GeolCarp_Vol70_No2_183

GEOLOGICA CARPATHICA

, APRIL 2019, 70, 2, 183–190

doi: 10.2478/geoca-2019-0010

www.geologicacarpathica.com

Dasycladalean alga Furcoporella diplopora Pia (1918) from

the type-locality (upper Paleocene–Eocene, Radstadt,

Austria): new remarks and taxonomic implications

MARIO DE MATTEIS

and FILIPPO BARATTOLO



Dipartimento di Scienze della Terra “A. Desio”, Università degli Studi di Milano, Via Mangiagalli 34, 20133, Milan, Italy;

mario.dematteis@unimi.it

Dipartimento di Scienze della Terra, dell’Ambiente e delle Risorse (DiSTAR), Università degli Studi di Napoli Federico II, Via Vicinale

Cupa Cintia, 21, 80126, Naples, Italy;



lippolo@unina.it

(Manuscript received November 13, 2018; accepted in revised form February 21, 2019)

Abstract: The dasycladalean green alga Furcoporella diplopora Pia (1918) exhibits a peculiar structure comprising pairs

of diverging and rather indistinct pores. Because of such structure, the genus Furcoporella has been a matter of debate.

Currently, there is still no agreement on its taxonomic position within Dasycladales. This paper provides a re-description

of the type-species F. diplopora Pia (1918), based on Pia’s type material from the upper Paleocene to Eocene reworked

carbonates of Radstadt/Pongau, Austria. New observations carried out on this material allowed the acquisition of

additional and more complete data on this quite neglected alga. Some morphological characteristics of the thallus,

specifically of the laterals are discussed and a suite of potential taxonomic implications are evaluated. Finally, according

to interpretations the genus Furcoporella is ascribed to the Family Triploporellaceae and more specifically it should be

placed within the tribe Thyrsoporelleae.

Keywords: micropaleontology, taxonomy, green algae, Dasycladales, Furcoporella.

Introduction

The dasycladalean alga Furcoporella diplopora was intro-

duced by Pia (in Trauth 1918), investigating the Paleocene to

middle Eocene detrital limestones, exposed near Radstadt/

Pongau, in Austria. Describing the species, the same author

erected the genus Furcoporella. Currently it is still considered

to be a poorly known genus (Deloffre & Génot 1982), despite

its reported occurrences in many Paleocene–Eocene Tethysian

shallow-water limestones. Besides the type-locality, F. diplo

pora specimens have been found in the Middle East (Elliot

1956, 1968), Libya, Egypt (Pfender & Massieux 1966; Kuss &

Leppig 1989), China (Yu Jing 1976) and Anatolia (Radoičić &

Özgen-Erdem 2011).

Presumably, the incomplete understanding of this taxon is

a consequence of the problematic structure exhibited by

F. diplopora that displays few pairs of diverging pores

arranged in whorls around the central stem cavity.

Discussions on this dasycladalean alga started, indeed,

with Pia’s original generic and specific descriptions

(in Trauth 1918): these were hypothetical (see Paleontological

description) and Pia clarified the necessity of further modi-

fications, admitting that his generic diagnosis was only

temporary. Additionally, although Pia figured two sections

of F. diplopora (pl. 1. fig. 1–2), he did not designate

a holotype.

A notable number of descriptions have followed the original

study, coupled with attempts to unravel the taxonomic rela-

tionships of the genus Furcoporella with other dasycladalean

algae (Pia, in Hirmer 1927; Pfender 1940; Elliott 1968;

Bassoullet et al. 1979; Deloffre & Génot 1982, Deloffre 1988;

Deloffre & Granier 1992). Consequently, this genus is central

to much debate, since there is no agreement on the tribe in

which it should be placed.

It is also worth mentioning that Furcoporella, which corre-

sponded formerly to a monospecific genus, encloses the spe-

cies Furcoporella vasilijesimici (Radoičić, 2005) described

from the lower Valanginian detrital limestones of the Metohija

Unit (Mirdita Zone, Kosovo).

Considering that the taxonomic placement of F. diplopora

remains unresolved, this taxon deserves to be the subject of

a more in-depth paleontological study. This paper provides

a formalized and detailed re-description of this species, made

on the basis of Pia’s type material.

