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c Cambridge University Press 2012
Geol. Mag. 149 (6 ), 2012, pp. 1023–1045. doi:10.1017/S0016756812000179
1023
The Late Palaeozoic trilobites of Iran and Armenia and their
palaeogeographical significance
R U DY L E R O S E Y- AU B R I L ∗
Forschungsinstitut Senckenberg, Senckenberganlage 25, D-60315 Frankfurt am Main, Germany
(Received 11 January 2011; accepted 13 February 2012; first published online 27 April 2012)
Abstract – The Iranian territory is composed of a mosaic of tectonic units, several of which
underwent in the Permian and Triassic periods a migration from northern Gondwana to southern
Laurussia associated with the opening of the Neo-Tethys Ocean. Although this broad outline of PermoTriassic palaeogeographical evolution of Iranian microplates is now widely accepted, the individual
timing of migration of these blocks, and their biogeographical relationships, remain insufficiently
known. Here I review the Late Palaeozoic record of trilobites in Iran and Armenia, and discuss
their palaeobiogeographical affinities in an attempt to shed light on the Permian palaeogeographical
evolution of Iranian and Armenian terranes. Seven Iranian or Armenian localities, representative of
five tectonic units, have yielded Carboniferous and Permian trilobites. Ten species are recognized,
including two new taxa, Persia praecox gen. nov. sp. nov. and Pseudophillipsia (s.l.) parvizii sp. nov.
P. praecox is the only Carboniferous (Tournaisian) species. The others are Wordian to Wuchiapingian
in age and can be separated into three morphological groups, probably representing clades. One is
composed of representatives of Acropyge, while the two others (armenica-group and paffenholzigroup) comprise species of Pseudophillipsia. Only P. (s.l.) parvizii sp. nov. from the Zagros Mountains
(Arabian Plate) is not attributed to one of these groups. The distribution of trilobites in Iran and
Armenia strongly suggests that the Alborz, Central Iran and Transcaucasia microplates represented a
single biogeographical unit in Middle and Late Permian times. Special relationships of this biochore
with South China can also be stressed.
Keywords: Trilobita, Carboniferous, Permian, Iran, Armenia, palaeobiogeography.
1. Introduction
Even if some of them possessed planktonic larval stages
facilitating dispersal, adult trilobites were primarily
benthic organisms restricted in their habitat to a certain
depth-range. Consequently, the palaeobiogeography of
trilobites, especially those living on the continental
shelf, was strongly influenced by the distribution of
continents, which permits, retrospectively, the use of
these fossils for inferring past continental configurations. Trilobites have thus proved to be critical in
constraining palaeogeographical models in the Lower
Palaeozoic (e.g. Fortey & Cocks, 2003). In the Permian
period, however, their taxonomic diversity (LeroseyAubril & Feist, 2012) and their abundance are so low
that these organisms are almost never considered in
reference to the evolution of palaeogeography (the
same is true for biostratigraphy). On the one hand,
with such a patchy fossil record in the Late Palaeozoic,
it may make sense to think that the potential of
trilobites for discriminating Permian biochores (i.e.
biogeographical units) may be negligible compared
with other fossil groups (e.g. brachiopods, fusulinid
foraminifera) and therefore that their palaeobiogeographical signal is worthless. However, this would be
ignoring the fact that the definition of a biochore,
∗
E-mail: rudy.lerosey-aubril@senckenberg.de
like that of a stratigraphical unit, is all the more
precise and robust if it integrates data from different
taxa (Westermann, 2000). This approach avoids bias
due to particular ecological characteristics of any one
given taxon and permits a good assessment of the
evolution through time of the geographical range of a
biochore.
The Iranian territory represents a particularly complex tectonic context, being composed of a mosaic of
more or less well-recognized tectonic units (Zanchi
et al. 2009). Unravelling the evolution through time
of these blocks and of their positions relative to
one another and to major continents (e.g. Gondwana,
Laurentia) is a difficult task, especially in the Permian period when several of these microplates were
separated from the northern Gondwana margin and
migrated towards the north during the opening of the
Neo-Tethys (Zanchi et al. 2009). Every potential source
of a palaeogeographical signal (e.g. palaeontological,
sedimentological, palaeomagnetical), even from a
minor component of fossil assemblages like trilobites,
requires consideration. The objective of this contribution is twofold: (1) to review the Late Palaeozoic record
of trilobites in Iran and Armenia and (2) to discuss their
palaeobiogeographical affinities with other Permian
taxa and their implications with regard to palaeogeographical models of the Iranian region in the Permian
period.
1024
R . L E R O S E Y- AU B R I L
Figure 1. Geographical map (a) and simplified tectonic scheme (b) of Iran and neighbouring countries, showing the location of the
outcrops that have yielded Late Palaeozoic trilobites. Five tectonic units (Iran: Alborz, Arabian Plate, Sanandaj–Sirjan Belt, Tabas;
Armenia: Transcaucasia) are associated with Late Palaeozoic trilobite faunules. Labels: 1 – Abadeh area (Capitanian); 2 – Chahriseh
area (Wordian); 3 – Dena Range (Wuchiapingian); 4 – Yush area (Wuchiapingian); 5 – Howz-e-Dorah area (Tournaisian); 6 – Vedi
area (Wordian); 7 – Ogbin area (Wordian). Abbreviations: Arm. – Armenia; Azerb. – Azerbaijan; Bah. – Bahrain; Koh. Boy.-Ahm. –
Kohgiluyeh and Boyer-Ahmad Province; Turk. – Turkey; U.A.E. – United Arab Emirates; AA – Araxian–Azerbaijanian Zone;
GKDF – Great Kavir–Doruneh Fault system; MZT – Main Zagros Thrust; PTSZ – Palaeo-Tethys Suture Zone; SSZ – Sanandaj–Sirjan
Zone; T – Talesh. Figure 1b simplified after Zanchi et al. 2009.
2. Materials, localities and structural units
2.a. The Abadeh area (Sanandaj–Sirjan Belt, central Iran)
Late Palaeozoic trilobites have been found in five
different localities in Iran (1 to 5 on Fig. 1), the
distribution of which covers an area of c. 360 000 km2
in the middle third (along a W–E axis) of the country.
Although limited in size compared with the whole
of Iran (c. 1 650 000 km2 ), this area includes parts of
five distinct tectonic units, four of which have yielded
trilobite remains (Fig. 1b). Several of these structural
units represent more or less independent microplates
(western Cimmerian blocks) during the Permian period,
separated from northern Gondwana by the opening of
the Neo-Tethys Ocean. In addition to these five Late
Palaeozoic outcrops in Iran, two southern Armenian
localities (6 and 7 on Fig. 1) are considered in this
work because of their proximity to the Iranian border
and notable similarities in their trilobite faunules. The
geographical and stratigraphical details for all these
localities are summarized in Table 1.
This review integrates the most recent stratigraphical
data concerning each locality but also, for some of
them, new data inferred from the study of thin-sections.
Although only two trilobite occurrences are new (from
Dena Mountain and Howz-e-Dorah; 3 and 5 on Fig. 1),
an effort has been made to figure or refigure all the Late
Palaeozoic trilobites discovered in Iran and Armenia
in an attempt to facilitate future investigations on this
material. Original pictures have been used to illustrate
the few specimens that could not be relocated in the
collections of their host institutions. Otherwise, new
pictures have been taken.
The first description of post-Devonian trilobites from
Iran was made by Kobayashi & Hamada (1978) and
concerned specimens found in the Permian beds exposed in the vicinity of Abadeh (Fars Province, Central
Iran; label 1 on Fig. 1). Initially, this material was
described as two new taxa, Acropyge lanceolata and
Iranaspidion sagittalis, for which a late Guadalupian
age had been proposed (Kobayashi & Hamada, 1978).
Of the nine specimens (four type specimens, five
additional ones) initially studied by these authors, only
two can still be found in the collections of the University
Museum of the University of Tokyo (T. Sasaki, pers.
comm. 2010): the holotype of A. lanceolata (PA
16766) and a pygidium of I. sagittalis (PA 16765). The
holotype and the two paratypes of the latter species are
missing. A. lanceolata is considered herein as a subjective junior synonym for A. encrinuroides (Weber, 1944).
