◄ Carnets Geol. 13 (L01) ►
Contents
[1. Introduction] [2. Taxonomical background]
[3. Description of the new material]
[4. Systematics]
[5. Discussion]
[Bibliographic references]
Département des Sciences de la Terre et de l'Univers,
UFR Sciences et Techniques,
Université de Bretagne Occidentale,
6 Avenue Le
Gorgeu, CS 93837, F-29238 Brest Cedex 3 (France)
Manuscript online since February 28, 2013
[Editor: Michel ;
language editor: Donald E. ]
When the species Heteroporella ? paucicalcarea (, 1970) was erected it was left in open nomenclature. Later on new combinations were introduced but did not meet a general agreement among paleophycologists. Considering the current acception of the Dasycladalean families, we ascribe it to the Family Polyphysaceae and subsequently to the genus Clypeina (, 1845). Its known stratigraphic range is rather brief (Late Hauterivian-Early Barremian), which makes it a good index fossil in Urgonian carbonate platform series. Its geographical distribution is apparently restricted to Western Europe (France, Spain and Switzerland).
Calcareous fossil algae; Dasycladales; Polyphysaceae; Heteroporella; Clypeina; Hauterivian; Lower Barremian.
B. (2013).- Heteroporella ? paucicalcarea (, 1970), an Urgonian Dasycladalean alga revisited.- Carnets de Géologie [Notebooks on Geology], Brest, Letter 2013/01 (CG2013_L01), p. 59-65.
Révision de l' Heteroporella ? paucicalcarea (, 1970), une algue dasycladale urgonienne.- À sa création l'espèce Heteroporella ? paucicalcarea (, 1970) a été laissée en nomenclature ouverte. Ultérieurement de nouvelles combinaisons ont été proposées mais aucune n'a rencontrée l'unanimité parmi les paléophycologues. Compte-tenu de l'acception actuelle des familles d'algues dasycladales, nous rapportons ce taxon à la Famille des Polyphysaceae et par contre-coup au genre Clypeina (, 1845). Sa répartition stratigraphique connue est plutôt courte (Hauterivien supérieur-Barrémien inférieur), ce qui en fait un bon marqueur fossile dans les séries de plates-formes carbonatées urgoniennes. Sa distribution géographique est apparemment réduite à l'Europe occidentale (Espagne, France et Suisse).
Algues calcaires fossiles ; Dasycladales ; Polyphysaceae ; Heteroporella ; Clypeina ; Hauterivien ; Barrémien inférieur.
Further to the 1994 revision of Heteroporella lepina , 1967, i.e., the type-species of the genus Heteroporella ( & ex , 1967), the latter was left with one species only (its type, by definition) out of its original sixteen representatives ( et al., 1994). Most species were ascribed either to Chinianella ex & , 1993, or to Otternstella in et al., 1994; few specific epiteths were considered nomina nuda whereas five species were left in open nomenclature, awaiting revision. Heteroporella ? paucicalcarea , 1970, is part of this last set and required our attention.
From its very beginning (, 1970), Heteroporella ? paucicalcarea was in open nomenclature. The question mark following the generic name was fully justified because it was not obvious whether the illustrated specimens were bearing both fertile (large) and sterile (thin) laterals as stated in the original diagnosis of the genus (, 1967), that is prior to its revision ( et al., 1994). Actually the pores corresponding to the so-called "sterile hairs, located within and/or between the whorls", which were reported by (1970), are merely defined in the interverticillar space by the incompletness of the calcification on the whorls of fertiles laterals.
On the basis of material questionably referred to the species, (2000) introduced the new combination, Similiclypeina paucicalcarea nov. comb. An option that was sound because the genus Similiclypeina , 1993, was erected to group Clypeina-like species the verticils of which are set rather close together ("les espèces clypeiniformes dont les verticilles sont relativement rapprochés", , 1994).