Geological setting

The type-locality is situated near Radstadt (see Trauth 1918)

in the Enns Valley area (47°22’48.3” N, 13°26’01.0” E),

adjacent to the Tauern tectonic Window and the Northern

Calcareous Alps. The whole area belongs to the Eastern Alps

realm. Here, the Northern Cal ca reous Alps constitute part of

the Austro-Alpine nappe complex and represent its most

prominent tectonic unit. The Eastern Alps stacked nappes con-

sist of sedimentary deposits of Late Permian to Eocene age

and the entire succession is strongly dominated by Triassic

carbonates (e.g., Wagreich & Faupl 1994; Mandl 2000).

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The Valley of the Enns river is filled by Oligocene to

Quaternary deposits (see Keil & Neubauer 2011 for a review).

It corresponds to a WE-striking fault-controlled valley deve-

loped after Neogene time (Keil & Neubauer 2009). Since that

period, the Eastern Alps have been affected by strike-slip

movements, parallel to the Alpine orogen, accommodating

lateral extrusion tectonics (Ratschbacher et al. 1991). Such

strike-slip scheme produced fault-induced lowlands and intra-

montane basins (Dunkl et al. 2005) and modified the drainage

pattern setting the stage for the modern river system (Frisch et

al. 1998).

Near Radstadt, exposed fluvial conglomerates contain

reworked shallow-water carbonate sediments (e.g., Trauth

1918). Reworked calcareous clasts comprise: i) platform-de-

rived limestone components (with corals, foraminifera and

calcareous algae); ii) other carbonate debris with nummulitids

(Trauth 1918; Moussavian 1984). Such reworked carbonates

also host Furcoporella diplopora specimens, and have been

interpreted as late Paleocene/early Eocene in age (J. Pignatti

2018, personal communication).

In the Eastern Alps, Early Paleogene shallow-water carbo-

nates are exposed only as small-scale outcrops or as reworked

clasts and boulders within younger poorly-dated conglome-

rates (Nebelsick et al. 2005), as in the case of Radstadt/Pongau.

These localized occurrences likely correspond to the remnants

of a former shallow-water shelf environment, developed since

the Middle Paleocene time on the southern edge of the Nor-

thern Calcareous Alps (e.g., Moussavian 1984; Nebelsick et

al. 2005). Such limestone beds were indeed repeatedly exposed

since the late Paleocene to the Eocene (Kázmér et al. 2003)

and subsequently subject to block segmentation and fluvial

erosion during Oligocene and Miocene times (Moussavian

1984).

Material

Furcoporella diplopora specimens in Fig. 1 represent Pia’ s

type material (in Trauth 1918). Such material consists of two

thin sections labelled CLXXII.1 and CLXXII.2., belonging to

the ‘Pia Collection’, stored at the Natural History Museum of

Vienna. One of the authors (F. Barattolo) took the photomicro-

graphs in 1993, during the meeting ‘Alpine Algae’, examining

Pia’s original material.

Repository and institutional abbreviation: types examined

in this study are deposited in the Natural History Museum of

Vienna, Department of Geology & Paleontology (NHMV-

DGP), Pia collection, slides labelled CLXXII.1 and CLXXII.2.

Paleontological description

Order Dasycladales Pascher, 1931

Family Triploporellaceae (Pia, 1920) Berger & Kaever 1992

Tribe Thyrsoporelleae Pia 1927

Genus Furcoporella Pia in Trauth 1918

Type-species Furcoporella diplopora Pia in Trauth 1918

Introducing the species F. diplopora, Pia erected the genus

Furcoporella (Pia in Trauth 1918). The original generic

description is reported hereafter (English translation by

Deloffre & Génot 1982):

“General cylindrical shape. The pores are widened towards

the exterior. At the level of the verticils, they are joined 2 by 2

at the base and are few in number. Unsegmented skeleton. This

definition must be considered as temporary, for it seems likely

that it will have to be modified as forms related to our Eocene

species will be discovered.”

The same author then provided the following short descrip-

tion for the type-species (from Deloffre & Génot 1982):

“The thallus seems to be always straight. The calcification

does not reach the axial cell.”

Elliott (1968) added further observations both on the genus

and on the type-species after the study of larger Middle East

specimens. In his revision, the author described the ramified

pores, observable in transversal sections, as short primary late-

rals subsequently dividing into two divergent secondaries.

Deloffre & Génot (1982) supplied the following comment/

remarks for the genus Furcoporella and the type-species:

“Furcoporella is still a littleknown genus, relatively and

rarely quoted. It is a hollow cylindrical calcareous tube with

numerous regular horizontallyset verticils with a very short

primary canal divided into two divergent secondary branches.

