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Two Permian sphinctozoan sponges from the

  Shotori Mountains (eastern Iran)








Institute of Paleontology, University of Erlangen-Nürnberg, Loewenichstr. 28, 91054 Erlangen, Germany;


University of Esfahan, Faculty of Sciences, Esfahan, Iran;;

(Manuscript received November 7, 2005; accepted in revised form March 16, 2006)

Abstract: Two sphinctozoan sponge taxa, Shotorithalamia tubulara nov. gen., nov. sp. and Amblysiphonella cf. A.
vesiculosa vesiculosa (de Koninck), are described from the Permian deposits of the Jamal Formation, exposed in Bagh-e
Vang Mountain, and from a locality near the town of Deh-e Mohammad, both in the Shotori Mountains, north of Tabas,
eastern Iran. The new genus Shotorithalamia is characterized by having ring-chambers – like Amblysiphonella – and
also by possession of multibranched tubes that extended from the imperforate exo- and endowalls into the chamber
interiors. The validity of families Amphorithalamiidae Senowbari-Daryan & Rigby and Girtyocoeliidae Finks & Rigby
is discussed.

Key words: Permian, Iran, Shotori Mountains, Bagh-e Vang, Taxonomy, Porifera, Sphinctozoa.


The Permian deposits, called the Jamal Formation, crop
out in several localities in the Shotori Mountains, in east-
ern, and also in central Iran (Stöckline et al. 1965). The
section of the Jamal Formation crops out on the western
flank of the Kuh-e Bagh-e Vang, located about 45 km NW
of Tabas, approximately 5 km south of the town of Shir-
gesht (geol. map Shirgest, 1 : 100,000 by Ruttner et al.
1968, see Fig. 1). Here the Jamal Formation reaches thick-
nesses of 193 m (Ruttner et al. 1968) to 300 m (Leven &
Vaziri Moghaddam 2004). The lower part (about 60 m
thick) of the section is composed of differently coloured
limestones and marls, with some olistoliths (up to 3 m in
diameter). This part was identified as an independent for-
mation by Partoazar (1995), who named it the Bagh-e
Vang Member, of Bolorian age (for more information see
Leven & Vaziri Moghaddam 2004; Senowbari-Daryan et
al. 2005).

Sponges, described by Senowbari-Daryan et al. (2005)

from the Bagh-e Vang locality, were collected mainly
from reefal beds of the Bagh-e Vang Member. The fol-
lowing taxa have been described by the authors:

  Amblysiphonella rigbyi Senowbari-Daryan,

Rashidi & Hamedani,  Baghevangia minima Senowbari-
Daryan,  Rashidi & Hamedani, Shotorispongia permica
Senowbari-Daryan, Rashidi & Hamedani, Shotorispon-
gia biserialis Senowbari-Daryan, Rashidi & Hamedani,
Thaumastocoelia? cf. Th. irregularis Weidlich & Se-
nowbari-Daryan, Sollasia ostiolata Steinmann, Sollasia
cylindrica Senowbari-Daryan, Rashidi & Hamedani, Gir-
tyocoelia beedei (Girty), and Lemonea cylindrica (Girty).

  Preperonidella recta grossa (Wu), Bicoelia spi-

nosa Senowbari-Daryan, Rashidi & Hamedani, Hep-
tatubispongia symmetrica Rigby & Senowbari-Daryan,

Maeandrostia kansasensis Girty, Maeandrostia radiata Se-
nowbari-Daryan, Rashidi & Hamedani, and Solutosspongia
tabasensis Senowbari-Daryan, Rashidi & Hamedani.

Shotorithalamia tubulara nov. gen., nov. sp., de-

scribed in this paper, was also collected from reefal beds
of the Bagh-e Vang Member (Fig. 1, locality 1) by one of
the authors (K. Rashidi).

Amblysiphonella cf. A. vesiculosa vesiculosa was

found in an outcop (33º 59´ 46.6´´ N and 57º 01´ 46´´ E)
located about 4 km northeast of the town of Deh-e Mo-
hammad, approximately 68 km north of Tabas, and
23 km from the Bagh-e Vang locality (Fig. 1, locality 2).
The age of the section, exposed in this locality corre-
sponds to the Bagh-e Vang section (Lower Permian,
Bolorian). The systematic classification, proposed by
Finks & Rigby (in Finks et al. 2004) is used in this paper.

