GeolCarp_Vol62_No6_501

www.geologicacarpathica.sk

GEOLOGICA CARPATHICA

, DECEMBER 2011, 62, 6, 501—517 doi: 10.2478/v10096-011-0036-5

Dasycladalean green algae and some problematic algae from

the Upper Triassic of the Nayband Formation (northeast Iran)

BABA SENOWBARI-DARYAN

, KOOROSH RASHIDI

and BEHNAM SABERZADEH

Geozentrum Nordbayern, University Erlangen-Nürnberg, Loewenichstrasse 28, 91054 Erlangen, Germany; basendar@pal.uni-erlangen.de

Payam-e Noor University (PNU), Ardakan/Yazd, Iran; Koo.rashidi@gmail.com

Islamic Azad University, Zarand Branch, Iran; Behnamsaber@yahoo.com

(Manuscript received January 17, 2011; accepted in revised form March 17, 2011)

Abstract: This paper describes the dasycladales green algae from two sections of the Rhaetian Howz-e Khan Member
of the Nayband Formation, northwest of the Dig-e Rostam motorway service area (south of the type locality of the
Formation near the town Naybandan). Both sections are composed of bedded fine-grained limestones containing partly
abundant dasycladales algae associated with foraminifers, which are mainly aulotortid types. Additionally scattered
samples were collected from several beds of the Howz-e Khan Member in this area. The following dasycladalean taxa
are described: Chinianella carpatica (Bystrický), Griphoporella curvata (Gümbel), Griphoporella lutensis nov. sp.,
some undetermined dasycladacean taxa, problematic algae like Lithocodium aggregatum Elliott, Bacinella irregularis
Radoicic, and Thaumatoporella parvovesiculifera (Raineri). While Chinianella carpatica is not numerous and the other
described algae are rare, Griphoporella curvata is extremely abundant in the investigated material. This paper describes
Ch. carpatica for the first time from the Triassic of Iran and also includes a discussion of the strong variability of
G. curvata. Additionally we include an informal description of a problematic fossil (animal: shell fragment?; plant: alga?).

Key words: Triassic, Iran, Nayband Formation, Green Algae, Dasycladales, Chinianella, Griphoporella, Gyroporella,
Lithocodium, Bacinella.

Introduction

Dasycladalean  green  algae  are  an  abundant  fossil  group  in
shallow-water carbonates, particularly in bedded limestones
of Upper Paleozoic and Lower Mesozoic deposits. They are
known from numerous localities around the world, especially
from the Tethyan Realm. General works about the environ-
mental  distribution  and  diversity  of  Permian,  Triassic,  and
Jurassic  dasycladales  are  given  by  Flügel  (1975,  1979,
1985), and the stratigraphical range of Triassic dasycladales
were  carried  out  by  Ott  (1972),  Flügel  (1991),  and  Bucur
(1999). A general work with synonymy-lists of all species of
Permian and Triassic dasycladaleans is published by Granier
& Grgasovic (2000).

Red algae (solenoporaceans) occur in the bio-construc-

tions of the Nayband Formation and are not described in this
paper.  Triassic  dasycladales  are  scarcely  known  from  Iran,
but they are relatively abundant in the bedded limestones of
some  Nayband  Formation  localities.  Senowbari-Daryan  &
Hamedani (2000) described the species of Diplopora, Gyro-
porella, and Acicullela from the Norian part of the Nayband
Formation (central Iran). The genus Acicullela is also report-
ed  from  the  Triassic  of  the  Zagros  Mountains  in  southwest
Iran  (Golestaneh  1979).  Foraminifers,  dasycladales,  in-
cluding  Poikiloporella  duplicata,  Clypeina  besici,  and
Teutloporella  herculea  and  some  problematic  fossils  were
described by Senowbari-Daryan (2003) from the Carnian of
the Nayband Mount, not far from the localities of dasyclad-
ales described in this paper. The problematic fossil Probolo-
cuspis  espahkensis,  described  by  Brönniman  et  al.  (1974)
was identified in a section near the town of Espahk (south of

Tabas) and interpreted as dasycladales by Senowbari-Daryan
& Majidifard (2003). Furthermore, P. espahkensis and an-
other species of the genus – P. aculeate Nittel – are known
from the Wetterstein limestones (Ladinian—Carnian) of the
Northern Calcareous Alps, Austria (Nittel 2006).

Localities

The dasycladales, described in this paper are from two

bedded  limestone  sections  of  the  Rhaetian  Howz-e  Khan
Member of the Nayband Formation. Both localities are situ-
ated  in  southwest  area  of  the  town  of  Naybandan,  a  small
town  east  of  the  type  locality  of  the  Nayband  Formation
south of the Mount Nayband. Additionally scattered samples
were collected from locality 3 in Fig. 1. The field work was
done by the last two authors.

The first sampled section is located about 7 km southwest of

the  town  of  Naybandan  (Fig. 1).  The  locality  can  only  be
reached  on  foot  or  by  motorcycle.  Approximately  160  rock-
pieces were sampled from the  ~ 246 m thick section of grey,
bedded limestones. The lower part of the section, indicated as
locality 1  in  Fig. 1  (N  32°18

’817”; E 57°27’213”) is com-

posed  mainly  of  thick-bedded  limestones  without  reef  build-
ing organisms. In the upper part of the section some sponges
and corals occur. Thin sections, numbered “H” in plate expla-
nations are derived from this locality. The second section is
located  about  15 km  southwest  of  the  town  of  Naybandan,
about  8 km  southwest  of  the  first  section  (Fig. 1)
(N  32°15

’917”; E 57°24’942”). The lowermost part of the

section is sandstone beds, continuing into carbonate beds of

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GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2011, 62, 6, 501—517

the Rhaetian Howz-e Khan Member. 76 rock-pieces were col-
lected from this locality and are numbered “khk” in plate ex-
planations. All other thin sections are taken from the different
beds of the valley, lying north of the section 1 locality, indi-
cated with a thick line numbered “3” in Fig. 1. Thin sections,
marked “n” and “J” were collected from different area of the
Howz-e Khan Member in valley between the localities 2 and 3
in Fig. 1. The investigated thin sections are deposited in the
“Staatssammlung für Paläontologie und historische Geologie
in München, Inventar-Nr: BSPG 2010 XIII 1-48”.