This genus is found between the Paleocene and the Lutetian of

Europe, Middle East and China”

“Furcoporella diplopora — Cylindrical dasyclad tube with

numerous horizontallyset verticils. Each verticil shows a single

very short and large primary branch divided almost at once

into two divergent secondaries widening to the exterior […]”

Fig. 1. Furcoporella diplopora Pia (1918), upper Paleocene–lower Eocene of Radstadt/Pongau (Enns Valley, Austria). A – Lectotype; trans-

versal section. It corresponds to the specimen figured by Pia (in Trauth 1918, pl. 1, fig. 2). Notice the evident and wide diverging pores, NHMV-

DGP Pia CLXXII.1. B – Transversal sections, NHMV-DGP Pia CLXXII.1. C – Transversal slightly oblique section, NHMV-DGP Pia

CLXXII.1. D – Oblique section. Notice the vertical section of the pores and the quite indistinct inner surface of the calcification, NHMV-DGP

Pia CLXXII.1. E – Longitudinal section of a specimen showing the vertical section of the pores. Notice their overall hourglass shape resulting

from the cut of the swollen secondary laterals (the section does not cut axially the primary laterals), NHMV-DGP Pia CLXXII.1. F – Oblique

section showing ordered doubled pores resulting from the divergent secondary lateral. Notice that they are arranged in continuity in simple

subsequent whorls, NHMV-DGP Pia CLXXII.2. G – Oblique section, NHMV-DGP Pia CLXXII.2. H – Oblique section, NHMV-DGP Pia

CLXXII.2. I – Oblique section, NHMV-DGP Pia CLXXII.2. J – Slightly oblique transversal section. The section transversally cuts two subse-

quent whorls, NHMV-DGP Pia CLXXII.1. K – Oblique-transversal section, NHMV-DGP Pia CLXXII.2. L – Oblique-transversal section,

NHMV-DGP Pia CLXXII.2.

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Description in this study of F. diplopora expands Pia’s first

description and includes the observations added by Elliott

(1968), interpreting diverging pores as first and second order

laterals. Furthermore, it supplies additional data to the remarks

provided by Deloffre & Génot (1982).

Furcoporella diplopora P

(in Trauth) 1918

Fig. 1 A–L

1918 Furcoporella diplopora n. gen., n. sp. – PIA, plate 1, fig. 1, 2,

text-fig. 4b.

1927 Furcoporella diplopora – PIA, p.86.

1940 Furcoporella diplopora – PFENDER, p. 242–243.

1956 Furcoporella diplopora – ELLIOT, plate 2, fig. 5–6.

1966 Furcoporella diplopora – PFENDER & MASSIEUX, text-fig. 4,

plate 4, fig. 8–9.

1968 Furcoporella diplopora – ELLIOT, plate 11, fig. 7–9.

1976 Furcoporella diplopora – YU-JING, plate 9, fig. 7–8.

1982 Furcoporella diplopora – DELOFFRE & GÉNOT, plate 8,

fig. 2–4.

1989 Furcoporella diplopora – KUSS & LEPPIG, fig. 9, g–h; fig. 10a.

2011 Furcoporella diplopora – RADOIČIĆ & ÖZGEN-ERDEM,

fig. 11, f–i.

Lectotype

Pia (1918) did not designate a holotype for

F. diplopora. Among photomicrographs, one of the original

specimens figured by Pia (in Trauth 1918, pl. 1, fig. 2) is

reported and selected as the lectotype. It corresponds to

the transversal section A of Figure 1 (NHMV-DGP Pia

CLXXII.1).

Description

The investigated dasycladalean alga is characterized by

a sim ple, cylindrical, non-segmented thallus with a relatively

large central stem. Large diverging pores are interpreted as

results of a short primary lateral splitting horizontally into two

divergent second-order laterals, swollen outwards (phloiopho-

rous). Laterals are arranged in a closely packed configuration

and are set in continuity in subsequent whorls (euspondyl).

Whorls are set orthogonally to the stem axis since both

the short, thick primary laterals and the secon dary ones are

generally horizontal. The latter flare out distally to make

a quadrangular cortex. Reproduction is supposed syringo-

sporate, less probably endosporate.