I. sagittalis is now referred to Pseudophillipsia (Carniphillipsia) sagittalis (Kobayashi & Hamada, 1978),
after Iranaspidion had been itself put in synonymy with
Pseudophillipsia (Lerosey-Aubril & Angiolini, 2009),
but only the holotype and its two paratypes (Kobayashi
& Hamada, 1978, figs 1, 3a–c) are herein regarded as
representatives of this taxon. The isolated pygidium
figured by Kobayashi & Hamada (1978, fig. 4a–c)
and refigured herein (Fig. 4b, d, e) is reassigned to
Pseudophillipsia (s.l.) armenica Weber, 1944.
According to Kobayashi & Hamada (1978), the
trilobites were found in the upper part of Unit 1 of
the Abadeh section as defined by Taraz (1971). Taraz’s
Late Palaeozoic trilobites of Iran and Armenia
1025
Table 1. Geographical, structural and stratigraphical details for each Armenian and Iranian locality that has yielded Late Palaeozoic trilobites
Localities
Structural Units Stratigraphy
1: Abadeh (Ir) Hambast
SSZ
Mountains Fars Province
2: Chahriseh/Kuh-e Kaftar
(Ir) Kaftar Mount
Esfahan Province
3: Kuh-e Dena (Ir) Zagros
Mountains Kohgiluyeh
& Boyer-Ahmad
Province
4: Yush (Ir) Central Alborz
Mountains Mazandaran
Province
5: Howz-e-Dorah (Ir)
Shotori Range Yazd
Province
6: Vedi (Ar) Ararat
Province
Abadeh Formation (Unit 4)
Guadalupian (Capitanian)
Surmaq Formation (upper
Unit 1) Guadalupian
(Capitanian)
Trilobite Taxa
References
‘Pseudophillipsia sp.’
Taraz, 1971
Kobayashi &
Hamada, 1978
SSZ
Jamal Formation Guadalupian
(Wordian?)
Arabian Plate
(Zagros Belt)
Dalan Formation Lopingian
(Wuchiapingian)
Acropyge encrinuroides
Pseudophillipsia (s.l.) armenica
Pseudophillipsia (Carniphillipsia)
sagittalis
Pseudophillipsia (Carniphillipsia)
sp. A
‘Pseudophillipsia sp.’
Pseudophillipsia (s.l.) parvizii sp.
nov.
Alborz
Nesen Formation Lopingian
(Wuchiapingian)
Acropyge weggeni
Hahn & Hahn, 1981
Pseudophillipsia (s.l.) aff. caucasica
Central Iran
(Tabas Block)
Shishtu Formation
Mississippian (Tournaisian)
‘Trilobites’
Persia praecox gen. nov. sp. nov.
Transcaucasia
Gnishik Formation Guadalupian Pseudophillipsia (Carniphillipsia)
(Wordian)
paffenholzi
Acropyge sp. A
Pseudophillipsia (s.l.) armenica
Pseudophillipsia (Carniphillipsia)
paffenholzi
Gnishik Formation?
Acropyge encrinuroides
Guadalupian (Wordian)
Pseudophillipsia (s.l.) armenica (?)
Pseudophillipsia (s.l.) caucasica
Permian
Pseudophillipsia (s.l.) armenica
7: Ogbin (Ar) Vayots Dzor Transcaucasia
Province
Tananam (Az) Nakhchivan Transcaucasia
Autonomous Republic
Mistiaen et al. 2000
Ernst et al. 2006
–
Yazdi, 1999
–
Weber, 1939, 1944
Arkhipova, 1965
Weber, 1944
Weber, 1944
Numbers in first column are the same as in Figure 1. Abbreviations: Ar – Armenia, Ir – Iran, SSZ – Sanandaj–Sirjan Belt. A single specimen
of P. (s.l.) armenica might have been found in the locality of Tananam (Nakhchivan Autonomous Republic; Weber, 1944) but, as it has never
been figured, it is not considered further in this work.
Units 1 to 3 were later considered parts of the Surmaq
Formation (Taraz et al. 1981) and the uppermost
part of Unit 1 has been recently dated as Capitanian
(Midian; Kobayashi & Ishii, 2003). Taraz (1971,
p. 1289) also mentioned the presence of trilobites
in the basal part of his Unit 4 (‘Pseudophillipsia
sp.’), now part of the Abadeh Formation (Taraz et al.
1981), which is also of Capitanian (Midian) age
(Mohtat-Aghai & Vachard, 2005). However, he did not
provide any description or figure of this material, and
accordingly, I do not consider it further in Section 3
below. Structurally, the Abadeh area is part of the
Sanandaj–Sirjan structural belt (SSZ; Fig. 1b), a region
rich in strongly deformed and metamorphosed rocks
sandwiched between Central-North Iran and the Zagros
Orogenic Belt (Zanchi et al. 2009).
2.b. The Chahriseh area (Sanandaj–Sirjan Belt,
central Iran)
Another locality belonging to the Sanandaj–Sirjan
structural belt has yielded Permian trilobites. Indeed,
Feist et al. (in Mistiaen et al. 2000) have described a
single entire specimen collected from a loose block in
Kuh-e Kaftar, a locality near Mount Kaftar and a village
called Chahriseh, some 55 km northeast of Esfahan
(label 2 on Fig. 1). This specimen, initially assigned
to Pseudophillipsia (?Carniphillipsia) sp. is below
referred to as Pseudophillipsia (Carniphillipsia) sp. A.
The age of the Permian sequence in this area is
not well-constrained, ranging from the Wordian to the
Wuchiapingian according to microfossils (middle–late
Murghabian to early Dzhulfian; Mistiaen et al. 2000).
In addition, Permian trilobites (‘Pseudophillipsia sp.’)
in a neighbouring section have been reported by Ernst,
Senowbari-Daryan & Hamedani (2006). These authors
referred to the Permian beds exposed in their section as
part of the Jamal Formation (see also Senowbari-Dayan
& Hamedani, 2002), a lithographical unit well known
in eastern Iran (e.g. Wendt et al. 2002). Unfortunately, a
comparison between this Permian sequence and that of
the Abadeh area (same structural unit, less than 200 km
to the south), and therefore its possible correlation
with one of the lithostratigraphical units known in
Abadeh (Surmaq, Abadeh or Hambast formations),
is nowhere to be found in the contributions of this
working group (Senowbari-Daryan & Hamedani, 2002;
Rigby, Senowbari-Daryan & Hamedani, 2005; Ernst,
Senowbari-Daryan & Hamedani, 2006). A Wordian
age (Murghabian) has been proposed for these trilobitebearing beds based on their bryozoan content (Ernst,
Senowbari-Daryan & Hamedani, 2006), because of its
great similarity with that of the Gnishik horizon in
Transcaucasia (see Section 2.f below for further details
1026
about this lithostratigraphical unit). Accordingly, a
similar age is tentatively given to the specimen
described in Mistiaen et al. (2000). However, the
assignment by Ernst, Senowbari-Daryan & Hamedani
(2006) of the trilobite remains to the genus ‘Pseudophillipsia’ is purely arbitrary, as they were only
observed in thin-sections (B. Senowbari-Daryan, pers.
comm. 2010).
2.c. The Dena range (Arabian Plate, southwestern Iran)
Some 120 km west of Abadeh and 40 km northwest of
Yasuj, in the Dena Range (Zagros Mountains; label 3
on Fig. 1), is another Permian outcrop where trilobite
remains occur. Two pygidia were recently discovered
there by M. Parvizi (University of Payam-e Noor)
on some loose blocks, which likely came from beds
belonging to the Dalan Formation. These specimens
are assigned to a new species Pseudophillipsia (s.l.)
parvizii sp. nov.