Systematic Paleontology

Phyllum:  Porifera Grant, 1872

Class:  Demospongea Sollas, 1875

Subclass:  Ceractinomorpha Levi, 1973

Order:  Agelasida Verrill, 1907

Family:  Sebargasiidae Laubenfels, 1955

Subfamily:  Sebargasiinae Senowbari-Daryan, 1990

Genus:  Amblysiphonella Steinmann, 1882

Amblysiphonella  cf. vesiculosa vesiculosa (de Koninck)


(Fig. 2.4—5)

*1863  Orthoceras vesiculosum n. sp. – de Koninck, p. 15
   1887   Amblysiphonella vesiculosa (de Koninck) – Waagen &
        Wentzel, p. 973—974, pl. CXXII, fig. 1
?1970  Amblysiphonella vesiculosa – Zhuravleva, fig. 3a—3b

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M a t e r i a l :

 Only one specimen from the Lower Permian

(Bolorian) Jamal Formation of Deh-e Mohammad locality
(Fig. 1/2).

O c c u r r e n c e :

 See synonymy.

D e s c r i p t i o n :

 The available part of the broken

sponge has a length of 50 mm. Its true length cannot be
given, because only a portion is preserved. The sponge
was cut in several pieces and later glued together. It has an
outer diameter of 50 mm and is composed of several cres-
cent-like ring-chambers. Because of the overlap of periph-
eral parts of preceding chambers by the younger chamber
walls, outer segmentation is not recognizable from the ex-
terior view, but internal chambered construction is clearly
evident in sections of the specimen. The chambers reach
heights of about 6—8 mm. The exowalls are doubled lay-
ered, reaching thicknesses of up to 3 mm, and the inter-
walls are about 1 mm thick. An axial spongocoel of
retrosiphonate type is 10 mm in diameter. Chamber walls
are pierced by numerous pores 0.3—0.4 mm in diameter.
Chamber interiors are filled by vesiculae, which appear
circular in longitudinal and tangential sections.

R e m a r k s :

  Amblysiphonella is a long-lasting genus

occurring from the Ordovician and Carboniferous to the
Late Triassic. To date, the genus has not been reported
from the Silurian and Devonian. More than 60 species are
known (Senowbari-Daryan & Garcia-Bellido 2002). The

majority of these have been described from Permian de-

According to the sponge diameter, the Iranian species is

comparable with A. vesiculosa vesiculosa, erroneously de-
scribed as a cephalopod Orthoceras vesiculosum by de
Koninck (1863), and later redescribed by Waagen &
Wentzel (1887) as a sponge from the Permian of the Salt
Range (Pakistan). Later, the species was described by
Zhuravleva (1970) from the former Soviet Union without
giving the exact stratigraphic level and locality of the oc-
currence. The Iranian specimen differs from other known
specimens described by de Koninck and by Zhuravleva
(diameter smaller than 30 mm) by having a larger diameter
(50 mm) and by different shapes of the chambers. Cham-
bers of the Iranian specimen, however, have almost the
same height as in A. vesiculosa minima described by
Zhang (1983), and Fan & Zhang (1985) from the Late Per-
mian of Lichuan, West Hubei, China. We are not sure
about the assignment of the Iranian specimen to the spe-
cies from Pakistan or from China, and have thus described
it as A.  cf. vesiculosa vesiculosa (de Koninck).

Amblysiphonella is an abundant sponge genus in Per-

mian deposits in eastern and central Iran. The species A.
iranica was described from the Permian at a locality near
the La Kaftari Mountain (south of Bagher-Abad, north of
Esfahan by Senowbari-Daryan & Hamedani 2002). Other
sponges from this locality were reported by Rigby, Senow-
bari-Daryan & Hamedani (2005).

A. rigbyi Senowbari-Daryan, Rashidi & Hamedani (2005)

is the most abundant sphinctozoan sponge in the Permian
Jamal Formation of the Bagh-e Vang locality. A. cf. vesicu-
losa vesiculosa, described in this paper, is the third species
of the genus Amblysiphonella occurring in Permian depos-
its in Iran, along with A. iranica and A. rigbyi.

Discussion about the validity of the families

Amphorithalamiidae and Girtyocoeliidae

Finks & Rigby (in Finks et al. 2004: p. 658) established

the new family Girtyocoeliidae designating Girtyocoelia as
the type genus of the family. Additional genera, including
Amphorithalamia Senowbari-Daryan & Rigby (1988), Ca-
lymenospongia Elliott (1963), Enoplocoelia Steinmann
(1882),  Henricellum Wilckens (1937), Phraethalamia Se-
nowbari-Daryan & Ingavat-Helmcke (1994), Polyedra Ter-
mier & Termier (1955), Sollasia Steinmann (1882), and
finally, with   question-mark Solenocoelia Cuif (1973), were
attributed to the Girtyocoeliidae. Later, an additional Or-
dovician genus, Girtyocoeliana Rigby, Karl, Blodgett &
Baichtal (2005) was added to this family.