Remarks

General features of the calcareous skeleton

The calcification corresponds to a simple massive calca-

reous sleeve. It preferentially envelops the laterals up to their

distally widened ends, approaching the cortical zone. The cen-

tral cavity is filled either by micritic matrix or blocky sparite.

The pores, remnants of the former laterals, are commonly

filled by matrix. Pores are connected to the central cavity by

relatively wide openings (Fig. 1C, D) and the inner surface of

the skeleton does not display sharp edges (Fig. 1C, E, I). This

suggests that the innermost part of the calcification was not

directly in contact with the central stem outer wall. Therefore,

the actual central cavity diameter is slightly larger than

the for mer axial stem diameter. Such a characteristic is pre-

sumably a primary, morphological feature (i.e., calcification

does not envelop laterals base).

Diverging laterals

Transversal sections of the calcareous skeleton (Fig. 1A–C)

offer the opportunity to clearly observe the pores splitting into

two divergent canals. In geometrical terms and in the case

of an even number of laterals, if the axial section of the ske-

leton cuts the primary laterals axially, it will not cut the diver-

gent secondaries at the same time. The longitudinal and

oblique sections (Fig. 1D, F), however, display elongated

pores with an overall hourglass shape. The hourglass-

shaped pores result from a cut passing longitudinally along

the primary pore and just one of the two divergent secondary

laterals. Such elongated hourglass shape of the pores may be

also produced by the relative high thickness of the thin

sections.

The number of primary laterals per whorl can be simply

obtained from transversal sections (Fig. 1A–C). It corresponds

to 5–6, a number markedly lower than that observed by

Radoičić & Özgen-Erdem (2011, fig. 11g), i.e., 10–12.

According to the geometry observed in the oblique sections

of the calcareous skeleton (Fig. 1F, H), the phloiophorous

secondary laterals are arranged horizontally, appearing elon-

gated and relatively thick at the same time. Their angle of

divergence has been obtained from the transversal sections

and corresponds to 43° (average value, see Table 1 for other

main biometrical parameters). At the far end, each secondary

lateral flares out and comes in touch with the others alongside,

making a cortex.

Since the calcification does not envelop the central stem and

the most proximal part of the laterals, the morphology of

their junction with the stem is still debated. Additionally, there

is no trace of narrowing between the large inner pore and

the diverging ones.

Reproduction

No evidence of reproductive organs has been observed

within the calcareous skeleton, neither within the central

cavity nor in the primary laterals. There is also no trace of

reproductive structures such as ampullae. According to

the current state of knowledge, the reproduction type of

F. diplopora remains unknown.

Reconstruction in this study (see Conclusions) proposes

large openings connecting both the laterals and with the cen-

tral stem. This kind of large connection is here considered to

be a primary, morphological characteristic. Presumably, these

wide portions hosted gametophores (cysts), thus, the repro-

duction type is herein supposed syringosporate (Granier 2010;

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Mathieu et al. 2011; Granier & Bucur in Granier et al. 2013)

(see Interpretation of the diverging pores and attribution to

higher rank).

Interpretation of the diverging pores

and attribution to higher rank

Studies of fossil dasycladalean algae are frequently met by

problems connected to the calcification. Since the calcifica-

tion degree is indeed strongly variable, it produces highly

different kinds of skeleton (e.g., extensive, scattered or largely

absent calcification). The structure of fossil dasycladaleans

must be interpreted principally on the basis of the pores left by

calcified parts. Consequently, the development of algal recon-

structions may be hard and challenging in the case of scattered

or even largely absent calcification.

In the specific case of Furcoporella, though it exhibits

an evident calcareous skeleton, the calcification of the diver-

gent pores remains indistinct. Indeed, the absence of clear

narrowing at the pore junctions opens the field to ample specu-

lation regarding the order of the laterals (i.e., first+second-

order laterals vs. first-order only). A suite of junction

combinations and potential arrangements of the laterals is

reported in Fig. 2. The first two solutions (Fig. 2A, B) show

two orders of laterals and consequently an euspondyl arrange-

ment. Both are geometrically consistent with the thin section

observations. The only difference between these first two

solutions lies in the width of the connections. The other solu-

tions (Fig. 2C, D) are instead representative of a metaspondyl

arrangement. In solution C, two first-order laterals are directly

connected to the stem axis, while the last solution (Fig. 2D) is

characterized by a vestibule supporting the tuft of laterals.