But, in turn, et al. (2009) introduced another new combination, Piriferella paucicalcarea nov. comb., which was fully backed by (in et al., 2009; , 2011). There are some side consequences:
1- for instance, the species Clypeina somalica et al., 1983, was successively transferred to Holosporella , 1930 ( et al., 1991), then to Similiclypeina , 1993 (, 1993), and finally to Piriferella , 1996 (, 1996; et al., 2009). et al. (1983) stated that its thallus bears verticils with noncontiguous branches, even near to the main axis ("Rameaux (...) jamais contigus même à proximité du siphon", ibid.); accordingly this species should have been excluded from genera Clypeina (, 1845) and Similiclypeina. The two remaining genera, Holosporella , 1930, and Piriferella , 1996, have similar characteristics, the only difference between them being the shape of the laterals. Though the species somalica better fits the diagnostic parameters of Holosporella ( et al., 1991; & , 1993), et al. (2009) favoured the Piriferella option. In addition, without giving much consideration to the measurements (particularly the spacing of the verticils and the relative size of the fertile pores), these authors also regarded Piriferella spinosa , 1996, the type-species, and Salpingoporella verrucosa , 1996, as junior synonyms of Piriferella somalica (et al., 1983), a statement we should not agree with;
2- the genus Similiclypeina , 1993, is downsized. Actually it might even be considered as monospecific due to the so far unique morphological trait of the proximal part of the laterals in its type-species, Similiclypeina conradi , 1993;
3- Rather than clearing up the confusion that was there et al. (2009) added more. Let us remember that Holosporella , 1930, Piriferella , 1996, and Salpingoporella in , 1917, belong to the Family Triploporellaceae (, 1920), Heteroporella ( & ex , 1967) is a representative of the Family Thyrsoporellaceae ( & in et al., 2012), whereas both Clypeina (, 1845) and Similiclypeina , 1993, are ascribed to the Family Polyphysaceae (, 1841).
In conclusion, except for one small detail, it would be safer to revert to 's(2000) view: Heteroporella ? paucicalcarea is a Clypeina-like species the verticils of which are set rather close together. Because the species lacks the typical pattern of the laterals in Similiclypeina conradi , 1993, it should "naturally" be relocated among the representatives of Clypeina (, 1845).
Material: The studied material consists of some tens of thin sections, each
containing one to several sections (mostly oblique and more or less well preserved) of the alga.
Part of the material (Fig. 1 
), the one from Haute-Savoie (France), is derived from rock
samples collected by
(1996) during field work to complete a PhD thesis: a first calcareous alga,
Falsolikanella danilovae ( ex ,
1978), was already revised based on the same set of thin sections (
et al., 2000). Additional material (Fig. 2 )
was collected by the author from a
section in Drôme (France), where algal remains transported in debris and grain
flows can be dated by the ammonites found in the adjacent marls and limestones (
et al., nearing completion).
| Measurements | , 1970 | This work ( Collection) |
| L | 2.5 mm | > 1.75 mm |
| D | 0.85 - 1.5 mm | 0.89 - 1.28 mm |
| d | 0.3 - 0.57 mm | 0.32 - 0.47 mm |
| d/D | 30% - 48% | 25% - 35% |
| h1 or h2 | 0.17 - 0.28 mm (h1) | 0.23 - 0.43 mm (h2) |
| l | 0.30 - 0.43 mm | |
| p | 0.08 - 0.15 mm | |
| e | 0.04 - 0.07 mm | |
| w | 9 - 18 | 8 - 14 |
Table 1: Biometric data of Clypeina paucicalcarea (, 1970) nov. comb. (L: maximum length; D: external diameter; d: diameter of the stem; h1: interverticillar spacing sensu stricto, i.e., from the top surface of one whorl to the bottom surface of the next; h2: interverticillar spacing sensu lato, i.e., distance from a reference plane in one whorl to the same plane in the next; l: length of the laterals; p: width of the laterals; e: thickness of the calcareous coating on the laterals; w: number of laterals per verticil).
Description: The algal thallus is roughly cylindrical with
a
cylindrical main axis bearing whorls (euspondyle type) consisting of 8 to 16 (w) laterals. The calcareous
skeleton is broken into segments with rarely more than ten verticils. These verticils are close set, but
not imbricated. The interverticillar spacing sensu stricto (h1), i.e.,
the distance from the top surface of one whorl to the bottom surface of the next whorl, is always a positive
value (see Table 1). The interverticillar
spacing sensu lato (h2), i.e., the distance from a reference plane in one
whorl (insertion points of a verticil, for instance) to the same plane in the next whorl, is more than twice the thickness of a
verticil (see Table 1), that is more than twice the diameter of the laterals (p). These laterals
are of the first order only, slightly phloiophorous in shape, i.e., increasing in diameter outward,
and supposedly fertile because they form quite wide pores.
Within a single verticil, they are first gently inclined (probably upward, in
the growth direction) with respect to the algal main axis (up to ? 60°), then
they rapidly bend outward and are nearly perpendicular in the medial and distal parts of the
whorl, which are almost planar. These laterals are close set in the proximal part of
the verticil itself and thus embedded in a joint calcareous coating. But, because their
diameter does not increase significantly, they rapidly diverge distally and as a result
each lateral gets an individual coating. The star pattern of the verticils
in transverse sections (Fig. 2.c )
reminds us that it is known in Clypeina and in Actinoporella (
in , 1881). The lack of corona structure
above or below the verticil allows us to ascribe the species paucicalcarea
to the genus Clypeina.