Amphorithalamia was designated as the type genus of

the family Amphorithalamiidae by Senowbari-Daryan &
Rigby (1988). According to the “International Code of
Zoological Nomenclature (ICZN)”, published in its Ger-
man version by Kraus (2000), it is incorrect to move Am-
phorithalamia to another family established later than the
first establishment of its family (“Priority Principle”). Am-
phorithalamiidae has priority. It would be correct to leave

Fig. 1. Geographical position of the Kuh-e Bagh-e Vang (1) and
Deh-e Mohammad (2) localities in eastern Iran.

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Amphorithalamia within the originally established family
of Amphorithalamiidae. The same is also true for the ge-
nus  Polyedra, which was attributed to the family Poly-
edridae Termier & Termier (in Termier et al. 1977).

Finks & Rigby (in Finks et al. 2004) have attributed to

the family Girtyocoeliidae genera with totally different
diagnostic characteristics. For example, the genera Girtyo-
coelia, Enoplocoelia, Phraethalamia or Polyedra are char-
acterized by possession of one or more axial canals, but
Sollasia or Henricellum are asiphonate genera. The “interi-
or of each chamber” of Amphorithalamia (an asiphonate
sponge) is “filled with a fine, meandriform, trabecular net
that outlines anostomosing, tubular spaces” (Finks & Rig-
by: in Finks et al. 2004: p. 658). The filling skeleton, ve-
siculae excepted, is lacking in all other mentioned genera.
Because of the rule of priority (ICZN), the family names
Amphorithalamiidae and Polyedridae cannot be replaced
by Girtyocoeliidae, or any other new family.

Calymenospongia kudistanensis,  a minor chambered

fossil described from the Kolosh Formation (Paleocene) of
northern Iraq by Elliott (1963), is another genus, attribut-
ed to the Family Girtyocoeliidae by Finks & Rigby (in
Finks et al. 2004). The construction of this microfossil is
similar to Sollasia, as noted by Elliott. However, the very
small dimensions, which according to Elliott are: maxi-
mum diameter of 0.72 mm, chamber heights of 0.42 mm,
and a wall thickness of the chambers of 0.03 mm, differs
from all other sphinctozoan sponges. The wall structure is
also different from other sphinctozoan sponges. The sys-
tematic position of Calymenospongia is uncertain, and an
attribution to foraminifers is not be excluded.

Cuif (1973) described the genera Solenocoelia and Retic-

ulocoelia without placing them in a higher taxonomic cate-
gory. He illustrated both genera only by tangential sections,
which unfortunately do not show the sphinctozoan con-
struction. Also the species name “cylindrica“ does not re-
flect the sphinctozoan construction of this sponge.
Solenocoelia, attributed, althought with some doubt, to the
family Girtyocoeliidae by Finks & Rigby (in Finks et al.
2004), shows a wall with well-preserved spherulitic micro-
structure, without pores or ostia (Cuif, pl. 9, figs. 3—4, pl. 10,
fig. 4). The systematic position of this fossil as either a seg-
mented sphinctozoan or an unsegmented inozoa is uncer-
tain. Also the systematic position of  Reticulocoelia
arborescens Cuif (1973), attributed to the Virgulidae – a
family of inozoid sponges – by Finks & Rigby (2004:
p. 598), is uncertain.

In our opinion establishment of a new family Girtyocoeli-

idae, and separation of the genera included there from the
rest of the genera of the family Thaumastocoeliidae seems
superfluous and not necessary. We use in this paper the
“classic” classification (Senowbari-Daryan 1990, Senow-
bari-Daryan & Garcia-Bellido 2002) of the sphinctozoan
sponges to classify the new genus Shotorithalamia.

Concerning the above problem, J.K. Rigby (Provo, Utah)

concurs with our opinion and states (pers. com. Rigby, 11


February 2005): “You are right, as you have interpreted the
International Rules of Nomenclature. I think that the Am-
phorithalamiidae should be separated out, like we did (Se-

nowbari-Daryan & Rigby 1988). I would encourage you to
publish basically what you have written to me, that the Am-
phorithalamiidae is a valid and useful family, separate from the
Girtyocoeliidae, and that it ought to be maintained because of
differences, but also because of the rules and priority”.