Thin-sections made from the surrounding matrix revealed that this sediment corresponds to a sandy wackestone with gastropods, brachiopods (e.g. productoids),
bryozoans (fenestellids), crinoids, ostracods and very
rare trilobites (D. Vachard, pers. comm. 2010). The rare
foraminifers (1–5 specimens per genus) observed were
representatives of Fusulinata (Neoendothyra parva,
Retroseptellina ex gr. decrouezae, Codonofusiella
ex gr. tenuissima), Miliolata (Pseudovermiporella ex
gr. nipponica) and Nodosariata (Pachyphloia ex gr.
ovata, Pachyphloia sp., Langella ex gr. perforata).
The inferred depositional environment is that of the
boundary between the inner- and middle ramp (10–
25 m deep). A Wuchiapingian age is indicated by
the presence of Codonofusiella ex gr. tenuissima
(D. Vachard, pers. comm. 2010). Separated from the
Sanandaj–Sirjan structural belt by the main Zagros
thrust, the Zagros fold belt is a part of the Arabian
Plate (Gaillot & Vachard, 2007; Zanchi et al. 2009).
2.d. The Yush area (Alborz, northern Iran)
The fourth locality that has yielded Permian trilobites
in Iran is in the Central Alborz Mountains, near a
small village called Yush (exact location: 36◦ 13.05 N,
51◦ 42.20 E; label 4 on Fig. 1). Hahn & Hahn (1981)
used one of the three specimens discovered there to
define a new species, Acropyge weggeni, and referred
to the other two as Acropyge? sp. and Iranaspidion?
sp. Reinvestigations of this material housed at the
Senckenberg Research Institute confirms that two of
these specimens belong to the genus Acropyge. It
seems overcautious to me not to refer to the very
small pygidium assigned to Acropyge? sp. by Hahn
& Hahn (1981) as A. weggeni (see Section 3 below).
The third specimen is a partial pygidium associated
with an external mould of its posterior region. After
preparation, the latter was used to create a latex
cast, which provides complementary data on the
R . L E R O S E Y- AU B R I L
morphology of this specimen. Below, this specimen
is referred to as Pseudophillipsia (s.l.) aff. caucasica.
The horizon that has yielded this material belongs to the Nesen Formation, which is of mainly
Wuchiapingian age (Gaetani et al. 2009). Three thinsections made from the associated matrix revealed that
it is a wackestone with remains of bivalves, brachiopods, bryozoans, crinoids, gastropods, ostracods and
trilobites (D. Vachard, pers. comm. 2010). In addition,
it contains the following foraminifera: Codonofusiella
cf. tenuissima, Climacammina sp., Dunbarula tumida,
Earlandia? sp., Geinitzina cf. taurica, Globivalvulina
sp., Midiella sp., Pachyphloia ex gr. ovata and
Retroseptellina decrouezae. This association of taxa
confirmed the Wuchiapingian age of the trilobite
faunule (D. Vachard, pers. comm. 2010). The Alborz
region is usually considered a structural unit of its own
(e.g. Torsvik & Cocks, 2004; Muttoni et al. 2009a;
Zanchi et al. 2009).
2.e. Howz-e-Dorah (Tabas Block, eastern Iran)
As far as has been determined, only one Carboniferous
trilobite has been found in Iran until now. This specimen
was collected by M. Yazdi (University of Esfahan)
during a conference excursion at the Howz-e-Dorah
section in the Shotori Range (see Yazdi, 1999 and
Wendt et al. 2005 for details about this locality; label 5
on Fig. 1). It is herein assigned to a new taxon, Persia
praecox gen. nov. sp. nov.
The trilobite-bearing horizons, already mentioned by
Yazdi (1999, fig. 5), belong to the Shishtu Formation
and are dated as Tournaisian (typicus conodont
biozone). Howz-e-Dorah is located near Tabas and
therefore on a structural sub-unit of Central Iran, the
so-called Tabas Block (Zanchi et al. 2009).
2.f. Ogbin and Vedi areas (Transcaucasia,
southern Armenia)
The occurrence of Permian trilobites in the Vedi area
(label 6 on Fig. 1) in southern Armenia was first
reported by Weber (1939). A few years later, these
specimens were described, along with others discovered
in the neighbouring locality of Ogbin (label 7 on Fig. 1)
(Weber, 1944). However, discrepancies in this work
between the text in Russian, his table 2, the text in
English and the captions of his figures are confusing,
and it requires particular attention to assess correctly
the faunal content of each locality. Weber’s table 2
(1944, p. 18) obviously contains mistakes and his text
in English does not provide all the data mentioned in
the Russian part. According to this Russian part and
the captions of the figures, the following summary
can be given. Two apparently close outcrops along
the Vedi River have yielded remains of Cyphinium
(?) paffenholzi, a species which is referred to as
Pseudophillipsia (Carniphillipsia) paffenholzi below.
In another locality near Ogbin village, a pygidium
of Pseudophillipsia (Anisopyge) encrinuroides and a
Late Palaeozoic trilobites of Iran and Armenia
pygidium of Pseudophillipsia armenica have been
found. In addition, a single pygidium of Pseudophillipsia elegans var. caucasica might have been discovered
in the same locality, but Weber does not provide
accurate geographical data for this specimen, which
is only mentioned in his Russian text. In his table 2,
however, the presence of this taxon is mentioned in this
locality. Pseudophillipsia (Anisopyge) encrinuroides,
Pseudophillipsia armenica and Pseudophillipsia elegans var. caucasica are hereafter referred to as Acropyge
encrinuroides, Pseudophillipsia (s.l.) armenica and
Pseudophillipsia (s.l.) caucasica. Apparently, a second
specimen of P. (s.l.) armenica has been collected in
Tananam (‘locality 63’), a third locality between Ogbin
and Karabaglyar in the neighbouring Azerbaijan (Nakhchivan Autonomous Republic), but it was not figured
by Weber and could not be considered in this work.
More recently, Arkhipova (1965, pp. 82–3, pl. 45,
figs 1–3) figured specimens coming from the Vedi
area (‘Vedi 2’, Gnishik horizon; label 6 on Fig. 1):
a cephalon and a pygidium that she attributed to
Pseudophillipsia armenica and a smaller pygidium she
assigned to Pseudophillipsia paffenholzi. The cephalon
is hereafter reassigned to P. (C.) paffenholzi and
the small pygidium to P. (s.l.) armenica. The larger
pygidium, however, has a peculiar morphology that
I consider evocative of the genus Acropyge and it
is therefore referred to below as Acropyge sp. A.
Unfortunately, these three specimens could not be
found again in the collections of the Palaeontological
Institute in Moscow and it seems likely that they are
definitively lost (E. Naimark, pers. comm. 2010).
Weber (1944) only mentioned a vague ‘Permian’ age
for the trilobites he described from southern Armenia,
while Arkhipova (1965) suggested a ‘Guadalupian’
age. According to Leven (1998), the fusulinids of the
Gnishik Formation are indicative of a Wordian (= late
Murghabian) age, and consequently, at least the
specimens figured by Arkhipova (1965) might be of
that age. Leven (1998) also stated that similarities
in fusulinid assemblages suggest that ‘the Gnishik
Formation corresponds approximately to the upper half
of Unit 1’ of Taraz (1971). However, it has been shown
recently that the uppermost part of this Unit 1 contains
the fusulinid Chusenella abichi, the index species of the
Midian in Transcaucasia (Kobayashi & Ishii, 2003).
Moreover, Partoazar (2002) affirmed that the microfauna of the ‘Gnishik beds’ in the Jolfa area (Northwestern Iran) indicates an early Wuchiapingian (=
early Dzuhlfian) age, but he did not specify which taxa
led him to this conclusion. Considering this, and the
fact that there is no precise correlation between the
Gnishik Formation and the upper part of Unit 1 of
the Surmaq Formation, I am more inclined to consider
the trilobites figured by Arkhipova (1965) as Wordian
in age, keeping in mind that they might be slightly
younger (i.e. Capitanian). Since two of the three species
represented by this material were originally described
by Weber (1944), it seems reasonable to consider a
similar age for the specimens described by this author.
