeris (exceptionally three, e.g., in the living Neomeris stipitata) and in the fossil Genotella, – more than three in Cymopolia. Tertiary segments are known in both Dasycladus and Chlorocladus, quaternary segments in Chlorocladus only. Afghanopolia and Indopolia are considered as junior synonyms of Cymopolia (Génot 1978, 1980 and RadoibiT 1998, respectively). Neomeris dates back to the Valanginian but the Wrst choristosporate could be Eodasycladus from Liassic times (Barattolo et al. 1995, 2012), as in the case of Granieria above. (4) The Tribe Batophoreae comprised only one genus, Batophora. Fertile laterals have two or more gametophores in terminal position on primary and secondary—occasionally tertiary—segments (choristosporate type). The sequence of segments continues with tertiary, quaternary, quinary, senary, and even septenary segments. With Brasiliporella, the tribe now includes a new—fossil—representative. (Remark: The genus Archaeobatophora of the Fig. 8 Oblique section (above) and short (two whorls only) axial section (below) of Brasiliporella nkossaensis, emend., nov. comb. R1: primary segment of the laterals; R2F: secondary (fertile) segments of the laterals, i.e., a gametophore plus its proximal peduncle; D: external diameter; d: internal diameter; l’: length of R1; l’’: length of R2F; h: height of a “whorl” (verticil) plus one interverticillar spacing Ordovician is “all but” a member of the Dasycladaceae because it lacks gametophores.) As the acquisition of reproductive traits such as goniospory and choristospory, as well as syringospory, is irreversible, we think that these features can be used to subdivide the Order Dasycladales (Fig. 9) into the following families: – Family Bornetellaceae nov. fam. (basionym Bornetella, type species: B. nitida), which comprises all goniosporate forms; – Family Thyrsoporellaceae nov. fam. (basionym Thyrsoporella, type species: T. cancellata) with syringosporate forms only. Accordingly, the families Dasycladaceae and Triploporellaceae are emended to exclude the taxa now included in the new families: – Family Dasycladaceae emend. (basionym Dasycladus, type species: D. vermicularis), restricted to choristosporate forms, and 123 Author's personal copy Facies Fig. 9 Comparison of Berger and Kaever’ classiWcation with the new classiWcation introduced herein, based on the reproductive features of the sole euspondyle forms (after Granier in Mathieu et al. 2011, and Granier 2011, modiWed). The table does not take into consideration aspondyle forms, i.e., the Family Seletonellaceae, nor metaspondyle forms, i.e., the Family Diploporaceae – Family Triploporellaceae emend. (basionym Triploporella, type species: T. fraasi) with endosporate and cladosporate forms. The emended classiWcation of the Dasycladales proposed herein comprises seven families, three with both modern and fossil representatives (Bornetellaceae, Dasycladaceae, and Polyphysaceae) and four with fossil representatives only (Diploporaceae, Seletonellaceae, Thyrsoporellaceae, and Triploporellaceae). Two of the families discussed here (i.e., Bornetellaceae—goniosporate—and Dasycladaceae— choristosporate—) possibly originated in Jurassic times; the oldest representative of the third family (i.e., Thyrsoporellaceae) could be Placklesia from the Rhaetian (Late Triassic). This last genus was considered by Schlagintweit (in Gawlick et al. 2006) as a junior synonym of Thyrsoporella, a position that we do not agree with (because both genera have discrete division formula for their branching patterns). Acknowledgments This paper is a contribution to the special volume of Facies dedicated to the “10th International Symposium on Fossil Algae” held in Cluj-Napoca (Romania, September 12–18, 2011). 