Family:  Thaumastocoeliidae Ott, 1967

Subfamily:  Enoplocoeliinae Senowbari-Daryan, 1990

Shotorithalamia nov. gen.

D e r i v a t i o   n o m i n i s :

 From the Shotori Mountains,

and  thalamos (Greek) = chamber.

D i a g n o n i s :

 Aporate sphinctozoan sponge with ring-

chambers arranged around an axial spongocoel of ambisi-
phonate type. The exo- and endowalls are pierced by ostia
extended inwards by multibranched tubes with perforated
walls. The openings of exowall are rimmed. Vesiculae and
other internal structures are lacking.

T y p e   s p e c i e s :

  Shotorithalamia tubulara nov. sp.

D i s c u s s i o n :

 Because of its imperforate chamber walls,

Shotorithalamia nov. gen. is attributed to the family Thau-
mastocoeliidae Ott. Further placement of the genus within
the sphinctozoan sponges is discussed above.

In general, Shotorithalamia is similar to the genus Gir-

tyocoelia  Cossmann (1909), an abundant sponge in Car-
boniferous and Permian deposits, but differs from it by
having tubes extended from the exo- and endowalls into
the chamber interior. In Girtyocoelia the tubes may extend
from the exowalls to the outside (see Senowbari-Daryan
1990, pl. 45, figs. 5—6, text-fig. 48, reillustrated in Finks et
al. 2004: fig. 434/1d—1e) but not into the chamber interi-
ors, where the chamber interiors are hollow. The structure
of the spongocoel (Girtyocoelia is also prosiphonate,
while  Shotorithalamia is ambisiphonate) and the perfora-
tion types of both genera are totally different.

Because of the tubes within the chamber interiors, Shoto-

rithalamia resembles Praethalamia described by Senow-
bari-Daryan & Ingavat-Helmcke (1994) from Upper Permian
strata of Phrae province, Thailand. Shotorithalamia, howev-
er, differs from Phraethalamia by having only one spongo-
coel and the latter genus by having two or more axial
spongocoels. In addition, tubes in Phaerethalamia extend
only from the endowall into the chamber interiors. Tubes
like these, extending from the exowall into the chamber in-
teriors, as in Shotorithalamia, are absent in Phraethalamia.

Tubes that extend from the endowall into the chamber

interiors are also developed in the Permian genus
Pseudoamblysiphonella (Demospongea?) described by
Senowbari-Daryan & Rigby (1988) from Upper Permian
reefs of Djebel Tebaga of southern Tunisia and they also
occur in the species Barroisia lehmani (Calcispongea), de-
scribed by Hillmer & Senowbari-Daryan (1986) from the
Cretaceous strata of northern Germany. Tubes that extend
from the exowall into the chamber interior, as developed
in  Shotorithalamia, are absent in both of the foremen-
tioned genera. In addition, Pseudoamblysiphonella and
Barroisia have perforated chamber walls and are attribut-
ed to the group of Porata of Seilacher (1962).

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Fig. 2. 1—3 – Shotorithalamia tubulara nov. gen., nov. sp. 1 – Holotype. Marginal axial longitudinal section show the ring-chambers,
spongocoel  and  the  rimmed  ostia  on  the  exowalls.  In  the  third  chamber  from  the  bottom  the  tubes  extended  from  the  exo-  and  endowalls
appear as circles. 2 – Magnification from Fig. 1 exhibits one rimmed ostium of the exowall. 3 – Magnification from Fig. 1 shows two
chambers with a rimmed ostium of the exowall and the tubes extended from the ostia into the chamber interiors. 4—5 – Amblysiphonel-
cf. vesiculosa vesiculosa de Koninck. 4 – Transverse oblique section shows the axial spongocoel and a few chambers filled with ve-
siculae. 5 – Longitudinal oblique section shows the crescent-like chamber filled with vesicular skeleton.

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Shotorithlamia nov. gen. differs from Girytyocoeliana, an-

other genus of the family, described by Rigby, Karl, Blodgett
& Baichtal (2005) by possessing of the internal tubes and by
the lack of “rimmed to tubular interpores in irregular rings
around central tube” (Rigby et al. 2005: p. 867).

Multiple tubes, as tubular filling skeleton, fills chamber

interiors of representatives attributed to the family Polytho-
losiidae Seilacher (1962). The kind of filling skeleton and
the perforated chamber walls distinguish Shotorithalamia
from the taxa attributed to the family Polytholosiidae. No
other sphinctozoan genera are comparable with Shoto-
rithalamia nov. gen.

Shotorithalamia tubulara nov. sp.