Among the reported morphologies, however, the euspondyl

arrangement is considered to be the most probable configura-

tion. It is worth mentioning that each of the four alternative

solutions provided in Fig. 2 may fit well the geometry of

the pores of Furcoporella. Of course, the choice of one among

the four-reported hypotheses leads to relevant taxonomic

consequences. According to the features observed in thin sec-

tion, the “channelled primary lateral” option (Fig. 2A) is

selected as the most probable one, since it requires fewer

assumptions.

A metaspondyl arrangement (with or without a vestibule)

would cause the shifting of the genus Furcoporella to

the Family Diploporaceae. In his first description, Pia actually

pointed out the morphological affinity of F. diplopora with

the genus Diplopora (in Trauth 1918): the specific name he

has chosen is certainly a clear reference to this similarity.

Anyway, the elements and the geometries figured in Fig. 2C, D

are typical of Upper Paleozoic and Triassic genera. Given

the remarkable time gap between these older genera and

the Eocene age of F. diplopora, it is considered these two solu-

tions less probable than the first two. Because of the euspon-

dyl configuration, the genus Furcoporella should be placed in

the Family Triploporellaceae: this attribution is indeed in

agreement with the one already proposed by Radoičić (2005).

The same author (Radoičić 2005) and Radoičić & Özgen-

Erdem (2011) assigned Furcoporella diplopora to the tribe

Salpingoporellae, which includes verticillated genera sharing

the occurrence of only primary laterals. Here, the occurrence

of two orders of laterals prompts us to exclude such

attribution.

Lastly, excluding the assignment of the genus to the subtribe

Macroporellinae initially proposed by Pfender (1940): actually,

this subtribe formerly included both euspondyl and aspondyl

forms (Pia 1927), but now is restricted to grouping only some

aspondyl genera (Deloffre 1988).

Specimen

Thin Section

(°)

d/D

CLXXII.1

175.3

317.6

69.2

35.1

0.55

CLXXII.1

174.7

383.7

81.6

44.3

0.46

CLXXII.1

195.2

370.5

68.9

33.8

0.53

CLXXII.1

186.9

350.6

0.53

CLXXII.1

138.9

254.7

86.7

0.55

CLXXII.2

152.2

258.1

68.3

0.59

CLXXII.2

167.1

284.5

68.2

0.59

CLXXII.2

161.3

287.8

88.4

0.56

CLXXII.2

173.2

285.9

78.7

0.61

CLXXII.1

147.6

274.6

60.9

31.8

0.54

CLXXII.2

147.5

256.4

0.58

CLXXII.2

154.8

291.1

67.0

34.9

0.53

Min

138.9

254.7

68.2

60.9

31.8

0.46

Max

195.2

383.7

88.4

81.6

44.3

0.61

N. of measures

Average

164.6

301.3

78.1

69.5

36.0

0.55

St. deviation

17.14359

44.56555

0.51640

9.66737

7.52627

4.81697

3.31662

0.03947

Table 1: Main biometrical parameters of Furcoporella diplopora Pia. All size parameters are given in µm, with the exception of

��, given in

degrees. d — inner diameter of the central cavity; D — outer diameter of the calcareous skeleton; w — number of primary laterals per whorl;

h — height between subsequent whorls; p — width of primary laterals; p’ — width of secondary laterals (measured at the base); α — average

angle of divergence of the secondary laterals.

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Among the Triploporellaceae, Furcoporella diplopora

shows a remarkable resemblance to some other Paleocene–

Eocene species belonging to the tribes Dissocladelleae and

Thyrsoporelleae. The occurrence of two orders of strong

late

rals allows comparison of Furcoporella with Disso

cladella. The latter genus shows indeed a short, globose

primary lateral and phloiophorous secondaries similar to

those of Furco porella, but in the case of Dissocladella

the primary lateral is attached to the central stem through

a small pore and the number of secondary laterals is always

more than two.

Differently, the short and strong primary lateral of

Furcoporella exhibits a wide opening with the central cavity.

Such a wide junction closely resembles that of Thyrsoporella

and Belzungia. However, these two genera can still be dif fe-

rentiated since they display four to five orders of laterals fining

outwards (e.g., Radoičić & Özgen-Erdem 2011).

It is suggested that the genus Furcoporella seems to be

strictly related to the genus Anatolia Radoičić & Özgen-Erdem

2011. According to the authors (p. 231), in Anatolia “the thal

lus consists of cylindrical articles with rather spaced whorls.