Systematic paleontology

Family: Triploporellaceae Pia, 1920; emend. LoDuca, 1977

D e f i n i t i o n : “Thallus with euspondyle ramifications;

productive bodies endosporate or cladosporate” (LoDuca
1977: p. 943).

Tribe: Triploporelleae (Pia, 1920), Bassoullet et al., 1979
Genus: Chinianella Ott (not 1967), ex Granier & Deloffre,

1993: not 1994 as given in Granier & Grgasovic, 2000

R e m a r k s : There is some confusion about the name of the

genera Chinianella and Heteroporella Praturlon (1967). Ott
(1967: p. 215; date of the journal 15

December 1967) intro-

duced  the  genus  name  Chinianella  with  type  species  Cylin-
droporella  ellenbergeri  Lebouché  &  Lemoine  (1963)  and
added the two further species, Ch. crosi Ott and Ch. zankli Ott
to  this  genus.  Ott  (1968:  date  of  the  journal  15

December

1968) emended the genus Heteroporella Praturlon and attrib-
uted the species Ch. crosi and Ch. zankli to this genus. Granier
&  Deloffre  (1993:  p. 26;  see  also  Granier  et  al.  1994  and
Granier  &  Grgasocic  2000)  re-introduced  the  genus  name
Chinianella Ott with type species Ch. ellenbergeri and also in-
cluded the species Heteroporella carpatica Bystrický (date of
the publication and reprint is 1967, but erronneous reference
as  cited  by  Granier  &  Deloffre  1993  is  1968;  Granier  et  al.
1994;  Granier  &  Grgasovic  2000)  to  this  genus.  Because
Chinianella zankli was attributed to a non valid genus at that
time by Ott (1967), therefore this species was also not valid.
Consequently  Granier  &  Deloffre  (1993),  Granier  et  al.
(1994),  and  Granier  &  Grgasovic  (2000)  correctly  synony-
mized Chinianella zankli Ott with Ch. carpatica (Bystrický).
The  complete  synonymy-list  of  Chinianella  carpatica
(Bystrický) is given by Granier & Grgasovic (2000).

Chinianella carpatica (Bystrický), 1967

(date not given 1967 on reprint)

(Fig. 2A—I, Fig. 3)

*1967 Heteroporella carpatica sp. nov. – Bystrický, p. 302, pl. 15,

fig. 1—5, pl. 16, fig. 4

*1967 Chinianella zankli n. sp. – Ott, p. 219, pl. 13, Fig. 2—3, text

fig. 5/1—14

1968 Heteroporella zankli (Ott) – Ott, p. 258 (without illustrations)
2000 Chinianella carpatica (Bystrický) – Granier & Grgasovic, p. 20

(without illustration, complete synonymy-list)

M a t e r i a l : Several specimens (for thin section numbers of

illustrated material see explanations of plates).

D e s c r i p t i o n : Cylindrical specimens of this alga with

regularly arranged sterile and fertile whorls. The outer diame-
ter  (D)  of  the  thallus  varies  between  1.44 mm  and  2.12 mm,
inner diameter (d) between 0.32 mm and 0.52 mm. Diameter
of all Iranian specimens corresponds the diameter of Ch. car-
patica  (Bystrický  1967)  from  the  Carpathians,  but  is  moder-
ately  smaller  than  those  data  given  by  Ott  (1967)  for
specimens from the Alps.

Groups of five—six sterile laterals end on the thallus surface

(Fig. 2H). Their number is given as possibly 4 by Ott (1967).

Fig. 1. Geographical position of localities southwest of the small
town of Naybandan, from which the algae of the Howz-e Khan
Member (Rhaetian) of the Nayband Formation are described. (Part
of geological map of Naybandan, Kluyver et al. 1983.)

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The number of laterals of sterile whorls is given 5—7 by
Bystrický (1967) in sections, but maybe up to 20 laterals ex-
tending to the surface. The diameter of fertile laterals varies
between 0.32 mm 0.60 mm. Both, the diameter of sterile and
fertile laterals on the thallus surface are with 0.040—0.080 mm
almost the same. In our material, there are 6 sporangia in each
whorl, but Ott (1967) gives their number as 7—8. The biomet-
rical data of Iranian specimens of Ch. carpatica (Bystrický) is
given in Table 1.

O c c u r r e n c e : The geographical distribution and the

stratigraphical range of Ch. carpatica are given by Granier &
Grgasovic (2000). The alga is described here for the first time
from the Upper Triassic (Rhaetian) of Iran.

