A new mid-Cretaceous Neomeris - Université de Bretagne Occidentale



A new mid-Cretaceous Neomeris - Université de Bretagne Occidentale
A new mid-Cretaceous Neomeris
(dasycladacean alga) from the Potiguar
Basin, Brazil
Bruno Granier, Dimas Dias-Brito & Ioan
I. Bucur
International Journal of Paleontology,
Sedimentology and Geology
ISSN 0172-9179
DOI 10.1007/s10347-012-0322-4
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DOI 10.1007/s10347-012-0322-4
A new mid-Cretaceous Neomeris (dasycladacean alga)
from the Potiguar Basin, Brazil
Bruno Granier • Dimas Dias-Brito
Ioan I. Bucur
Received: 6 July 2012 / Accepted: 16 July 2012
Ó Springer-Verlag 2012
Abstract Neomeris (Lamouroux, 1816) is an extant
taxon, the origin of which can be tracked back into Early
Cretaceous times. The introduction of a new mid-Cretaceous species from Brazil, i.e., Neomeris srivastavai n. sp.,
offers the opportunity to review the subdivision of the
genus into three subgenera, to complete the catalogue of
the fossil calcareous algae of Brazil, and to point out the
huge stratigraphic gap and lack of documentation between
the first occurrence of the dasycladacean model of reproduction, i.e., choristospory, and the oldest record so far
known of an undescribed fossil Neomeris (from Portugal).
Keywords Dasycladales Dasycladaceae Neomeris Choristosporate Cretaceous
Many dasycladalean genera are monospecific or comprise
only a few species. In comparison, with more than 50
species over almost 120 myr, i.e., ranging from the Early
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
Departamento de Geologia Aplicada, UNESPetro,
Universidade Estadual Paulista, Rio Claro, IGCE, Caixa-Postal
178, av. 24 A 1515, Bela Vista, Rio Claro, SP 13506-900, Brazil
I. I. Bucur
Department of Geology, Babeş-Bolyai University,
Str. M. Kogalniceanu nr.1, 400084 Cluj-Napoca, Romania
Cretaceous times up to the present day, Neomeris
(Lamouroux, 1816) has a relatively stable and successful
body plan. This paper deals with the description of a new
fossil species, an early representative found in Cenomanian
strata of Brazil.
A summary review of the genus Neomeris (Lamouroux,
1816) and its representatives
Originally described as a coral-like animal by Lamouroux,
(1816, p. 241–243), the living Neomeris dumetosa was
later correctly assigned to the algae by Decaisne (1842,
p. 142). The generic name itself, after that of one of the
Nereids in the Greek mythology, has known some vicissitudes; this epithet was also given to polychaetes (Costa
1844) and even to a dolphin (Gray 1846). The main feature
of this algal genus is a main stalk bearing regularly spaced
verticils of laterals with two (exceptionally three, e.g., in
the living N. stipitata Howe, 1909) secondary sterile segments and, where they are fertile, one gametophore in
terminal position (choristosporate type) per primary segment. Aragonite precipitation on the outer side of the
cytoplasmic envelope of these large unicellular benthic
algae may form calcareous coatings that may be ‘‘preserved’’ in the sediments: the calcification process is
mostly intercellular, i.e., extracellular sensu lato (Granier
2012), as it occurs in the open space between the stalk and
the cortical layer formed by the juxtaposed distal parts of
the laterals. The finds of such remains (‘‘tubes’’ or isolated
‘‘rings’’) in the fossil record led to the introduction of
numerous synonyms and eventually some subgenera:
Larvaria Defrance, 1822, Vaginopora Defrance in Bronn,
1825, Haploporella Gümbel, 1872, Herouvalina MunierChalmas ex Steinmann, 1899, Lemoinella L & J Morellet,
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1913, Meminella L & J Morellet, 1913, Sakkionella
Segonzac, 1976, non 1970, and Drimella Radoičić, 1984.
In addition to the type species, Neomeris dumetosa
(Lamouroux, 1816), there are six extant species living in
modern tropical and subtropical seas (Berger and Kaever
1992; Berger 2006), at least 37 Cenozoic species according
to Deloffre and Granier (1992), and a small number of
Cretaceous forms.