1027
Along with the majority of Azerbaijan and Georgia,
Armenia is frequently regarded as part of an independent tectonic unit variously named Transcaucasia
(e.g. Leven, 1998; Gaillot & Vachard, 2007; Théry,
Vachard & Dransart, 2007; Gaetani et al. 2009;
herein), Transcaucasus (e.g. Ezaki, 1993; Kobayashi
& Ishii, 2003), Lesser Caucasus (e.g. Ruban, 2007a,b;
Ruban, Al-Husseini & Iwasaki, 2007) or the Araxian–
Azerbaijanian Zone (Angiolini et al. 2007; Muttoni
et al. 2009b; Zanchi et al. 2009). In this work,
Transcaucasia and the Araxian–Azerbaijanian Zone are
used interchangeably in reference to this tectonic unit.
3. Systematic palaeontology
Terminology and repositories. Morphological terms and
abbreviations used herein follow those defined by Whittington et al. (1997). In addition, ‘main glabellar lobe’
designates the part of the glabella in front of the preoccipital
lobes, while ‘anterior glabellar furrows’ refers to glabellar
furrows S2 to S4. The term ‘enrollment’ and ‘volvation’
are considered hereafter as interchangeable (for further
details about the term ‘volvation’ see Feist, Lerosey-Aubril
& Johnson, 2010). The specimens discussed herein are
housed in the collections of the ‘A. P. Karpinsky’ Russian
Geological Research Institute (Saint Petersburg; VSEGEI),
the Esfahan University (EUIT), the Nanjing Institute of
Geology and Palaeontology (NIGPAS), the Palaeontological
Institute of the Russian Academy of Science (Moscow; PIN),
the Senckenberg Research Institute (Frankfurt am Main;
SMF) and the University Museum of the University of Tokyo
(Tokyo; PA).
Order PROETIDA Fortey & Owens, 1975
Family PHILLIPSIIDAE Oehlert, 1886
Subfamily CUMMINGELLINAE Hahn & Hahn, 1967
Genus Persia gen. nov.
Type species.
Persia praecox sp. nov.
The type species is the only known
representative to date.
Assigned species.
Etymology.
From the ancient name of the country of origin.
Diagnosis. A genus within the Cummingellinae displaying
the following unique combination of characters: glabella
hourglass-shaped with anterior and posterior parts of subequal width (tr.), long (sag.) and rather slender and flat,
anterior border rather wide (sag. and exs.) and not overhung
by glabella, S1 broadly curved backwards to meet SO, L1
sub-circular, S2–S4 inconspicuous, anterior fixigena rather
wide (tr.), high cephalic border, short (exs.) genal spines,
pygidium with 10 + 1 axial rings and 8 pleural ribs devoid
of conspicuous inter-pleural furrows.
The well-defined cephalic anterior border and
the rather slender glabella can be regarded as primitive
characters within the Cummingellinae, being observed in
Comptonaspis, Liobolina, Moschoglossis or Richterella. On
the other hand, the new species also exhibits morphological
traits that are more frequent in younger representatives of
the group (e.g. Bedicella, many species of Cummingella,
Paraphillipsia): an hourglass-shaped glabella with a clear
constriction separating the preoccipital glabella into two
parts of roughly equal width (tr.), S1 curved backwards to
meet SO and delimiting L1 that somewhat encroaches on
the occipital ring, S2–S4 barely visible (if present), and to
a lesser extent, a rather limited number of axial rings and
Discussion.
1028
R . L E R O S E Y- AU B R I L
pleural ribs (10 and 8, respectively) in the pygidium. As a
consequence, it does not fit with the description of any of the
cummingelline genera known to date as they are presently
conceived. All in all, it looks like a derived cummingelline
(e.g. most species of Cummingella, Bedicella), which would
be devoid of the typically massive glabella. In doing so, it
more closely resembles two Carboniferous cummingellines:
Xiangzhongella (Liu, 1987) and Zhegangula (Hahn, Hahn &
Yuan, 1989). From the Bashkirian of South China (Guangxi
Province), Zhegangula exhibits, like the new taxon, an
elongate glabella. However, it is much more slender and
almost reaches the anterior margin in the Chinese species.
In addition, its anterior border is extremely narrow (sag.
and exs.), as are its anterior librigenal fields (tr.), and it
still displays a pronounced S2. The pygidium of Zhegangula
resembles that of Persia in its general aspect and in the
number of axial rings (10) and pleural ribs (7 instead of 8) it
possesses. However, it has a wider (tr.) axis and, despite the
claim of Hahn, Hahn & Ramovš (1990, p. 161), shallow but
obvious inter-pleural furrows split its pleural ribs into anterior
and posterior bands of sub-equal width (exs.); these furrows
are inconspicuous in Persia. Reduced posterior bands of
pleural ribs are observed on the pygidium of Xiangzhongella,
another Carboniferous (Tournaisian) cummingelline from
South China (Hunan Province). Xiangzhongella also has
in common with the new taxon a moderately expanding
forwards glabella separated by a furrow from an anterior
border that it does not or only weakly overhangs, barely
visible S2–S3, most genal characters, including the course
of the facial suture, and a narrow (at best) pygidial border.
However, Xiangzhongella differs from Persia in having a
wider (tr.) glabella, a narrower (sag., exs., tr.) and especially
lower (in lateral view) cephalic border, no genal spines,
thoracic axial rings apparently with pre-annuli, a greater
number of axial rings (13 or more) and pleural ribs (9 or
more) in the pygidium, and the presence of tubercles on the
axial rings of both the thorax and the pygidium.
Persia praecox sp. nov.
Figure 2a–e
Material, locality and horizon. Holotype, complete and articulated specimen, partially exfoliated (EUIT 7–850; Fig. 2);
thick-bedded limestone with solitary corals, brachiopods,
crinoids, bivalves and trilobites; Gnathodus typicus conodont
biozone (see Yazdi, 1999), Shishtu Formation, Howz-eDorah area, South Shotori Range, Yazd Province, Eastern
Iran (label 5 on Fig. 1).
In reference to the fact it resembles a derived
cummingelline but with a rather slender glabella, a trait generally observed in primitive taxa and in juvenile specimens
of derived taxa in this subfamily.
Etymology.
Diagnosis.
As for the genus.
Cephalon of semi-circular outline (Fig. 2b).
Glabella rather long (maximum width/maximum length
ratio: 0.6), hourglass in outline due to a strong constriction
of the rather shallow and broad dorsal furrows opposite γ
adaxially enabling the recognition of two parts of roughly
equal width (tr.). Occipital ring wide (sag. and exs.);
SO rather well incised and composed of a long median
portion curved forwards and short abaxial portions running
backwards from axial furrows. Posterior part of preoccipital
glabella slightly extending outwards; S1 shallow and evenly
curved from axial furrows to SO; L1 sub-circular in outline
and only faintly inflated; short and almost inconspicuous S2–
S4 possibly present near glabellar constrictions; preglabellar
furrow shallow but narrow. Anterior border short (sag. and
Description.
exs.) all along and curved backwards abaxially. Fixigena
composed of a particularly narrow (tr.) posterior field,
narrow (tr.) but long (exs.) palpebral lobe and narrow (tr.)
anterior field that more than doubles in width forwards.
α located opposite glabellar constriction anteriorly; α–γ
strongly curved outwards so that β has a position similar
to ζ relative to the sagittal line, i.e. the most abaxial position
of the suture after ω; γ–ε gently curved outwards with δ
being slightly more abaxially positioned than ε, like the
latter relative to γ and γ relative to α; ε–ζ almost straight,
short and diverging; ζ–ω long and strongly diverging with η
barely defined and ω located at the mid-width (tr.) of the
pleural lobe. Eye kidney-shaped, long (exs.), narrow (tr.)
and surrounded by a broad depressed subocular groove.