(Figs. 2.1—3, 3, 4)

D e r i v a t i o   n o m i n i s :

 Named from the occurrence of

the tubes extending from the endo- and exowalls into the
chamber interiors.

H o l o t y p e :

 Fig. 2.1 (magnifications of holotype are il-

lustrated in Figs. 2.2—3).

L o c u s   t y p i c u s :

 Kuh-e Bagh-e Vang, about 45 km

NW of Tabas (see Fig. 1, locality 1).

S t r a t u m   t y p i c u m :  

Lower Permian, most probably


D i a g n o s i s :

 See diagnosis of the genus.

M a t e r i a l :  

One specimen (holotype) only.

D e p o s i t o r y :

 The material is filled together with

sponge material described by Senowbari-Daryan et al.
(2005) in the collection of the Institute of Paleontology,
University of Erlangen-Nürnberg (Material: Senowbari-
Daryan, Permian, Bagh-e Vang, thin section BVA/1 and a
rock piece with the other half of the sponge).

D e s c r i p t i o n :

 Because there is only one available

specimen of the species the description of holotype corre-
sponds to the description of the species.

The 50 mm long sponge with a diameter of 18 mm is

composed of 9 spherical to rectangular ring-like chambers
arranged around an axial spongocoel (Fig. 2.1). The last
two chambers are incompletely developed or broken.
Chamber height varies between 5 and 7 mm. The chamber
walls (exo-, inter- and endowalls) have the same thickness
of 0.8—1.0 mm. Because of the thickening of chamber
roofs by doubling of the walls (especially at the peripheral
part) these may appear thicker than 1 mm. An axial spon-
gocoel of ambisiphonate type, 4 mm in diameter, passes
through the whole sponge length. The exowalls are
pierced by rimmed ostia of about 0.6 mm (Figs. 2.2—3).
The most characteristic features of the sponge are the
branched tubes extended from the ostia and those from the
endowall into the chamber interiors (Figs. 2.3, 3—4). The
tubes are multibranched and pierced by numerous pores
connecting the tubes interior with the chambers (Fig. 3).
The tubes from exowalls and those from endowalls are not
connected with others. Tubes in oblique sections through
the chambers appear as circles (Fig. 2.1, in lower cham-
bers, compare with Fig. 3). The openings of the endowall
and the tubes have the same diameter as the ostia on the
exowalls. Skeletal mineralogy, microstructure and spi-

Fig. 3.  Shotorithalamia tubulara nov. gen., nov. sp. (holotype,
counterpart to Fig. 2.1). Drawn from thin section. Scale bar = 1 cm.

Fig. 4. Reconstruction of Shotorithalamia tubulara nov. gen.,
nov. sp., showing the ring-like chambers around the axial spon-
gocoel, tubes within the chamber interiors, and the rimmed ostia
within the exowalls. Schematic (not to scale).

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cules are unknown. A reconstruction of Shotorithalamia
tubulara showing the chambers, tubes within the interior
of the chambers and the rimmed ostia is given in Fig. 4.


A total number of 12 sponge genera and 15 species, in-

cluding sphinctozoans (7 genera and 9 species) and inozoid
(5 genera and 6 species), were described by Senowbari-
Daryan et al. (2005) from the reefal limestone of the Permi-
an Jamal Formation exposed in Kuh-e Bahg-e Vang. With
documentation of Shotorithalamia tubulara the number of
sponge taxa in this locality is raised to 13 genera and 17
spieces, including Amblysiphonella cf. vesiculosa. The reefal
limestones of the Jamal Formation exposed in Kuh-e Bagh-e
Vang should be classified as sponge-Tubiphytes-Archaeolo-
thoporella-boundstones (Senowbari-Daryan et al. 2005).

As discussed by Senowbari-Daryan et al. (2005), the

sponge association of the Permian Jamal Formation in
Kuh-e Bagh-e Vang shows considerable similarity to the
sponge association known from the Lower Permian reef
boulders imbedded within the Lercara Formation, western
Sicily (Senowbari-Daryan & Di Stefano 1988).


 The investigations were carried out

within the framework of Project Se 416/11, supported by
the Deutsch Forschungsgemeinschaft (DFG). The authors
thank the University of Esfahan for logistic support of our
field work. Our special thanks are addressed to Prof. Dr. J.
Keith Rigby (Provo, Utah) for the improvement of the first
draft of the manuscript. The authors express their gratitude
to Diego C. Garcia-Bellido (Madrid), Jozef Michalík (Bra-
tislava), and Oliver Weidlich (Berlin) for their fair and
constructive comments as reviewers.


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