Genus: Griphoporella Pia, 1915

T y p e s p e c i e s : Gyroporella curvata Gümbel, 1872.
R e m a r k s : The attribution of Griphoporella within the

dasycladales  is  controversial.  Pia  (1915,  1920)  did  not  place
Griphoporella  in  any  family.  Following  Deloffre  (1988),
Berger & Kaever (1992) attributed Griphoporella to the fami-
ly Seletonellaceae (tribe: Mastoporeae Pia, 1920). Barattolo et
al.  (1993)  discussed  the  differences  between  Griphoporella

and other similar looking dasycladales attributing Gripho-
porella to the family Dasycladaceae, tribe Salpingoporelleae.
The family Triploporellaceae was emended by LoDuca (1977)
and this systematic is followed here.

Griphoporella curvata (Gümbel, 1872), Pia, 1915

(Fig. 4A—G, Fig. 5A—L, Fig. 6A—G, Fig. 7A—K, Fig. 8, Fig. 12)

*1872 Gyroporella curvata n. sp. – Gümbel pl. D.IV, fig. 2a—d

1915 Griphoporella curvata (Gümbel) – Pia, pl. 1, fig. 11
1988 Griphoporella curvata (Gümbel) – Sartorio & Venturini, p. 56/2
1988 Macroporella retica Zanin-Buri – Sartorio & Venturini, p. 56/3
1993 Griphoporella curvata (Gümbel) – Barattolo, De Castro &

Parente, p. 26, pl. 1—7

1997 Griphoporella curvata (Gümbel 1872) – Grgasovic, pl. 2, fig. 8,

13 (selected syn.).

2000 Griphoporella curvata (Gümbel) – Granier & Grgasovic, p. 67—70

(com. syn.)

2000 Griphoporella curvata (Gümbel, 1872) Pia, 1915. – Senowbari-

Daryan & Hamedani, p. 102, pl. 1, fig. 1, 2A, 4, 6, 7, pl. 2,
fig. 1B, 2 B, 3B, pl. 3, fig. 2B, 4, pl. 4, fig. 9

M a t e r i a l : Numerous specimens. We have illustrated nu-

merous specimens cut in different directions of the thallus to
show the strong variability of this alga.

D e s c r i p t i o n : The simple calcareous thallus of this alga is

straight or moderately curved and slightly club shaped, reach-
ing outer diameters between 2.6 mm and 4.0 mm, the inner
diameter between 1.8 mm and 3.2 mm. Two specimens with
diameters of 7.5 (Fig. 6A: type 2) and 10 mm (Fig. 12F/A) are
extremely large.

The very wide axial stem is surrounded by a thin wall com-

pared with the diameter of the alga. The thallus wall contains
wide laterals arranged in alternating verticils (whorls), recog-
nizable in sections oblique to the wall. The laterals are proxi-
mally  narrow,  becoming  distally  wider  (phloiophore).  They
are  always  closed  by  a  thin  and  convex  wall.  In  the  convex
wall of some specimens very small pores are perceptible, but
unverifiable. The partial lack of the convex wall in some spec-
imens is interpreted as a possibly result of post-mortem weath-
ering.  The  wall  between  the  individual  laterals  is  proximally
relatively  thick,  becoming  thinner  in  the  middle  part  of  the
walls.  Because  of  the  relatively  thin  wall  of  the  large  thalli,
most specimens are broken and the thallus fragments are ex-
tremely abundant in investigated thin sections.

We distinguished two types (based on variability?) of the

thalli with moderately different biometrical data. Type 1 with

Fig. 3. Chinianella carpatica (Bystrický). Two longitudinal sec-
tions and a cross section exhibiting the fertile and sterile laterals, as
well as the fine, distal laterals (drawn from specimens illustrated in
Fig. 2A—B and E).

Section

DSF

d/D

KHK3

1.6

0.4

0.04–0.06

0.04

0.32

0.04–0.05

0.25

cross

KHK19

1.6

0.32

0.12

0.04

0.48

0.04

0.2

cross

KHK19

1.68

0.44

0.08–0.1

0.04

0.6

0.04–0.05

0.26

cross

KHK19

2.12 0.48

0.08

0.04

0.48–0.5

0.06–0.08 0.22 oblique

KHK19

1.8

0.52

0.08

0.04–0.06

0.6

0.06–0.08

0.28

cross

KHK19-2

1.44

0.44

0.04–0.06

0.06

0.44–0.6

0.04–0.06

0.3

oblique

KHK19-3

–

0.4

0.08

–

tangential

Table 1: Biometrical data of Chinianella carpatica (Bystrický) from the Rhaetian Howz-e Khan Member of the Nayband Formation, northeast
Iran. D – outer diameter of the thallus, d – diameter of the central stem, T – thickness of the wall around central stem, DS – diameter of
the sterile laterals at the base, DF – diameter of sporangia, DSF – diameter of sterile and fertile laterals ending on the surface of thallus,
d/D – ratio of central stem/outer diameter of the thallus, PS – position of section. All measurements are in mm (d/D in %).

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pear-shaped  laterals  (Fig. 4A—G,  Fig. 5A—L,  Fig. 6C,E,
Fig. 7A—B,E,G—J,  Fig. 12F/A);  type 2  with  slender  laterals
(Fig. 6A—B,D,F—G,  Fig. 7C—D,F,K,  Fig. 12F/B).  Biometric
data of the thallus of both types are separately summarized in
Table 2.  Illustrations  in  Fig. 8  exhibit  the  variability  of  the
alga in the investigated material. Fig. 9 shows the ratio of the
central stem the thallus diameter (d/D) of type 1 (triangle) and
type 2  (rectangle)  and  Fig. 10  the  ratio  of  the  thallus  wall  to
the outer diameter (t/D) of type 1 (triangle) and type 2 (rectan-
gle) of Griphoporella curvata.