Librigenal field anterior to the eye rather narrow (tr.),
only moderately widening (tr.) posterior to eyes, bounded
posteriorly by a rather deep but shallow posterior border
furrow and laterally and anteriorly by a very shallow to
inconspicuous lateral border furrow. Posterior border of
equal width all along representing about two-thirds of the
width of the occipital ring. Lateral border rather narrow
(tr.) and barely differentiated from librigenal field. Short
genal spine. In lateral views (Fig. 2a, c), glabella rather low
compared with the genal lobe; occipital ring slightly convex
dorsally although culminating at roughly the same height as
adjacent part of the preoccipital glabella, which is almost
flat-topped in this region, but anteriorly it gently curves
downwards until it abuts the slightly inflated anterior border.
Eye is low, especially anteriorly, while the lateral border is
high. In frontal view (Fig. 2d), cephalon moderately arched
(tr.) with glabella only slightly rising above the fixigenal
fields (the cephalon is slightly tilted anteriorly on the figure,
suggesting erroneously that the occipital ring is higher than
the preoccipital glabella). As indicated by an internal cast
of the doublure on the left side, it is at least as wide as the
cephalic border.
Thorax composed of nine segments. Dorsal furrows
gently converging backwards and delimiting an axial lobe
slightly wider (tr.) than pleural lobes. Segments are very
similar to one another. Axial ring rather narrow (sag. and
exs.) all along. Pleura extending abaxially until fulcrum
then abaxially and posteriorly, especially in anteriormost
segments; rather well-incised pleural furrow running in a
straight direction from a short distance from dorsal furrow
to fulcrum, then rapidly vanishing; outer portion of pleura
apparently modified anteriorly into a smooth articulating
facet. In lateral view, pleurae straight and particularly high
posteriorly, where they represent almost two-thirds of total
height of the thorax, decreasing in height anteriorly while
outer portions become increasingly curved forwards. In
posterior or frontal views, axial lobe strongly arched dorsally.
Fulcrum subdivides pleura into a sub-horizontal inner portion
(c. 40 % of transversal length of pleura) and a strongly
downwards flexing (more than 70◦ ) outer portion.
Pygidium semi-circular in outline (Fig. 2b), slightly wider
(tr.) than long (sag.). Axis wide (tr.) anteriorly (c. 40 %
of the maximal width of the pygidium) but increasingly
tapering backwards; apparently 10 + 1 axial rings delimited
by broad axial furrows that rapidly shallow posteriorly; no
conspicuous post-axial furrow. As far as it can be observed,
the pleural field almost reaches the pygidial margin and
therefore a border, if present, may be either narrow (tr.)
or poorly differentiated from the rest of the pleura; seven,
probably eight pleural ribs defined by broad and shallow
pleural furrows increasingly curving backwards from front to
rear; up to three extremely faint furrows that occur abaxially
along the posterior margin of the first three ribs may represent
interpleural furrows, suggesting a subdivision of the rib into a
Late Palaeozoic trilobites of Iran and Armenia
1029
Figure 2. Persia praecox gen. nov. sp. nov.; Tournaisian, Gnathodus typicus conodont biozone, Shishtu Formation, Howz-e-Dorah
area, South Shotori Range, Eastern Iran. (a–e) EUIT 7-850, holotype specimen, right lateral, dorsal, left lateral, frontal and posterior
views.
broad (exs.) anterior branch and a very narrow (exs.) posterior
branch. Laterally (Fig. 2a, c), axis low, gently lowering
rearwards until almost merging with the post-axial region
(only a small piece of the latter is preserved); anteriormost
axial rings are moderately convex dorsally. Pleural region
high (c. 80 % of total height) and provided with a faceted
antero-lateral corner, which probably received the pleural
tip of the posteriormost thoracic segment during enrollment.
In posterior view (Fig. 2e), axis low and broadly elliptical in
profile. Pleural region subdivided into a horizontal inner third
and strongly flexed downwards (60◦ ) outer two-thirds. Mould
of inner surface of the doublure indicates that it is broad,
almost reaching the level of the tip of the axis sagittally.
Sculpture: seven to ten low terrace ridges run sub-parallel
to the margin on the cephalic border; at least nine of them
also occur on the pygidial doublure. Otherwise, no sculptural
features could be observed with confidence; granules of
various sizes can be observed here and there (e.g. Fig. 2d),
but no clear distribution pattern could be identified, which
suggests they may be preservational in origin.
Subfamily DITOMOPYGINAE Hupé, 1953
Genus Acropyge Qian, 1977
Type species. Acropyge multisegmenta Qian, 1977, from the
upper Changhsingian of Guizhou, South China.
Diagnosis.
See Owens, 1983, p. 28.
Acropyge encrinuroides (Weber, 1944)
Figure 3a, d, g, j, m
1944 Pseudophillipsia (Anisopyge) (?) encrinuroides
Weber, pp. 14, 23, 24, pl. 2, fig. 11a, b.
1978 Acropyge lanceolata Kobayashi & Hamada, p. 160,
fig. 5a, b.
1981 Acropyge lanceolata Kobayashi & Hamada; Hahn
& Hahn, pp. 220, 222, table 1.
1983 Acropyge encrinuroides (Weber); Owens, p. 29.
1983 Acropyge lanceolata Kobayashi & Hamada; Owens,
p. 29.
1984 Acropyge lanceolata Kobayashi & Hamada; Kobayashi & Hamada, p. 73, pl. 14, fig. 5a, b.
Materials, localities and horizons. Poorly preserved holotype pygidium (VSEGEI 83–5217, Fig. 3m); probably
Wordian, vicinity of Ogbin village, Vyots Dzor Province,
southern Armenia (label 7 on Fig. 1). Well-preserved
pygidium (PA 16766, Fig. 3a, d, g, j); uppermost part of Unit
1 of Taraz (1971), Surmaq Formation, Capitanian (Midian;
Kobayashi & Ishii, 2003), southwestern part of the Kuh-ehambast Range, Abadeh area (Fars Province), Central Iran
(label 1 on Fig. 1).
Diagnosis. A species of Acropyge displaying the following
combination of characters: pygidium escutcheon-shaped,
mucronate, particularly flat; axis strongly and evenly narrowing (tr.) posteriorly, not reaching border, highest at the fifth
or sixth-most anterior ring, devoid of muscles scars, with 23
± 1 axial rings; long and large post-axial ridge; 12 pleural
ribs, pleural furrows slightly curved rearwards abaxially and
running almost sub-parallel to sagittal axis posteriorly.
Remarks. When they described Acropyge lanceolata from
the Capitanian of Central Iran (Fig. 3a, d, g, j), Kobayashi
& Hamada (1978) compared it to the type species A.
multisegmenta, which had just been discovered in the
Changhsingian of South China (Qian, 1977). However, they
did not consider a very similar pygidium of equivalent
age that had been found in the neighbouring Armenia and
described as Pseudophillipsia (Anisopyge) (?) encrinuroides
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R . L E R O S E Y- AU B R I L
Figure 3. (a, d, g, j, m) Acropyge encrinuroides (Weber, 1944). (a, d, g, j) SMF 90658 (gypsum cast; original PA 16766, University
of Tokyo), pygidium with deformed right pleura; Capitanian, Surmaq Formation, Abadeh area, Kuh-e-hambast Range, Central Iran.
(a) Dorsal, (d) right lateral, (g) posterior, (j) left lateral views. (m) VSEGEI 83-5217, poorly preserved holotype pygidium in dorsal
view; Wordian (?), Gnishik Formation (?), vicinity of Ogbin, southern Armenia. (b, c, e, f, h, i, k, l) Acropyge weggeni Hahn & Hahn,
1981; Wuchiapingian, Nesen Formation, vicinity of Yush, Central Alborz Mountains, Northern Iran. (b, e, h, k) internal mould (latex)
of the holotype pygidium (SMF 282251 ) in (b) dorsal, (e) right lateral, (h) posterior and (k) left lateral views. (c, f, i, l) SMF 282252 ,
small partially broken pygidium on the same block in (c) dorsal, (f) right lateral, (i) posterior and (l) left lateral views. (n, o) Acropyge
sp. A, PIN 2321-19 (specimen missing), reproduced from Arkhipova (1965, pl. 45, fig. 2) with the permission of the Palaeontological
Institute, Moscow, slightly exfoliated pygidium possibly broken behind axis in (n) dorsal and (o) right lateral views; Wordian, Gnishik
Formation, Vedi area, southern Armenia. Scale bars = 5 mm, except for (c), (f), (i) and (l), where scale bars = 1 mm.