Most Cenozoic specimens were collected in loose calcareous sand, which makes it easy to sort individual calcareous coatings and to understand the structure of these
algae. This work was even eased by the use of scanning
electron microscopy (SEM) since the 1980s (Génot 1980,
1985, 1987, 2009; Deloffre and Génot 1982; Génot and
Granier 2011). Last but not least, some specimens may
‘‘have retained their original aragonite’’ (Génot 1985). On
the contrary, the remaining specimens and Cretaceous ones
are found embedded in limestones, commonly ‘‘recrystallized’’ (Granier 2012), which required thin-sections, either
random or oriented, to reconstruct the algal structures
resulting in lesser morphological detail. The catalogue of
Jurassic and Cretaceous algae published by Bassoullet et al.
(1978) lists four Cretaceous representatives: N. budaense
Johnson, 1968, N. cretacea Steinmann, 1899, N. occidentalis (Johnson and Kaska, 1965), and N. pfenderae Konishi
and Epis, 1962. In their inventory, Granier and Deloffre
(1993) added three species to the list: N. circularis Badve
and Nayak 1984, N. drimi Radoičić, 1984, and N. jerinae
Radoičić, 1984; but they also removed three: N. pfenderae,
which was transferred to the genus Genotella (Granier et al.
1991), as well as Larvaria occidentalis and Neomeris
budaense, which are both considered nomina nuda
[Granier et al. (1994) quote N. budaense in their synonymy
list for Heteroporella lepina (Praturlon, 1967, non 1966)].
Fig. 1 Location map of the Petrobras well RNS-11
Fig. 2 All photomicrographs from sample 1,638-m depth, Petrobras c
well 1-RNS-11, State of Rio Grande do Norte, Brazil. Scale bar is
1 mm (a), 250 lm (b–e). a The microfacies is a floatstone of large
aggregates (ag) and ‘‘solenoporacean’’ nodules (n) with a pelletoidalooidal grainstone matrix. Geopetal (?) internal sediment is visible (m).
b, d–e Neomeris srivastavai nov. sp. b, d Holotype: Oblique section
of a rather highly calcified portion of a thallus. Both the axial stem
and the primary segments of the laterals are calcified. b Natural light.
d Contoured black and white negative image. e Random section
cutting several verticils. c Thin-walled hyaline (? planktonic)
foraminifers in the geopetal infilling of the primary intergranular
Further to the revision of Acroporella assurbanipali Elliott
1968 by Barattolo and Romano (2001) this fifth species is
added to the list. Finally, Radoičić and Schlagintweit (2007)
described one species: N. mokragorensis, while Radoičić
(2002) introduced two new more: Neomeris conradi and
N. decapoaae. Sokač (2004) considered these last two
species as junior synonyms of N. cretacea Steinmann, 1899,
a view that we cannot support because Steinmann’s species
is ascribed to the subgenus Neomeris whereas both
Radoičić’s species belong to the subgenus Drimella. To
summarize, to date, eight Cretaceous species are valid, the
subgeneric assignment of which is given in Table 1.
Material and geological information
The material comes from a Petrobras exploration well
(1-RNS-11) drilled through the Ponta do Mel Formation in
the Potiguar Basin, offshore of Macau, Rio Grande do
Norte (Fig. 1).
Material The material studied consists of two acetate
peels and five thin-sections cut in a core sample at 1,638-m
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Locality Petrobras well 1-RNS-11, 30 km North of
Macau, State of Rio Grande do Norte, SAD 69 (Brazil):
UTM 24S 776981.92E 9461584.57 N as indicated by ANPBDEP (latitude 4°510 59.19800 S, longitude 36°300 9.65500 W;
see Google Maps: -4.866444, -36.502682).
Stratigraphic level The specimens are from the uppermost part of the Ponta do Mel Formation, (? Late Albian-)
Cenomanian in age (Tibana and Terra 1981; Granier et al.
Facies, assemblage, and paleoenvironment: The
microfacies (Fig. 2a) displays a grain-dominated fabric; it
is a floatstone of large aggregates and ‘‘solenoporacean’’
nodules (with Marinella and Pycnoporidium) with a
poorly sorted grainstone matrix. The smaller grains are
pelletoids, followed by micritic ooids that are slightly
larger. Bioclasts are few, partly micritized and commonly
microbially coated: echinoderms, small gastropods, mollusc shells, few benthic foraminifers, including Trocholina
sp., and calcareous green algal remains (the new
Neomeris, a Clypeina sp., and a Linoporella ? sp., see
Granier et al. 2008, pls. 1–2). The intergranular space is
filled by calcitic cements, mostly drusy calcite and some
syntaxial cement surrounding echinoderm remains. There
are also local patches consisting of micrite and small
grains, which are geopetally arranged. Among these
grains, we identified thin-walled hyaline (? planktonic)
foraminifers (Fig. 2c). Part of this material possibly percolated through the grainy column from the discontinuity
(transgressive surface) sited a few meters above and
which marks the boundary between the shallow-water,
moderate-energy facies of the Ponta do Mel Formation
and the pelagic facies of the Jandaira Formation (Granier
et al. 2008).