Click on thumbnail to enlarge the image.
Figure 1: a-c and e-m: Clypeina paucicalcarea (, 1970) nov. comb.
a. oblique to tangential section. GT 1.065 (Upper Hauterivian, Ha6 HST) Rocher de Cluses, Haute-Savoie, France; b. oblique to axial section. GT 1.122 (Upper Hauterivian, Ha7 LST) Rocher de Cluses; c. oblique to transverse section. FL 2.09 (Lower Barremian, Ba1 LST) Flaine, Haute-Savoie, France; e. axial section. GT 1.066 (Upper Hauterivian, Ha6 HST) Rocher de Cluses; f. tangential section. GT 1.047 (Upper Hauterivian, Ha6 TST) Rocher de Cluses; g. oblique section through 3 verticils. GT 1.067 (Upper Hauterivian, Ha6 HST) Rocher de Cluses; h. oblique section. GT 1.098 (Upper Hauterivian, Ha7 LST) Rocher de Cluses; i. subtransverse section. BA 19a05 (Upper Hauterivian, Ha6 LST) Combe de Balme, Haute-Savoie, France; j. oblique to transverse section. GT 1.101 (Upper Hauterivian, Ha7 LST) Rocher de Cluses; k. oblique section through 3 verticils. GT 1.066 (Upper Hauterivian, Ha6 HST) Rocher de Cluses; l. oblique section through 4 verticils. FL 2.09 (Lower Barremian, Ba1 LST) Flaine; m. oblique section through (?) 2 verticils. GT 1.122 (Upper Hauterivian, Ha7 LST) Rocher de Cluses.
d. oblique to tangential section of a "look-alike" Triploporellacean alga (Clypeina is a Polyphysacean alga). Note that laterals are arranged in quinconxes. C5.709 (Upper Hauterivian, Ha7 LST) Plateau d'Andey, Haute-Savoie, France.
All thin sections are stored at the Muséum d'Histoire naturelle, Genève [Scale bar: 250µm]: G. (a-c and e-m) and J. (d) collections [Scale bar: 250µm].
Click on thumbnail to enlarge the image.
Figure 2: a-e: Clypeina paucicalcarea (, 1970) nov. comb. L'Estellon section, Drôme, France. Ages are directly constrained by ammonites found in strata below and above.
a. tangential section through 4 verticils. EST 26.5 (Nicklesi Zone, Lower Barremian, LST Ba1); b. oblique section through 5 verticils. EST 92.3 (Moutonianum Zone, Lower Barremian, LST Ba3); c. transverse section of a verticil with 8 primary laterals. EST 49.5 (Pulchella Zone, Lower Barremian, LST Ba2); d. oblique section through 3 verticils. EST 92.3 (Moutonianum Zone, Lower Barremian, LST Ba3); e. oblique section through 4 verticils. EST 49.7 (Pulchella Zone, Lower Barremian, LST Ba2).
All thin sections from the author's collection [Scale bar: 250µm].
Phylum Chlorophyta
Class Dasycladophyceae et al., 1995
Order Dasycladales , 1931
Family Polyphysaceae (, 1841)
Genus Clypeina (, 1845)
Clypeina paucicalcarea (, 1970)
nov. comb.