The whorl consists of three orders of horizontal to subhori

zontal laterals arranged in vertical rows. The primary laterals

divide into two divergent, strong and stocky secondaries. Each

secondary lateral gives rise to 4–5 slender tertiaries, distally

enlarged. Reproductive organs are unknown.” Apart from

the shape of the thallus (presumably not articulated vs. articu-

lated), the two genera share similar features of the primary and

secondary laterals. The only remarkable difference consists of

the absence and presence of tertiary phloiophorous laterals

respectively. In Anatolia the sites of reproduction were see-

mingly located within the primary and secondary laterals and

the relatively thin phloiophorous tertiary laterals functioned as

assimilators. In Furcoporella secondary laterals alone pro-

bably performed both reproduction and assimilation. Such

hypothesis coupled with the previously listed morphological

similarities lead us to assign the genus Furcoporella to tribe

Thyrsoporelleae.

Concerning the issue of family level attribution, recently

Granier & Bucur in Granier et al. (2013) proposed the erection

of the new family Thyrsoporellaceae to include syringosporate

genera. This clade is directly derived from the family

Triploporellaceae that consequently becomes paraphyletic.

Actually, all Dasycladales clades at family level (Berger &

Kaever 1992: Granier & Bucur in Granier et al. 2013) except

Seletonellaceae and Dasycladaceae, must be considered para-

phyletic if not polyphyletic and need a general re-examination

that takes into account DNA data from extant forms (Berger et

al. 2003; Verbruggen et al. 2009). For this reason, instead of

following the classification proposed by Granier & Bucur in

Granier et al. (2013), we prefer the classic Berger & Kaever

(1992) classification, assigning the tribe Thyrsoporelleae to

the family Triploporellaceae.

Conclusions

This paper presents a detailed revision of Furcoporella

diplopora Pia and new paleontological data on the genus

Furcoporella. Overall, the re-description reported herein

expands Pia’s first description and matches the observations

added by Elliott (1968) on the laterals. A comprehensive

reconstruction of Furcoporella diplopora was provided as

a result in which all the previously examined features are

coherently represented (Fig. 3).

This study focused on this problematic and neglected

alga, with the aim of facilitating a complete taxonomic

assignment of the genus Furcoporella in Dasycladalean

systematics. Besides simply modelling the thallus in this

reconstruction, we have taken into account a suite of diverse

lateral arran gements, evaluating potential taxonomic impli-

cations. Finally, according to this interpretation the genus

Furcoporella was assigned to the Family Triploporellaceae,

also sug gesting that this taxon should be placed within

the tribe Thyrsoporellae.

Fig. 2. Alternative arrangements of the laterals fitting the geometry of

the divergent pores (in transversal view): A — euspondyl, showing

a single primary lateral bearing two divergent secondaries, both

with wide connections; B — euspondyl, showing a single primary

lateral bearing two divergent secondaries with narrow connections;

C — meta spondyl arrangement, showing a tuft of two primary laterals

directly attached to the central cylindrical stem; D — metaspondyl

vestibulated, showing primary lateral set in a tuft of two, sustained by

a vestibule.

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In sum, our observations confirm the presence of two orders

of laterals: the wide divergent pores are interpreted as the

result of short primary laterals bearing two divergent phloio-

phorous secondaries. These are connected to the primary

lateral by large openings (Fig. 2A). Accordingly, laterals with

no tapered bases were reproduced. The laterals and the subse-

quent cortical meshes are arranged in ordered repetitive rows.

The cortical meshes, corresponding to the uncalcified outer

ends or edges of secondary laterals, have been drawn as

sub-rectangles set horizontally; this solution is geometrically

consistent given the number of secondary laterals and the arran-

gement in tangential and oblique section (Fig. 1F). Addi-

tionally, according to the evidence observed in thin section

(see General features of the calcareous skeleton), the calcifi-

cation has been reproduced without directly touching the cen-

tral stem wall.

Acknowledgements: The paper benefited from comments by

reviewers Bruno Granier, Ioan I. Bucur and Marc André Con-

rad. The authors also wish to thank Mathias Harzhauser and

Andreas Kroh (Natural History Museum, Vienna) for chec-

king the labels of the type-material. Johannes Pignatti

(University of Rome) is acknowledged for suggestions about

the age of the microfacies of F. diplopora. We are grateful to

Sarah M. Jacquet (University of Missouri) for her valuable

comments and review of the English.

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