D i s c u s s i o n : On the basis of numerous specimens from

the  type  locality  Barattolo  et  al.  (1993)  re-described  Gripho-
porella  curvata  (Gümbel  1872)  designating  the  neotype  and
emending the species and genus diagnosis. A complete syno-
nymy-list  of  the  species  is  given  by  these  authors  and  by
Granier  &  Grgasovic  (2000).  Barattolo  et  al.  (1993)  synony-
mized  Macroporella  retica  described  by  Zanin  Buri  (1965)
from  the  Rhaetian  of  “Prealpi  Lombardi”,  north  Italia.  Con-
firming the opinion of Barattolo et al. (1993) Pugliese (1995)
re-described  Macroporella  retica  from  the  Upper  Triassic  of
the  type  locality  near  the  town  of  Aviatic  (Lombardy,  Italy).
Also  those  specimens,  illustrated  as  Macroporella  retica  by
Ciarapica et al. (1987) were synonymized with Griphoporella
curvata (Gümbel) by Barattolo et al. (1993). The holotype of
Macroporella  retica,  illustrated  in  pl. 44/a  by  Zanin  Buri
(1965)  is  different  from  several  specimens  of  Griphoporella

Fig. 8. Griphoporella curvata (Gümbel). Fig. A—E (type 1) and F—H
(type 2). Drawn from specimens illustrated in Fig. 4A—B, Fig. 5A—C,
Fig. 6B and Fig. 7C and F. Scale in all Figs. is 1 mm.

Type 1

Section No:

Position

J-39

2.64

1.93

0.4

0.2

0.04

0.03

Cross

J-39

4.00

2.96

0.48

0.12

0.2

0.04

0.12–0.18

Longitud.

J-30

1.60 1.00 0.2–0.21

0.05–0.12

0.12–0.16 0.04 0.12

–

J-1-3

2.44

2.04

0.32

0.16

0.2

0.04

0.08

Cross

J-1-3

4.00

3.20

0.4

0.12–.16

0.2

0.04–0.06

0.06–.08

Cross

J-2-4

2.96 2.40 0.24–.28 0.08 0.16 0.04 0.04 Oblique

J-2-4

2.96 2.48 0.2–0.24 0.12

0.16

0.04 0.04–0.09 Oblique

J-14-2

2.80 2.16

0.32–0.36 0.16 0.24 0.06 0.06

Cross–oblique

J-14-2

3.20 2.48

0.28–0.32 0.16 0.28 0.04 0.08–0.12 Cross

J-18-2

3.40

2.64

0.4

0.12

0.04–0.08

0.16

Oblique

J-2-7

3.08 2.20 0.24 0.08 0.16–0.32

0.16–0.64 0.04

Cross–oblique

J-2-7

2.60

2.00

0.2

0.08–0.12

0.16–0.2

0.16–0.4

0.06–0.08

Oblique

J-2-7

2.60

1.8

0.32–0.4

0.08

0.36

0.16–0.2

0.04

Cross

J-15-1

2.08

1.64

0.24–0.4

0.16

0.2

0.08–0.4

0.04

Oblique

J-9-1

3.56 2.84 0.36 0.08–0.12 0.24 0.16 0.04

Cross

J-2-7

2.60

1.8

0.32–0.4

0.08

0.36

0.16–0.2

0.04

Cross

J-15-1

2.08

1.64

0.24–0.4

0.16

0.2

0.08–0.4

0.04

Oblique

J-9-1

3.56 2.84 0.36 0.08–0.12 0.24 0.16 0.04

Cross

Type 2

Section No:

Position

J-39

4.64

3.92

0.36

0.16

0.08

Cross

J-39

10.00

9.2

0.4–0.42

0.16

0.08

0.04

Cross

J-34

3.84

3.12

0.41

0.12

0.2

0.06

0.04

Oblique

J-34

2.64

2.08

0.4

0.1

0.06

0.08

Cross

J-2-4

4.00

3.36

0.32–0.36

0.16

0.2–0.24

0.04

Oblique

J-18

4.72

3.92

0.32–0.4

0.16

0.16–0.2

0.04–0.06

0.08–0.12

Oblique

J-18

3.00

2.20

0.32

0.16

0.16–0.18

0.04–0.12

0.12

Oblique

J-15-1

3.76

3.00

0.4

0.08–0.16

0.2–0.24

0.4–0.6

0.04

Longitudinal

J-10-1

3.20

2.48

0.4–0.48

0.12

0.12–0.16

0.08–0.12

0.04

Cross

J-10-1

1.64

1.16

0.24–0.28

0.16

0.04

Cross

J-3-2

3.6

3.00

0.28–0.32

0.08–0.12 0.2

0.12–0.16

0.04

Oblique

Table 2: Biometrical data of Griphoporella curvata (Gümbel). The data of both distinguished types are listed separately. D – outer diameter
of the thallus, d – inner diameter of the thallus, T – thickness of the wall, PP – proximal diameter of laterals, PD – distal diameter of laterals,
TB – thickness of the wall between the laterals, TF – thickness of the wall covered distally the laterals. All measurements are in mm.

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GEOLOGICA CARPATHICA, 2011, 62, 6, 501—517

Fig. 9.  Ratio  of  the  central  stem  to  the  thallus  diameter  (d/D)  of
type 1  (triangle)  and  type 2  (rectangle)  of  Griphoporella  curvata
(Gümbel). There are no serious differences between types 1 and  2
discussed in the text. One specimen was observed (see Fig. 12F/A)
with an outer diameter of 10 mm, much larger than all other speci-
mens. Units for all data are in mm.