Late Palaeozoic trilobites of Iran and Armenia
by Weber (1944; Fig. 3m herein). This specimen was then
successively ignored by Yin (1978) and Hahn & Hahn (1981).
More surprisingly, it has also been mistreated by Kobayashi
& Hamada (1984), despite the fact that Owens (1983) had
just recognized in this specimen all the characteristics of
Acropyge and had accordingly assigned Weber’s species to
this genus. Actually, the holotypes of A. lanceolata and A.
encrinuroides are extremely similar with regards to all the
characters that can be observed on both specimens (compare
Fig. 3a and 3m). The only noticeable difference is the absence
of a border in the holotype of A. encrinuroides but this is
owing to bad preservation, the specimen being broken along
its left margin, while its right one apparently remains overlain
by the surrounding matrix. Accordingly, A. lanceolata is here
considered a junior synonym of A. encrinuroides.
Discussion. The escutcheon shape, the particularly flat
profile and the thick and long (sag.) post-axial ridge are
some traits, among others, that easily enable this species to
be distinguished from A. brevica Yin, 1978, A. multisegmenta
Qian, 1977 and A. weggeni Hahn & Hahn, 1981.
Acropyge weggeni Hahn & Hahn, 1981
Figure 3b, c, e, f, h, i, k, l
1981 Acropyge weggeni Hahn & Hahn, pp. 219–22,
table 1, pl. 1, fig. 1a–d.
1981 Acropyge? sp. indet. Hahn & Hahn, p. 223, pl. 1,
fig. 2.
1983 Acropyge weggeni Hahn & Hahn; Owens, p. 29.
1984 Acropyge weggeni Hahn & Hahn; Kobayashi &
Hamada, p. 72.
Material, locality and horizon. The external mould of a
large and well-preserved pygidium (holotype, SMF 282251 ,
Fig. 3b, e, h, k) and a small, partially broken pygidium on
the same block (SMF 282252 , Fig. 3c, f, i, l); Nesen Formation, Wuchiapingian, Central Alborz Mountains, near Yush
village (36◦ 13.05 N, 51◦ 42.20 E; Mazandaran Province),
Northern Iran (label 4 on Fig. 1).
A species of Acropyge displaying the
following combination of characters: pygidium elongate,
ogival but not mucronate; axis with 28 axial rings, bearing
two rows of paired muscle scars and a sagittal furrow at
its posterior tip; post-axial ridge large but short; 12 pleural
ribs; border narrow (tr.) and hardly widening (sag. and exs.)
posteriorly; single row of granules along posterior margin of
each axial ring and each pleural rib.
Emended diagnosis.
Remarks. After comparison with juvenile pygidia of
Pseudophillipsia (Carniphillipsia) lipara Goldring, 1957
deposited at the Senckenberg Research Institute, I follow
Hahn & Hahn (1981) in assigning the tiny juvenile pygidium
found in the Central Alborz (Fig. 3c, f, i, l) to the genus
Acropyge. This is justified chiefly by the fact that the
specimen, despite its very small size (less than 2.5 mm),
already exhibits a high number of axial rings (16/17)
and pleural ribs (13) and its general outline tends to be
ogival as in the holotype of A. weggeni. However, it seems
to me overcautious not to assign this specimen to A.
weggeni considering that (1) representatives of Acropyge are
extremely rare, (2) this species is the only species of Acropyge
known from this locality, and (3) this small pygidium and the
holotype of this species are on a single small rock sample.
Discussion. A. weggeni is easily differentiated from the
other species of Acropyge by its greater number of axial
rings (28). It more closely resembles A. encrinuroides, in
particular in having a large post-axial ridge, but this ridge is
1031
shorter than in the Guadalupian taxon. Moreover, it has an
ogival not escutcheon-like outline and rows of muscle scars
on the axis. A. brevica Yin, 1978 and A. multisegmenta Qian,
1977 exhibit a much narrower (tr.) axis and a very different
disposition of the pleural furrows (e.g. posteriormost ones
are not sub-parallel to the dorsal furrows), which permits
them to be easily distinguished from A. weggeni.
Acropyge sp. A
Figure 3n, o
1965 Pseudophillipsia armenica Weber; Arkhipova,
pp. 82–3, pl. 45, fig. 2a, b.
Material, locality and horizon. A slightly exfoliated pygidium, possibly broken behind axis (PIN 2321–19, missing
specimen; Fig. 3n, o); Gnishik Formation, Wordian, Vedi area
(Ararat Province), Southern Armenia (label 6 on Fig. 1).
Pygidium elongate and ogival in outline
(Fig. 3n), with a superficially re-entrant (likely to be damaged,
see remarks below) posterior margin reaching the axis.
Axis rather long (c. 80 % of the sagittal length of the
pygidium) but possibly slightly broken posteriorly; 23 axial
rings discernable, with three to five additional ones probably
present in poorly preserved posteriormost portion, all bearing
a pair of conspicuous muscle scars forming two rows that
subdivide the axis into three lobes; rather deep inter-ring
furrows of sigmoidal course from sagittal axis to dorsal
furrows. No post-axial region visible (likely broken, see
remarks below). Pleural field representing approximately
a third of the width (tr.) of the pygidium anteriorly and
composed of 11, possibly 12 ribs; pleural furrows particularly
deep, faintly curved, running abaxially and increasingly
backwards from front to rear. Pygidial border separated from
pleural field by a strong break in slope, narrow anteriorly,
but gently widening posteriorly. In lateral view (Fig. 3o),
axis and pleura are of about the same height; axis gently
declining and slightly flexing downwards posteriorly; axial
rings rather prominent dorsally, while pleural ribs are flattopped. Border flat. No posterior or ventral views available.
Sculpture: single row of granules apparently occurs along
posterior margin of each pleural rib.
Description.
Remarks. The pygidium discovered in the Permian outcrop
near Vedi fits well with the diagnosis of the genus Acropyge,
except in the fact that it lacks ‘a posterior margin extended
into point’ (Owens, 1983). Actually, the morphology of its
post-axial region is peculiar and to my knowledge is unique
amongst the Proetida. Only in a few Cambrian trilobites of
the family Burlingiidae can a re-entrant posterior margin
be observed that merges with the posterior tip of the axis
as in this specimen (Ebbestad & Budd, 2003). Moreover,
other characteristics of burlingiid pygidia rule out any close
phylogenetical relationships with the Permian taxon. This
raises the question of whether this unusual morphology
might have resulted from a breakage or from some kind of
teratological phenomenon. A rapid review of the literature
dealing with teratology in trilobites shows that a similar
modification of the post-axial region has never been reported
to date. I am, therefore, more inclined to believe it has
resulted from a breakage, despite the symmetrical and
curved margins of the supposedly broken area. This strange
morphology might be explained if the breakage lines have
followed the margins of a strong post-axial ridge near the
tip of the axis and then increasingly diverged posteriorly,
probably accommodating the topography of the border in the
immediate proximity of the ridge. The fact that the specimen
is missing in the collections of the Palaeontological Institute
of Moscow has prevented further observations from being
1032
R . L E R O S E Y- AU B R I L
made, but the small pygidium from the Alborz Mountains
tentatively assigned to Acropyge weggeni exemplifies, to
some extent, how a breakage in this region can appear
symmetrical (Fig. 3c, i). The presence of a posterior axial
ridge in the Armenian specimen is also probable, given its
close resemblance to the holotype of A. weggeni (compare
Fig. 3b and 3n).