Table 1 Subgeneric
assignment of the Cretaceous
species of Neomeris
Fig. 3 Neomeris srivastavai nov. sp. All specimens from sample c
1,638.0 m depth, Petrobras well 1-RNS-11, State of Rio Grande do
Norte, Brazil. Scale bar is 250 lm (a–f). Thin-sections (a, c, e–f) and
acetate peels (b, d; made prior to the thin-sectioning). a–b Paratype:
Transverse sections cut some tens of micrometers apart. c–d Subtransverse sections cut some tens of micrometers apart. The axial
stem is not calcified. e–f Oblique sections cutting several successive
Elements of comparison and description
The discussion on the subgeneric assignment of the new
species can be abridged because it was already addressed
by Granier et al. (2008, p. 313, pl. 1, fig. A). The
arrangement of the pair of secondary segments with respect
to the ampulla, that is they are presently sited on both side
of the ampulla, is characteristic of the subgenus Neomeris
(Table 1). Accordingly comparisons can only be made with
two Cretaceous species: N. (N.) cretacea Steinmann, 1899,
and N. (N.) mokragorensis Radoičić & Schlagintweit,
The degree of calcification has often been regarded as a
diagnostic character at the species level. Génot (1985,
figs. 2, 3; 1994, tables 1–3) illustrated the variability of the
intercellular calcification between discrete species and
within a single species. Out of the eight Cenozoic representatives of the subgenus Larvaria he studied, none are
calcified on the primary segment of their laterals, three are
rarely strongly calcified along the axial stem: N. (L.)
auversiensis (L & J Morellet, 1913)*, N. (L.) encrinula
(Defrance, 1822), and N. (L.) montiensis (L & J Morellet,
1922), while fragments of the central stem are exceptionally
preserved in a fourth species: N. (L.) filiformis (L & J
Morellet, 1913). Similarly for his twelve representatives of
the subgenus Neomeris (Génot, 1994), one only is commonly
heavily calcified: N. (N.) larvarioides (L & J Morellet, 1913),
Neomeris (N.)
conradi Radoičić, 2002
cretacea Steinmann, 1899
decapoaae Radoičić, 2002
N. (Drimella)
Sketch [r2st = secondary segment
of the lateras, sterile;
= gametophore (sporangium)]
assurbanipali (Elliott, 1968)
N. circularis Badve & Nayak,
1984, has not been included
here because it is only known
from a single very poor random
section lacking diagnostic
N. (Larvaria)
drimi Radoičić, 1984
jerinae Radoičić, 1984
mokragorensis Radoičić
& Schlagintweit, 2007
The new species
described herein
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and a second species may exceptionally be: N. (N.) scrobiculata (Gümbel 1872).
*Note: according to Génot and Le Renard (2011),
Neomeris (Larvaria) auversiensis (L & J Morellet, 1913)
is a junior synonym of N. (L.) encrinula (Defrance,
Our new species has calcified stalks, but not the other
Cretaceous species, i.e., neither Neomeris (N.) cretacea nor
N. (N.) mokragorensis. However, we should not retain this
feature as a main diagnostic character because it appears
that the calcification of the main axis is related to the
external diameter of the portion studied. The smaller
illustrated sections (Figs. 2b, d,3a, b,4c) show a calcified
main axis; they represent probably the proximal and ?distal
ends of the thallus. In conclusion, the degree of calcification not only varies from one individual to the other within
the same species (see discussion in Granier 2012), but as
for their modern counterparts it also varies within a single
Figure 3a and b illustrate transverse sections of the same
specimen. They were ‘‘cut’’ some micrometers apart, a
result which was obtained through the making of an acetate
peel prior to the making of the thin-section from the same
rock slice. They show that over a short distance the axial
stem may widen, while the external diameter of the thallus
remains almost constant. Figure 4a illustrates an oblique
section through a large portion of a thallus. The general
shape of the alga is probably not strictly cylindrical, but it
could be either fusiform (thinning at both ends) or more
probably club-shaped (with a narrow proximal end and a
large distal end), a morphology still present in living
Neomeris (Berger and Kaever 1992; Berger 2006).