Fig. 10. Ratio of the thickness of thallus wall to the thallus diameter
(t/D) of type 1 (triangle) and type 2 (rectangle) of Griphoporella
curvata (Gümbel). Units for all data are in mm.

curvata (Gümbel) illustrated by the same author in pl. 63. The
type species of Macroporella retica, illustrated by Zanin Buri
(1965)  in  pl. 44/a,  shows  the  distally  closed  laterals  in  part
like  in  most  specimen  (type 1)  from  Iran,  a  characteristic,
which is not visible in specimens illustrated by Barattolo et
al.  (1993).  The  closed  laterals  are  also  not  visible  in  speci-
mens of Griphoporella curvata (Gümbel) illustrated by other
authors  (e.g.  Flügel  1981:  fig. 10/E;  Pugliese  1995:  pl. 1,
fig. 6; Grgasovic 1997: pl. 2, fig. 8, 13). The laterals of speci-
mens  from  central  Iran,  described  as  Griphoporella  curvata
Senowbari-Daryan & Hamedani (2000) are also distally open
and the shape of the laterals is different. The differences be-
tween our specimens are shown in Table 2 (type 1 and type 2).

In summary, based on the shape of laterals two types of

Griphoporella can be distinguished in the investigated materi-
al. As shown in Table 2, the differences of other specific data
are  diffuse  and  do  not  allow  us  to  describe  them  as  separate
species. Because both types are united in Griphoporella cur-
vata (Gümbel) by previous authors, particularly in the synony-
my-list  of  Barattolo  et  al.  (1993)  and  Granier  &  Grgasovic
(2000)  we  do  not  separate  them  and  describe  both  types  as
Griphoporella curvata (Gümbel).

O c c u r r e n c e : Griphoporella curvata (Gümbel) is known

from the Norian—Rhaetian of numerous localities on the world
(see Grgasovic 1997; Granier & Grgasovic 2000). It was de-
scribed from the Norian of central Iran by Senowbari-Daryan
& Hamedani (2000) and occurs extremely abundantly in the
Rhaetian Howz-e Khan Member of the Nayband Formation in
northeast Iran (this paper).

Griphoporella? sp.

(Fig. 11C,J)

M a t e r i a l : Six specimens, from which only two are illus-

trated.

D e s c r i p t i o n : The two illustrated specimens are united in

one species, but may be different taxa. The outer diameter of
the specimens illustrated in Fig. 11C is 4.6 mm; inner diame-
ter is 2.9 mm. It is surrounded by white incrustation (possibly
spongiomorphids?). The polygonal laterals are about 0.3 mm
in diameter and arranged in two rows.

The specimen, illustrated in Fig. 11J is cut in longitudinal

section reaching an outer diameter of 3.0 mm; inner diameter
2.4 mm. Compared to the first specimen the laterals are circu-
lar or oval.

Griphoporella lutensis nov. sp.

(Fig. 2J—N, Fig. 12G—H, Fig. 13)

2005 Chains of spores of diploporid dasycladales – Fürsich et al., pl. 10,

fig. 8

D e r i v a t i o n o m i n i s : Named from the Lut desert in east

Iran.

H o l o t y p e : Specimen illustrated in Fig. 2K (compare

Fig. 13).

P a r a t y p e s : All specimens in Fig. 2J,L—N, Fig. 12G—H.
L o c u s t y p i c u s : See locality 2 in Fig. 1 (it was also

found in other two localities in Fig. 1).

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Table 3: Biometrical data of Griphoporella lutensis nov. sp. (for abbreviations see Table 2). P – Diameter of laterals. All measurements
are in mm.

S t r a t u m t y p i c u m : Howz-e Khan Member (Rhaetian)

of the Nayband Formation.

D i a g n o s i s : Cylindrical, possibly club-shaped alga with

extremely wide central stem and thin thallus wall with modest
extension of laterals. Single laterals are distributed equally
through the thallus wall. Euspondyl arrangement of the laterals.

M a t e r i a l : At least four, more or less well preserved and

some broken specimens.

D e p o s i t o r y : See chapter localities above.
D e s c r i p t i o n : Compared to the outer or inner diameter of

the thallus, the thallus wall is very thin reaching a thickness of
0.1—0.18 mm.  The  outer  diameter  of  the  thallus  varies  be-
tween  2.02—2.85 mm,  the  inner  diameter  between  1.77 mm
and 2.62 mm. The wall contains numerous single laterals dis-
tributed equally within the wall and oriented vertically to the
wall. The diameter of laterals is about 0.036—0.075 mm. Some
of  laterals  are  proximally  or  distally  closed  (due  to  the  sec-
tion), other are on both ends open being significant of all later-
als. The wall between the laterals is about 0.01—0.03 mm. In
oblique  sections  through  the  thallus  wall  the  arrangement  of
the laterals seems to be of euspondyl type (Fig. 2J—L). The bi-
ometrical data of the thallus and thallus elements of G. luten-
sis nov. sp. are given in Table 3.

D i s c u s s i o n : Senowbari-Daryan & Hamedani (2000) de-

scribed the species Diplopora iranica from the Norian part of
the Nayband Formation, central Iran. Numerous coherent ga-
metangia,  arranged  like  hoses  within  the  central  stem  were
found  within  D.  iranica.  Fragments  of  such  sporangia-hose
were mentioned first as “Mikroproblematikum 1” by Flügel &
Flügel-Kahler  (1963)  and  later  were  described  in  detail  by
Flügel  (1964)  as  “Problematikum 4”.  Such  “cell-aggregates”
were  also  illustrated  from  the  Norian-Rhaetian  carbonates  of
the Northern Calcareous Alps by Sadati (1981) as “Problema-
tikum 4”  or  by  Wurm  (1982)  as  “Griphoprella  curvata”  or
“micritic tubes” in page 274 by the same author. The dimen-
sions  of  the  “cells”  in  “Problematikum 4”  Flügel  and  other
specimens from the Alps correspond to the dimensions of the
sporangia in D. iranica and are identical to our specimens de-
scribed in this paper.