123 The material from the Sergipe Basin was collected with the support of ANP-PRH 05. Thin-sections from the Barreirinhas Basin were initially studied by Carozzi et al. (1973) before the creation of the ANP (Brazilian Petroleum National Agency). This research was supported by the “Carbonatos do Brasil Project” linked to the Brazilian Sedimentology/ Stratigraphy Net sponsored by Petrobras. We thank Petrobras, ANP and FUNDUNESP (Fundação para o Desenvolvimento da Universidade Estadual Paulista “Julio de Mesquita Filho”, Rio Claro, São Paulo, Brazil). Thanks go also László Bujtor, Marc Conrad, Olga Piros, and Rajka RadoibiT for information they provided regarding the type material of Hungariporella baconica. Finally, we are grateful to Rajka RadoibiT, Franz Fürsich, and Felix Schlagintweit for reading the original manuscript and making useful suggestions. Appendix List of taxa cited in the text Acetabulariaceae Hauck, 1885 Afghanopolia Kaever, 1969 Archaeobatophora Nitecki, 1976 Batophora J. Agardh, 1854 Batophoreae Valet, 1969 Author's personal copy Facies Bornetella Munier-Chalmas, 1877 Bornetella nitida (Harvey, 1857) Bornetellaceae nov. fam. Bornetelleae (L. & J. Morellet, 1913) Brasiliporella nov. gen. Brasiliporella nkossaensis P. Masse, 1995, emend., nov. comb. Bryopsidales SchaVner, 1922 Carpenterella (Munier-Chalmas ex L. & J. Morellet, 1922) Caesalpinia echinata Lamarck 1789 Chlorocladus Sonder, 1871 Cymopolia Lamouroux, 1816 Cymopolia satyavanti (Pia in Pia & L.R. Rao, 1936) Cymopolia tibetica L. Morellet in H. Douvillé, 1916 Dactylopora Lamarck, 1816 Dasycladaceae (Kützing, 1841), emend. Dasycladales Pascher, 1931 Dasycladaceae Pia, 1920 Dasycladus C. Agardh, 1828 Dasycladus vermicularis (Scopoli 1772) Digitella L. & J. Morellet, 1913 Diploporaceae (Pia, 1920) Elianella Pfender & Basse, 1948 Eodasycladus Cros & Lemoine in Granier & DeloVre, 1993 (non 1966) Genotella (Granier et al., 1991) Genotella pfenderae (Konishi & Epis, 1962) Granieria Barattolo & Romano in Barattolo et al. 2008 Granieria iberica (Dragastan & Trappe, 1986) Holosporella nkossaensis P. Masse, 1995 Holosporella Pia, 1930 Hungariporella Conrad et al., 2002, emend. Hungariporella baconica Conrad et al., 2002 Indopolia Pia, 1936 Indopolia satyavanti Pia in Pia & L.R. Rao, 1936 Jodotella L. & J. Morellet, 1913 Marinella Pfender, 1939 Marinella lugeoni Pfender, 1939 Montiella Munier-Chalmas ex L. & J. Morellet, 1922 Morelletpora Varma, 1950 Neomeris Lamouroux, 1816 Neomeris stipitata Howe, 1909 Parkerella Munier-Chalmas ex L. & J. Morellet, 1922 Parkerelleae (L. & J. Morellet, 1922) Placklesia Bilgütay, 1968 Polyphysaceae (Kützing, 1841) Pycnoporidium Yabe & Toyama, 1928 Seletonellaceae (Korde, 1950) Thyrsoporella Gümbel, 1872 Thyrsoporella cancellata Gümbel, 1872 Thyrsoporellaceae nov. fam. Triploporellaceae (Pia, 1920), emend. Triploporella (Steinmann, 1880) Triploporella fraasi (Steinmann, 1880) Uragiella ? liasica (Lebouché & M. Lemoine, 1963) Zittelina Munier-Chalmas ex L. & J. Morellet, 1913 References Barattolo F, Parente M (2000) New observation on Uragiella liasica (Lebouché & Lemoine) (Green Algae, Dasycladales) from the type-locality (Canders, Languedoc-Southern France). Boll Soc Paleont It 39:123–137 Barattolo F, De Castro P, Parente M (1995) Some remarks on the genera Palaeodasycladus (Pia, 1920) Pia, 1927 and Eodasycladus Cros & Lemoine, 1966 ex Granier & DeloVre, 1993 (Green Algae, Dasycladales). In: Piller WE (ed) Proceedings of the International Symposium and Field-Meeting “Alpine Algae ‘93”. Beitr Paläontol 19:1–11 Barattolo F, Shirazi MPN, Vecchio E (2012) The genus Eodasycladus (Lower Jurassic dasycladalean green alga) and its relationship with Palaeodasycladus. Facies: p 11 (Online First: 11 April 2012). doi:10.1007/s10347-012-0304-6 Bathurst RCG (1966) Boring algae, micrite envelopes and lithiWcation of molluscan biosparites. Geol J 5:15–32 Berger S, Kaever MJ (1992) Dasycladales. An illustrated monograph of a fascinating algal order. Georg Thieme Verlag, Stuttgart, p 247 Carozzi AV, Tibana P, Tessari E (1973) Estudo das microfacies da Formação BonWm (Cenomaniano) da bacia de Barreirinhas, Brasil. Ciência técnica petróleo: Seção Exploração de Petróleo, Petrobras, Rio de Janeiro 6: p 86 Conrad MA, Bodrogi I, RadoibiT R (2002) Hungariporella baconica n gen, n sp (Dasycladales, calcareous green algae) from the Santonian of the Southern Bakony Mountains (Transdanubia, Hungary). 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