Morphological features and measurements (summarized
in Table 2) help us to easily discriminate the new species
from the two Cretaceous representatives of the subgenus
Neomeris. Génot (1994) stated that ‘‘features of the
reproductive organs appear to be one of the most useful
criteria to separate species or groups of species’’. It applies
to N. (N.) cretacea, which has ‘‘ovoidal to ellipsoidal
ampullae’’ (Barattolo 1990), though both the new species
and N. (N.) mokragorensis have subspherical ampullae. In
addition, the dimensions of the Brazilian species are also
commonly closer to those of N. (N.) mokragorensis.
However, (1) this ‘‘Alpine’’ species bears more laterals per
verticil, (2) its fertile ampullae are smaller, and (3) its
verticils are set closer, consequently in tangential or oblique sections the gametophores look rather imbricated while
in the Brazilian species they are relatively wider spaced.
For a while, some specimens of N. (N.) cretacea were
erroneously considered as a genuine representative of the
genus Macroporella (Pia, 1912), i.e., Macroporella aptiensis Sokač, 1989, until recently when this author
Fig. 4 Neomeris srivastavai nov. sp. All specimens from sample c
1,638-m depth, Petrobras well 1-RNS-11, State of Rio Grande do
Norte, Brazil. Scale bar is 250 lm (a–d). a Paratype: Oblique section
of a large thallus (Granier et al. 2008, p. 313, pl. 1, fig. a). b Oblique
section of a poorly calcified thallus, only the distal part of the laterals
was calcified. c, d Sub-transverse sections, partly micritized (c) and
coated (d)
acknowledged his mistake (Sokač, 2004). The calcareous
skeleton of N. (N.) cretacea is rather thick and isopachous,
and both secondary laterals and the ampullae are embedded
in the ‘‘calcitic wall’’ where they form pores; in contrast, in
both the new species and N. (N.) mokragorensis, the
ampullae are enveloped in a relatively thin crust and the
morphology of the internal surface of the calcareous tube is
covered with little bumps corresponding to individual
ampullae (Figs. 2b, d, e,3c, d,4a).
Phylum Chlorophyta
Class Dasycladophyceae Hoek et al., 1995
Order Dasycladales Pascher, 1931
Family Dasycladaceae (Kützing, 1841)
Tribe Dasycladeae Pia, 1920
Genus Neomeris Lamouroux, 1816
Subgenus Neomeris (Lamouroux, 1816) Pia in
Hirmer, 1927
Neomeris (Neomeris) srivastavai nov. sp. (Figs. 2b,
d, e, 3a–f, 4a–d)
2008 Neomeris (Neomeris) sp.—Granier et al., Potiguar
Basin (NE Brazil), Ponto do Mel Formation (Upper
Albian-Cenomanian): 313, pl. 1, fig. A
2008 Cymopolia perkinsi—Granier et al., Potiguar
Basin (NE Brazil), Ponto do Mel Formation (Upper
Albian-Cenomanian): 313, pl. 1, fig. C
Origin of the name The species is dedicated to Prof. Dr.
Narendra Kumar Srivastava (1973, 1982a, 1982b, 1984),
Universidade Federal do Rio Grande do Norte (UFRN),
Natal, Brazil, for his contribution to the knowledge of
Cretaceous dasycladalean algae.
Type material The holotype is the specimen illustrated by
the oblique section (Fig. 2b, d) from a thin-section cut in a
core sample at 1,638 m, well 1-RNS-11 (offshore Rio
Grande do Norte, Brazil). Two paratypes were selected to
help in completing the description: one is illustrated by two
transverse sections (Fig. 3a, b), the other by an oblique
section (Fig. 4a). This material is currently housed in the
first author’s collection (it will be later transferred to the
national collections at the Faculty of Sciences in Lyons
(FSL), University Claude Bernard—Lyon I, Villeurbanne).
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Table 2 Biometric data of the
three Cretaceous species
assigned to the subgenus
(L maximum length, D external
diameter, d diameter of the
stem, ‘‘d’’ diameter of the axial
cavity, l0 length of the primary
segment of the laterals, ls length
of the ampulla, w number of
laterals per verticil, h height of a
‘‘whorl’’ (verticil) plus one
interverticillar spacing, p0 width
of the primary segment of the
laterals, p00 width of the
secondary segment of the
laterals, ps width of the
In mm
Neimeris (N.)
srivastavi n. sp.