“Problematikum 4” Flügel (1964) and other specimens

from  the  Alps  and  the  sporangia-hoses  in  D.  iranica  appear
dark  micritic  in  transmitted  light.  As  shown  by  Senowbari-
Daryan  &  Hamedani  (2000)  the  crystal  sizes  of  the  wall  are

micro-grains of about 5 µm and the mineralogy of Mg-calcite.
The alga, described above as Griphoporella lutensis nov. sp.
was  never  found  within  the  central  stem  of  other  algae  and
D. iranica was not found in investigated material. G. lutensis
appears light-coloured in transmitted light, indicating the cal-
cite mineralogy of the alga, different from those described by
Flügel  (1964)  and  Senowbari-Daryan  &  Hamedani  (2000).
Further differences are the thickness of the wall between the
individual  laterals  in  Griphoporella  lutensis.  The  wall  be-
tween  the  gametangia  in  D.  iranica  is  much  thinner  than  in
Griphoporella  lutensis.  Also  the  laterals  in  the  new  alga  are
proximally and distally open, the gametangia in D. iranica are
closed at both ends. These differences are against the equality
of this alga and the sporangia-hoses of D. iranica.

Diplopora cf. D. interiecta Fenninger, 1969

(Fig. 11F—G)

1969 Diplopora interiecta n. sp. – Fenninger, p. 24—25, pl. 1, fig. 1—9

M a t e r i a l : Two specimens.
D e s c r i p t i o n : Two specimens of this alga are cut in lon-

gitudinal and oblique sections showing the character and the
branching pattern of the laterals (metaspondyle) at the base
of the thallus wall. The outer diameter of the alga is 1.5—
2.9 mm; the inner diameter 1.1—2.0 mm; diameter of laterals
0.07—0.2 mm.

R e m a r k s : Diplopora is an abundant algal genus in

Triassic  carbonates.  Numerous  specimens  are  known  from
Permian  and  Triassic  strata  (Granier  &  Grgasovic  2000).
Three species of Diplopora are known from the Upper Trias-
sic of Iran: Diplopora interiecta Fenninger (1969), D. iranica
Senowbari-Daryan & Hamedani (2000), and D. phanerospora
Pia (in: Fürsich et al. 2005). The dimensions and other charac-
teristics of our species are similar to that species, described as
D.  interiecta  by  Fenninger  (1969)  from  the  Upper  Triassic
(Norian—Rhaetian)  of  the  Golpaygan  area  (northwest  Iran).
Similar algae to D. interiecta were also reported from the No-
rian—Rhaetian of Austria and Sicily (see synonymy-list of D.
interiecta in Granier & Grgasovic 2000).

Clypeina? sp.

(Fig. 11K)

Section No:

Position of section

J-2-6

2.85

2.62

0.11

0.056–0.075

0.037

Oblique

KHK-14

2.05 1.80 0.1–0.11 0.04 0.01–0.02 Oblique

H-9

2.02

1.77

0.10–0.11

0.049

0.034

Oblique-cross

J-29

2.13

1.92

0.09–0.117

0.04–0.045

0.27

Oblique

J-2-6

–

0.11

0.05

0.037

Broken

J-2

–

0.15

0.056

0.0375

Longitudinal-broken

J-14-2

–

0.18

0.04

0.027

Longitudinal-broken

J-2-6

–

0.109

0.036–0.042

–

Oblique-broken

J-9-2

– –

0.075–0.011

0.03

0.02–0.03

Oblique-broken

J-8-2

–

0.09–0.1

0.03–0.04

0.028

Longitudinal-broken

J-2

– – 0.15

0.03–0.05

0.3

Longitudinal-broken

H-1

– – 0.02–0.05

0.033–0.05

Tangential

J-20

– – 0.11

0.054–0.059

0.01–0.03

Oblique-broken

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M a t e r i a l : One specimen only.
The only cross section of this species is strongly re-crystal-

lized and the detail of the alga is not ascertainable. The outer
diameter of the alga is 1.0 mm; the inner diameter 0.36 mm.

R e m a r k s : Clypeina is an abundant algal genus in Creta-

ceous deposits. From Triassic only the species Clypeina besici
is described by Pantic (1965) from the Carnian of the Dinarids
(Montenegro). Questionable Clypeina from Upper Triassic
(Norian—Rhaetian) is reported by Flügel (1979: pl. 2, fig. 9)
and by Senowbari-Daryan (1980: pl. 14, fig. 6).

Similar algae to Clypeina? sp., described above are illustrat-

ed by Bodrogi et al. (1993) as Clypeina sp. from the Lower
Cretaceous of Hungary. Nittel (2006) also illustrated from the
Middle Triassic of the Northern Calcareous Alps, Austria a
similar section identified as bryozoans. Most probably the Ira-
nian species is a new algal taxon.

Dasycladales gen. et sp. indet 1

(Fig. 11H)

M a t e r i a l : One specimen only.
D e s c r i p t i o n : The only one specimen is cut in longitudi-

nal section. The U-shaped thallus exhibits a thin wall. The dia-
meter of the thallus is about 1.6 mm. The outer surface
appears to be annulated, caused by the narrow and short later-
als in relatively wide distances.