The pygidium of Acropyge sp. A is best
compared with that of A. weggeni with which it has the
following characters in common: an elongate and ogival
outline, a similar structure and shape of the axis, comparable
shape and orientation of pleural furrows, a flat border,
a similar general profile and rows of granules along the
posterior margin of each rib (Fig. 3b, n). The Armenian
pygidium exhibits only 23 axial rings and 11 pleural ribs, but
considering the poor preservation of the posterior portion of
the axis and the likely breakage of the post-axial region,
it seems possible that in fact this specimen possessed a
similar number of axial rings (28) and pleural ribs (12) as
the holotype of A. weggeni. However, a few differences
between the two pygidia suggest they belong to different,
though phylogenetically close, species. Indeed, compared
to the holotype of A. weggeni, the Armenian specimen
has a slightly narrower (tr.) axis that only moderately
flexes downwards posteriorly (Fig. 3o), an apparently wider
(tr.) border posteriorly, flat-topped pleural ribs and it is
slightly flatter in general profile. It differs more from A.
encrinuroides, from which it can be differentiated by its
ogival outline and the presence of rows of muscle scars on the
axis, and even more from A. brevica and A. multisegmenta,
which have a much narrower (tr.) axis and a very different
structure of pleural segmentation.
Discussion.
Genus Pseudophillipsia Gemmellaro, 1892
Type species. Phillipsia sumatrensis Roemer, 1880, from
the Wordian of the western coast of Sumatra, Indonesia.
Diagnosis.
See Owens, 1983, p. 28.
Remarks. In the absence of cranidia, it is not possible to
discriminate the three subgenera of Pseudophillipsia. Indeed,
no diagnostic characters of Pseudophillipsia (Nodiphillipsia) concern the pygidial morphology (Lerosey-Aubril &
Angiolini, 2009), and the ranges of the number of axial
rings and pleural ribs in Pseudophillipsia (Carniphillipsia)
or Pseudophillipsia (Pseudophillipsia) frequently overlap
(Hahn & Brauckmann, 1975). The species concerned are,
therefore, attributed to Pseudophillipsia sensu lato (s.l.)
hereafter.
Pseudophillipsia (s.l.) armenica (Weber, 1944)
Figure 4a–f
1944 Pseudophillipsia armenica Weber, pp. 13, 23, pl. 2,
fig. 1a, b.
1965 Pseudophillipsia paffenholzi Weber; Arkhipova,
pp. 82–3, pl. 45, fig. 3a, b.
1978 Iranaspidion sagittalis Kobayashi & Hamada,
p. 158, fig. 4a–c.
1984 Iranaspidion sagittalis Kobayashi & Hamada;
Kobayashi & Hamada, pp. 68–9, pl. 14, fig. 4a–c.
The holotype pygidium,
which has its left side broken (VSEGEI 73–5217; Fig. 4a);
probably Wordian, vicinity of Ogbin village (Vyots Dzor
Province), southern Armenia (label 7 in Fig. 1). A pygidium
with both pleural fields slightly broken anteriorly (PIN 2321–
15, missing specimen; Arkhipova, 1965, pl. 45, fig. 3a, b and
Material, locality and horizon.
Fig. 4c, f herein); Gnishik Formation, Wordian, Vedi area
(Ararat Province), southern Armenia (label 6 in Fig. 1). A
well-preserved pygidium (PA 16765; Fig. 4b, d, e); upper part
of the Unit 1 of Taraz (1971), Surmaq Formation, Capitanian,
Abadeh area (Fars Province), Hambast Mountains, Central
Iran.
Diagnosis. A species of Pseudophillipsia displaying the
following combination of characters: axis downwardly flexed
anteriorly and especially posteriorly, 21–22 prominent axial
rings, 11 prominent pleural ribs, border narrow and inclined
about 30–40◦ outwards, transverse rows of coarse tubercles
on axial rings and abaxial two-thirds of pleural ribs.
Pygidium semi-circular in outline (Fig. 4e,
f). Axis long, reaching border furrow; 21–22 axial rings
bearing a pair of marked muscle scars, which form two
rows subdividing the axis into a wide (tr.) median lobe
and two narrow lateral lobes; deep and rather broad interring furrows slightly posteriorly deflected abaxially. Pleural
field representing about a third of the width (tr.) of the
pygidium anteriorly and composed of 11, possibly 12 ribs;
pleural furrows particularly deep, broad and sigmoidal;
well-developed antero-lateral extensions with articulating
facets (Fig. 4b). Border narrow anteriorly, slightly widening
posteriorly. In lateral view (Fig. 4a–c), axis hardly more
than a third of pygidial total height, downwardly flexed
anteriorly and especially posteriorly; axial rings and pleural
ribs particularly prominent. In posterior view (Fig. 4d), axis
semi-circular in cross-section, pleural fields composed of
a narrow horizontal adaxial part and a wider and strongly
sloping (50–60◦ ) downwards abaxial part; border inclined
outwards at c. 45◦ . Sculpture: transverse row of tubercles on
each axial ring as well as on the abaxial two-thirds of pleural
ribs.
Description.
Remarks. Except for the inclination of the border, which
could not be verified on the specimen figured by Arkhipova
(1965), all the characters of the diagnosis are exhibited by
the two pygidia from southern Armenia and the Iranian
specimen from Abadeh, justifying their reassignment to the
same species. The original assignment to Pseudophillipsia
paffenholzi of the specimen from the Vedi area is difficult to
explain, since this species can be easily differentiated from P.
(s.l.) armenica by its more elongate general outline, its flattopped axial rings or its greater number (12) of flat-topped
pleural ribs. Even if the two species bear similarly distributed
tubercles, these are coarse in P. (s.l.) armenica while they are
barely more than small granules in P. paffenholzi.
Our attribution of the isolated pygidium from the Abadeh
area to P. (s.l.) armenica instead of P. sagittalis (Kobayashi
& Hamada, 1978) raises the question of a possible synonymy
of the two species. As far as the pygidial morphology
is concerned, only two of the four specimens figured
by Kobayashi & Hamada (1978) can be considered: the
pygidium here reassigned to P. (s.l.) armenica (Fig. 4b, d,
e) and the holotype of P. sagittalis (Fig. 4n). The pygidium of
the holotype is damaged along most of its margin, especially
posteriorly, but compared with the other, it has a narrower
(tr.) border (where visible), and a narrower (tr.) axis with
straight and convergent axial furrows instead of increasingly
convergent ones. On the other hand, the two specimens share
similar pleural furrow courses, and prominent axial rings
and pleural ribs, which might indicate they belong to the
same species. Unfortunately, the sculpture or lateral profile
of the holotype can hardly be appreciated from Kobayashi &
Hamada’s figures. Until the holotype specimen of P. sagittalis
is found again, or its loss confirmed, it seems more reasonable
to conserve this taxon as a separate species, but it should be
Late Palaeozoic trilobites of Iran and Armenia
1033
Figure 4. (a–f) Pseudophillipsia (s.l.) armenica Weber, 1944. (a) VSEGEI 73–5217, holotype pygidium in right lateral (slightly
oblique) view; Wordian (?), Gnishik Formation (?), vicinity of Ogbin, southern Armenia. (b, d, e) SMF 90659 (gypsum cast; original
PA 16765, University of Tokyo), pygidium in (b) right lateral, (d) posterior and (e) dorsal views; Capitanian, Surmaq Formation, Abadeh
area, Kuh-e-hambast Range, Central Iran. (c, f) PIN 2321–15 (specimen missing), reproduced from Arkhipova (1965, pl. 45, fig. 3)
with the permission of the Palaeontological Institute, Moscow, pygidium partially broken anteriorly in (c) right lateral and (f) dorsal
views; Wordian, Gnishik Formation, Vedi area, southern Armenia. (g–j, l, m) Pseudophillipsia (s.l.) aff. caucasica; Wuchiapingian,
Nesen Formation, vicinity of Yush, Central Alborz Mountains, Northern Iran. (g, i, l) SMF 28226, partial pygidium in (g) posterior,
(i) dorsal and (l) right lateral, slightly oblique views. (h, j, m) latex cast made from external mould (SMF 28227) of specimen SMF
28226 in (h) posterior, (j) dorsal and (m) right lateral views. (k, n) Pseudophillipsia (Carniphillipsia) sagittalis (Kobayashi & Hamada,
1978), PA 16764 (specimen missing), enrolled holotype specimen, dorsal views of (k) cephalon and (n) pygidium; Capitanian, Surmaq
Formation, Abadeh area, Kuh-e-hambast Range, Central Iran. Reproduced from the Proceedings of the Japan Academy, Series B, with
the permission of the Japan Academy. Scale bars = 5 mm.