N. (N.) cretacea
(according to
Barattolo 1990)
N. (N.) mokragorensis
Radoičić and Schlagintweit 2007
(according to Schlagintweit
and Ebli 1995)
1.387 ± 0.302
1.29 ± 0.49
0.32, 0.38, 0.42 (3)
Not applicable
0.44 (1)
0.774 ± 0.220
0.99 ± 0.37
e = (D - ‘‘d’’)/2
0.305 ± 0.053
0.16 ± 0.04
30 %, 33 %, 32 % (3)
29 % (1)
56–68 %
42.9–63.7 %
60–88 %
16 - 34
25 - 34
0.10 - 0.20
0.08 ± 0.06
0.14, 0.16, 0.22 (3)
*0.04 (proximal)
to *0.08 (distal)
0.040 ± 0.006
0.034 ± 0.009
ls (Gametophore)
0.229 ± 0.037
0.13 ± 0.03
ps (Gametophore)
0.088 ± 0.014
(3) = Three measurements
0.097 ± 0.033
(1) = Based on their Fig. 7
Fig. 5 All photomicrographs from sample ALCL 10-3bas 9/83 (Pierre-Yves Berthou Collection), Estoi-Pão Branco, Algarve, Portugal, Upper
Berriasian or Lower Valanginian. Scale bar is 250 lm (a–d). a–d Various random (oblique) sections of a Neomeris sp.
Author's personal copy
Diagnosis Roughly cylindrical (fusiform or club-shaped)
thallus with simple verticils (euspondyle) arranged in a
regular alternation but without a strong imbrication.
Bumpy axial cavity, i.e., inner side of the outer calcareous
tube. Main axis locally calcified, at the proximal and
?distal ends of the thallus, forming an inner calcareous
tube. Laterals, where fertile, consisting of a primary segment bearing a pair of secondary sterile segments set on
both sides of a single spheroidal ampulla (a trait that refers
it to the subgenus Neomeris) in terminal position (choristosporate type). The biometric data (Table 2) should also
be taken into consideration to discriminate this species
from other representatives of the subgenus.
Table 2.
Keys for prospective research
The introduction of this new taxon represents an advance in
the ongoing inventory of the fossil calcareous algae from
Brazil (Granier et al. 2008, 2012). It is also a new step in
the knowledge of the representatives of the genus Neomeris
during the Cretaceous. In Brazil, records are from Albian
and Cenomanian strata (Srivastava 1982a; Granier et al.
1991a, b). According to Barattolo and Romano (2001), the
first worldwide record ever would date back from the
Valanginian or the Hauterivian times with N. (Larvaria)
assurbanipali (Elliott 1968) of the Garagu Formation of
Iraq. However, because Pseudocyclammina lituus (Yokohama, 1890) is a typical foraminifer of this stratigraphic
unit, the age uncertainty regarding this first occurrence
could be reduced from two stages to half a stage, i.e., to the
Early Valanginian. N. (L.) assurbanipali is competing for
the oldest record of the genus with an undescribed species
from Algarve, Portugal (Fig. 5). The latter could be Early
Valanginian or even Late Berriasian in age because it is
found in a phycological assemblage consisting of Clypeina
sp., Fourcadella sp., ? Falsolikanella sp., Permocalculus
sp., Rajkaella minima (Jaffrezo, 1973), Terquemella sp.,
and Zergabriella embergeri (Bouroullec and Deloffre,
1968), above a level with Pseudocymopolia jurassica
(Dragastan 1978, non 1968); it is associated with numerous
foraminifers with agglutinated tests, and Trocholina gr.
alpina (Leupold 1935), Mohlerina basiliensis (Mohler,
1938), and Protopeneroplis trochangulata Septfontaine,
1974. The earliest representative of Neomeris should have
been ‘‘born’’ at the transition between the Jurassic and the
Cretaceous. Now considering the choristosporate model of
reproduction, which is the main trait of the family Dasycladaceae, as emended by Granier and Bucur (Granier et al.
2012), the ancestor could be tracked back to Eodasycladus
from Liassic times (Barattolo et al. 1994, 2012), which
represents a huge stratigraphic gap and accordingly which
leaves us with a large stratigraphical field to seek the
missing links.
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). We thank Petrobras for ceding the investigated material to the
second author of this paper previously to the creation of the Brazilian
Petroleum National Agency—ANP. 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 to the reviewers, Patrick
Génot, Felix Schlagintweit, and an anonymous reviewer, for making
useful suggestions, to Franz T. Fürsich for editorial work and to
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