Dasycladales gen. et sp. indet 2

(Fig. 11B, I)

M a t e r i a l : Two specimens only.
D e s c r i p t i o n : From this alga only two incomplete speci-

mens in cross section are available. The description is based
mainly on specimen illustrated in Fig. 11B.

The thick-walled thallus exhibits a thin wall at the base of

laterals.  Laterals  become  continuously  thinner  to  the  thallus
periphery,  resembling  representatives  of  the  genus  Physopo-
rella.  The  outer  diameter  of  alga  is  1.65 mm;  inner  diameter
0.90 mm; wall thickness 0.82 mm.

R e m a r k s : Physoporella is an abundant algal genus in the

Permian  and  in  Middle  Triassic,  particularly  in  the  Anisian.
Only  two  species  of  Physoporella  are  known  from  the
Norian—Rhaetian:  Physoporella  jomdaensis  Flügel  &  Mu
(1982)  from  eastern  Tibet  and  Ph.  zamparelliae  Parente  &
Climaco  (1999)  from  south  Italy.  Most  probably  the  Iranian
species is a new algal taxon.

Dasycladalean algae

(Fig. 11A,D—E)

In addition to the described taxa other dasycladales occur

very rare, illustrated in Fig. 11A,D—E. Specimen in A is a very
small alga exhibiting the laterals corresponding to the genus
Griphoporella. The specimen in D seems to be a representa-
tive of the genus Macroporella. The specimen in Fig. 11E is
an indet one.

Division: Chlorophyta Kützing, 1843

Family: Incertae sedis

Genus: Lithocodium Elliott, 1956

E m e n d e d   d i a g n o s i s : See Schlagintweit et al. (2010).
T y p e   s p e c i e s : Lithocodium aggregatum Elliott, 1956.
D i s c u s s i o n :  Elliott  (1956)  assigned  Lithocodium  aggre-

gatum  to  codiacean  algae.  Lithocodium  aggregatum  was  at-
tributed  to  luftusiid  foraminifer  by  Schmid  &  Leinfelder
(1996).  Banner  et  al.  (1990)  synonymized  Bacinella  irregu-
laris  Radoicic  (1959)  and  Pseudolithocodium  carpaticum
Mišík  (1979)  with  Lithocodium,  attributing  these  to  the  new
subfamily  Lithocodioidea  within  the  family  Codiaceae
(Siphonales,  Clorophyceae).  For  Bacinella?  sterni  Radoicic
(1972)  the  new  genus  Radoicicinellopis  was  established  by
Banner et al. (1990). Cherchi & Schroeder (2006) favour, that
Lithocodium is a calcified cyanobacterium rather than a fora-
minifer.  Recently  Schlagintweit  et  al.  (2010)  discussed  the
systematic position of Lithocodium attributing it to “filamen-
tous-septate  heterotrichale  ulvophycean  alga”.  Assigning  to
“endolithic ulvophycean alga” these authors separate Bacinella
from  Lithocodium,  which  was  synonymized  by  Banner  et  al.
(1990). With separation of Lithocodium and Bacinella the sys-
tematic classification proposed by Schlagintweit et al. (2010)
is followed here to describe the two following organisms.

?Lithocodium aggregatum Elliott, 1956

(Fig. 11N—O)

R e m a r k s : According to Schlagintweit et al. (2010) the

Triassic organisms, known as “Lithocodium aggregatum”
Elliott is excluded from Lithocodium aggregatum Elliott
(1956) by these authors. Here only the synonymy of Triassic
“Lithocodium aggregatum” is listed below. For the synonymy
of Jurassic—Cretaceous Lithocodium see Schlagintweit et al.
(2010).

Selected synonymy

*1959 Problematicum A – Ohlen, p.73, pl.10, fig.1, pl.17, fig. 3

1979 Lithocodium – Senowbari-Daryan & Schäfer, pl. 1, fig. 8
1981 Lithocodium aggregatum Elliott – Sadati, p. 206, pl. 59, fig. 8
1984 Lithocodium aggregatum Elliott – Senowbari-Daryan, p. 33—34,

pl. 10, fig. 8 (cum. syn.)

D e s c r i p t i o n : Etching the substrate (bio- or lithoclast)

Lithocodium grows as encrustation or as aggregates between
other organisms. It is characterized by hollow cavities covered
by a layer with thin-walled alveoli. A detailed description of
Lithocodium and discussion about its systematic position are
given by Banner et al. (1990), Schmid & Leinfelder (1995,
1996), Cherchi & Schroeder (2006), and by Schlagintweit et
al. (2010).

R e m a r k s : In Triassic deposits, as in Jurassic and Creta-

ceous, Lithocodium aggregatum is usually associated with
Bacinella irregularis. In the investigated material we did not
find them in association. Lithocodium with association of Ba-
cinella irregularis is, however, reported by Fürsich et al.
(2005) from the Nayband Formation of the same area.

Order: Ulotrichales

Family: Incertae sedis

Genus: Bacinella Radoicic, 1959

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T y p e s p e c i e s : Bacinella irregularis Radoicic, 1959.

Bacinella irregularis Radoicic, 1959

(Fig. 11L,P)

Selected synonymy
*1959 Bacinella irregularis n. sp. – Radoicic, p. 89, pl. 3, fig. 1—2

1984 Bacinella irregularis Radoicic – Senowbari-Daryan, p. 38, pl. 9,

fig. 2 (with synonymy)

2005 Bacinella irregularis Radoicic – Fürsich et al., pl. 11, fig. 3

D e s c r i p t i o n : The aggregates of Bacinella occur in mic-

rite sediments forming bubble-like cavities with thin and mic-
rite walls. The outer walls and the tabulae-like elements
within the cavities are, like Thaumatoporella partly perforated
(Fig. 11L). Bacinella do not occur together with Lithocodium
in the investigated material.