1970 Heteroporella ? paucicalcarea n. sp.- , p. 68-69, Fig.
5 (holotype, duplicated in Fig. 3
herein), Pl. III, figs. 1-4; Pl. IV, figs. 1-3
1970 Heteroporella ? aff. paucicalcarea n. sp.- , p. 68-69, Pl. IV, fig. 4
1973 Heteroporella (?) paucicalcarea.- , p. 80, Pl. 3, figs. 9-11 & 16
1976 Heteroporella ? paucicalcarea.- , Pl. XXIV, figs. 14-16
1976 Heteroporella ? paucicalcarea.- & , p. 185-186, Figs. 10.a-b & 13a
1976 Heteroporella (?) paucicalcarea.- , p. 177, Pl. 4, fig. 7
1978 Heteroporella ? paucicalcarea.- et al., p. 133, Pl. 15, figs. 5 (= Fig. 5 in , 1970), 6 (= Pl. III, fig. 1 in , 1970), 7 (= Fig. 10.a in & , 1976) & 8 (= Fig. 10.b in & , 1976)
1980 Heteroporella ? paucicalcarea.- , Pl. 111, figs. 3–5
1980 Heteroporella (?) paucicalcarea.- , p. 252-253, Pl. XIX, figs. 1, 2 (= Pl. 3, fig. 9 in , 1973), 3 & 4 (= Pl. 3, fig. 11 in , 1973)
1989 Heteroporella ? paucicalcarea.- & , p. 281-282, Pl. II, fig. 11
1993 Heteroporella ? paucicalcarea.- , Pl. 2, fig. 1 (= Pl. 4, fig. 7 in , 1976)
1993 Heteroporella ? paucicalcarea.- & , p. 33
non 1993 Heteroporella ? aff. paucicalcarea.- et al., p. 64, Pl. 3, fig. 3
non 1993 Heteroporella (?) paucicalcarea.- et al., Pl. II, fig. 4
non 1993 Heteroporella ? paucicalcarea.- & , Pl. 5, figs. 1-2
1994 Heteroporella ? paucicalcarea.- et al., p. 135
non 1994 Heteroporella ? paucicalcarea.- , p. 152, Pl. VI, fig. 13
non 2000 Similiclypeina paucicalcarea nov. comb.- , p. 60, Pl. IV, fig. 7
non 2000 Similiclypeina aff. paucicalcarea.- et al., Pl. VIII, figs. 3-4
2007 Similiclypeina paucicalcarea.- et al., Figs. 4.6 & 4.9
2007 Salpingoporella genevensis.- et al., Pl. 6, fig. o
2007 Piriferella paucicalcarea.- et al., Pl. 6, fig. p
non 2007 Piriferella paucicalcarea.- et al., Pl. 6, fig. q
2009 Piriferella paucicalcarea nov. comb.- et al., p. 26
2011 Piriferella paucicalcarea.- , p. 628, Pl. 1, figs. 6-7; Pl. 2, fig. 13; Pl. 6, figs. 5 & 8
Emended diagnosis: Clypeina representative with close set verticils consisting of 8 to 16 slightly phloiophorous laterals. Corresponding pores first gently inclined, then rapidly bending outward and almost horizontal distally. Biometric measurements (Table 1) might help discriminating this species from the other representatives of the genus.
Click on thumbnail to enlarge the image.
Figure 3: Holotype of Clypeina paucicalcarea (, 1970) nov. comb. Oblique section through 6 verticils. Sample 319, slide 6 (Upper Hauterivian, TST Ha5); Rocher-des-Hirondelles, La Rivière, Ain, France; M.A. Collection, Muséum d'Histoire naturelle, Genève. Scale bar: 250µm [Some rights reserved].
Clypeina paucicalcarea (, 1970) nov. comb. has been reported from strata ranging in age from the Hauterivian to the Bedoulian: (? Upper) Hauterivian (& , 1989), Lower Barremian (, 1970; , 1976; & , 1976; , 1976, 1993; , 1980; et al., 2007; et al., 2007), undifferencied Barremian (, 1980), undifferencied Barremian-Bedoulian (, 1973). The authors reporting the early finds were not able to discriminate the age of the Urgonian strata they were dealing with. Today the known range can be reduced to the (?) Late Hauterivian - Early Barremian interval. The first occurrence is not accurate because we lack information on Lower Hauterivian (and even Valanginian) shallow-water carbonates. The last occurrence is better approximated. According to (1993) the species is not known above the Pulchella Zone (Early Barremian) although according to (personal communication, Dec. 2012) it is still present in the Moutonianum Zone, i.e., in the last ammonite zone of the Early Barremian.
This Urgonian species was found in several localities of W Switzerland (Genève: , 1970), S France (Eastern Pyrenees: , 1973, 1980, , 1976, & , 1976; Provence: , 1976, 1993; Ardèche and Drôme: , 1980, et al., 2007, et al., 2007, , 2011), and N Spain (Eastern Pyrenees: & , 1976, not illustrated). There it is commonly found associated to the classical "lower" Urgonian species: Falsolikanella danilovae ( ex , 1978), Pseudoactinoporella fragilis , 1970, Salpingoporella genevensis ex et al., 1973, S. polygonalis , 1996, S. muehlbergii (, 1902), S. melitae , 1967, ... Records outside that area, i.e., in Hungary ( et al., 1993), Romania ( et al., 1993; , 1994, 2000), Slovakia ( & , 1993) and Turkey ( et al., 2000), are not certified. Therefore the 's (1993) hypothesis regarding a possible algal provincialism in Western Europe remains valid.
The author thanks F. and I. for their review of the original manuscript, J. for having given him access to his collection and to that of G. , and B. for providing information on the stratigraphic position (including stratigraphic sequence) of the illustrated specimens. Field work (at L'Estellon, Drôme) in 2012 was supported by "Carnets de Géologie" Association.
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