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
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International Journal of Paleontology,
Sedimentology and Geology
ISSN 0172-9179
Facies
DOI 10.1007/s10347-012-0312-6
1 23
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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-
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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.
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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.,
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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.
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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).
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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.
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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
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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,
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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-
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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
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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
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