R e m a r k s : Occurring in association, the separation of Li-

thocodium aggregatum and Bacinella irregularis is in most
cases not possible. They were also described as a single organ-
ism by some authors or as separate genus/species by others
(for more information see Banner et al. 1990; Schlagintweit et
al. 2010). Bacinella and Lithocodium were not found in asso-
ciation in the investigated material. Both organisms were also
reported by Fürsich et al. (2005) from the Nayband Formation.

Incertae sedis

Genus: Thaumatoporella Pia, 1927

Type species: Gyroporella parvovesiculifera Raineri, 1922.

Thaumatoporella parvovesiculifera (Raineri), 1922

(Fig. 11M)

Selected synonymy
*1922 Gyroporella parvovesiculifera n. sp. – Raineri, p. 83, pl. 13,

fig. 17—18

1927 Thaumatoporella parvovesiculifera (Raineri) – Pia, p. 69
1984 Thaumatoporella parvovesiculifera (Raineri) – Senowbari-

Daryan, p. 35—36, pl. 7, fig. 1 (cum. syn.)

2002 Thaumatoporella parvovesiculifera (Raineri) – De Castro, text-

fig. 2, pl. 1, fig. 1—11

D e s c r i p t i o n : Thaumatoporella parvovesiculifera oc-

curs, like the preceding species in micritic sediments or be-
tween the reef organisms. The fossil is characterized by well
perforated sheets of aggregates. It is a rare fossil in the inves-
tigated material.

R e m a r k s : The systematic position of Thaumatoporella

parvovesiculifera is disputed. It was attributed to dasycladales
by  (Raineri  1922),  solenoporacean  (Elliott  1957),  codiaceans
(Johnson  1966),  possibly  sponges  (Flügel  1983).  De  Castro
(1990)  compared  Thaumatoporella  with  some  volvocacean
(planktonic  green  algae)  and  with  rivulariacean  cyanophyta.
He attributed Thaumatoporella to the new family Thaumato-
porellaceae  and  the  new  order  Thaumatoporellales  (see  also
De Castro 2002).

Problematicum 1 (shell fragment?, alga?)

(Fig. 12A—E)

M a t e r i a l : Several fragments.
D e s c r i p t i o n : The specimen illustrated in Fig. 12A

shows  7  cohered  elements  (“columns”)  in  calcite  (primary
aragonite?)  preservation.  Other  specimens  illustrated  in
Fig. 12 show only fragments. Each element exhibits numer-
ous spine-like to     circular peripheral elements with regularly
arrangement.  Some  of  these  peripheral  elements  are  almost
closed, building circular cavities of about 0.085 mm in dia-
meter (Fig. 12D). The specimen illustrated in Fig. 12D also
shows  that  these  peripheral  elements  originate  from  a  thin
wall (0.04 mm thick) around an axial tube with a diameter of
0.085 mm.  Other  specimens  show  that  these  elements  are
connected laterally with others. The thickness of the individual
“columns”  is  about  1.85 mm;  their  distance  varies  between
0.11 mm  and  0.3 mm.  The  distance  between  peripheral  ele-
ments  around  the  “columns”  varies  between  0.7 mm  and
0.8 mm.

D i s c u s s i o n : The specimen with an axial stem and cyst-

like  elements  around  it,  illustrated  in  Fig. 12D  shows  that
this  fossil  resembles  dasycladales.  The  lack  of  the  pores  or
laterals of the wall around the axial cavity does not confirm
its  interpretation  as  dasycladales.  Other  specimens,  particu-
larly the specimen illustrated in Fig. 12A also fail to support
this interpretation.

The cohered columns with laterally connected elements

support the interpretation of this fossil as sections through the
ripped  shell  of  juvenile  bivalves,  brachiopods  or  ostracods.
The sections could be oriented parallel to fragments of shell.
The axial cavity in Fig. 12C, however, does not support the in-
terpretation of this fossil as shell fragments. Another interpre-
tation  could  be  the  aptychus  of  cephalopods.  We  describe  it
here  as  “Problematicum 1”,  because  ambiguous  characteris-
tics make the exact interpretation of this fossil impossible.

Fig. 13. Griphoporella lutensis nov. sp. (Holotype). The specimen
shows extremely thin wall, compared with the thick axial stem
(drawn from Fig. 2K).

516

SENOWBARI-DARYAN, RASHIDI and SABERZADEH

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GEOLOGICA CARPATHICA, 2011, 62, 6, 501—517

Acknowledgments:  The  fieldwork  was  conducted  by
Koorosh Rashidi and Behnam Saberzadeh as part of a master
degree from the “Islamic Azad University of Zarand Branch,
Iran”. Limited financial support came from the “Sonderfonds
für  wissenschaftliche  Arbeiten  an  der  Universität  Erlangen-
Nürnberg” to B. Senowbari-Daryan. We thank the Ministry of
Environment Protection in Teheran and Yazd, Mr. H. Akbari
for permission to conduct fieldwork. Our thanks are addressed
to  Rowan  Martinale  (Los  Angeles)  for  improvements  of  the
English. We thank I.I. Bucur (Cluj-Napoca) and F. Barattolo
(Napoli), whose very helpful comments as the journal review-
ers improved the manuscript.

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