1034
kept in mind that it might prove to be a junior synonym of
P. (s.l.) armenica. In that case, the Iranian specimens would
then permit the cephalic and thoracic morphology of this
species to be known.
Discussion. The partial pygidium of Pseudophillipsia (s.l.)
caucasica (Weber, 1944) found in the North Caucasus
(Weber, 1944, pl. 2, fig. 4a–c) somewhat resembles P.
(s.l.) armenica in having the following traits: a strongly
downwardly flexed axis posteriorly, prominent axial rings
bearing a row of tubercles, 11 prominent pleural ribs also
bearing tubercles, and a similar curvature of the pleural field
in posterior view. However, it has more axial rings (25),
a more complex (sigmoidal) course of inter-ring furrows,
pleural tubercles restricted to the fulcral region and an inflated
border, all characters that demonstrate it is a different species.
Pseudophillipsia (s.l.) caucasica (Weber, 1944)
1944 Pseudophillipsia elegans var. caucasicus Weber,
pp. 12–13, table 2.
A pygidium of unknown state
of preservation; probably Wordian, vicinity of Ogbin village
(Vyots Dzor Province), southern Armenia (label 7 in Fig. 1).
Material, locality and horizon.
Remarks. In the Russian part of his work, Weber (1944,
pp. 12–13, table 2) mentioned that a pygidium of P. caucasica
has been found in the Ogbin area. This information is
completely omitted in the English section of his text dealing
with this species. As only the type specimen from the North
Caucasus is figured, it is difficult at present to confirm or deny
the claimed occurrence of P. (s.l.) caucasica in the Permian of
southern Armenia. A rather similar taxon, Pseudophillipsia
(s.l.) aff. caucasica, apparently occurred in Central Alborz
and therefore it is possible that the specimen mentioned by
Weber might in fact belong to this Iranian species.
Pseudophillipsia (s.l.) aff. caucasica (Weber, 1944)
Figure 4g–j, l, m
1981 Iranaspidion sp. Hahn & Hahn, p. 223, table 1,
fig. 3.
Material, locality and horizon. A partial pygidium (SMF
28226; Fig. 4g, i, l) and its external mould (SMF
28227; Fig. 4h, j, m); Nesen Formation, Wuchiapingian,
Central Alborz Mountains, near Yush village (36◦ 13.05 N,
51◦ 42.20 E; Mazandaran Province), Northern Iran (label 4
in Fig. 1).
Pygidium of roughly semi-circular outline
(Fig. 4i, j). Axis particularly long, overhanging the border
posteriorly and delimited laterally by straight axial furrows
evenly converging backwards; 17 axial rings discernable
corresponding laterally to the six posteriormost pleural ribs,
which permits an estimation of a total of 21–22 axial rings
for the whole axis; two rows of faintly impressed muscle
scars; inter-ring furrows of strongly sigmoidal course from
the sagittal axis to dorsal furrows. Pleural field of about the
same width (tr.) as axis anteriorly, but rapidly narrowing
(tr.) posteriorly; ten pleural ribs; pleural furrows deep but
narrow, with most of them having a particularly well-marked
sigmoidal course. Pygidial border separated from pleural
field by a strong break in slope, rather narrow anteriorly
but significantly widening posteriorly until the pygidial midlength (sag.), remaining of roughly equal width thereafter.
In lateral view (Fig. 4l, m), posterior part of axis strongly
flexed downwards; pleural field especially high, composed of
prominent pleural ribs, and provided with a smooth anterolateral projection. Border rather thick but flat dorsally. In
Description.
R . L E R O S E Y- AU B R I L
posterior view (Fig. 4g, h), axis elliptical in section (tr.),
bearing a very short sagittal furrow at its posterior tip. Pleural
ribs particularly prominent at fulcrum, which subdivides
the pleural field into a roughly horizontal inner third and
strongly flexing downwards (more than 60◦ ) outer two-thirds.
A slightly inflated area split in two by a broad sagittal furrow
occurs behind the axial tip. Ventral morphology unknown.
Sculpture: a single row of up to seven low tubercles on
the outer two-thirds of the four most posterior pleural ribs;
numerous thin terrace ridges along border margin, which are
sub-parallel to it posteriorly, while they have a more exsagittal
course anteriorly.
Discussion. The specimen is best compared with P. (s.l.)
caucasica (Weber, 1944, pl. 2, fig. 4a–c), with which it
shares similar inter-ring and pleural furrow courses, an axis
overhanging the border and bearing a short sagittal furrow at
its tip, broad pleural ribs that are prominent at fulcrum and
strongly sloping downwards abaxially, and a slightly inflated
area on the border behind axis. However, it differs from the
North Caucasian (and possibly Armenian) species in having
only 10 instead of 11 pleural ribs and a rather wide border
posteriorly. This character might be explained by ontogenetic
differences, the Iranian specimens being some 30 % larger
than the pygidium described by Weber (1944). The difference
in the number of pleural ribs, however, is more problematic,
which explains why I refer to this Iranian specimen as P. (s.l.)
aff. caucasica.
As discussed above, P. (s.l.) caucasica and P. (s.l.)
armenica are morphologically, and in all likelihood, phylogenetically close species. The similarities described between
these two species can also be mentioned for P. (s.l.) aff.
caucasica. In this form, however, the small tubercles are
not restricted to the fulcrum but borne by the entire outer
portion of the pleural ribs. Moreover, this specimen exhibits
medially a swollen area on the border split in two by a
furrow (Fig. 4h), a feature that can also be observed on
the isolated pygidium found in the Abadeh area (Fig. 4d)
and herein assigned to P. (s.l.) armenica. However, like P.
(s.l.) caucasica, the pygidium from the Nesen Formation
has a more complex course of inter-ring and pleural furrows
than that of P. (s.l.) armenica, and a thick border that is
not as inclined. In addition, the tubercles on the pleurae are
small and only visible on the posteriormost pleural ribs, the
axis notably overhangs the border posteriorly and the border
strongly widens posteriorly, all characters that exclude a
possible attribution of this specimen to P. (s.l.) armenica.
Neverthess, the similarities between P. (s.l.) aff. caucasica,
P. (s.l.) caucasica and P. (s.l.) armenica suggest that all these
forms originated from a common ancestor.
Pseudophillipsia (s.l.) parvizii sp. nov.
Figure 5
Well-preserved holotype
pygidium (SMF 90686; Fig. 5a, c, e, g) and a wellpreserved paratype pygidium (SMF 90687; Fig. 5b, d, f,
h); both from a dark-grey sandy wackestone with gastropods, brachiopods (e.g. productoids), bryozoa (fenestellids),
crinoids, ostracods, rare trilobites and foraminifers, Dalan
Formation, Wuchiapingian (indicated by Codonofusiella ex
gr. tenuissima), Dena Mountain, about 58 km northwest of
Yasouj (Kohgilouye and Boyrahmad Province), SE Iran (label
3 in Fig. 1).
Material, locality and horizon.
In reference to M. Parvizi (University of Payame Noor) who collected the two specimens and made them
available for study.
Etymology.
Late Palaeozoic trilobites of Iran and Armenia
1035
Figure 5. Pseudophillipsia (s.l.) parvizii sp. nov.; Wuchiapingian, Dalan Formation, Dena Mountain, about 58 km northwest of Yasouj,
SE Iran. (a, c, e, g) SMF 90686, holotype pygidium in (a) dorsal, (c) right lateral, (e) posterior and (g) left lateral views. (b, d, f, h)
SMF 90687, paratype pygidium in (b) dorsal, (d) right lateral, (f) posterior and (h) left lateral views. Scale bars = 5 mm.
Diagnosis. A species of Pseudophillipsia displaying the following combination of characters: anterior half of pygidium
almost parallel-sided, 21 axial rings, 13 pleural ribs, pleural
field with strongly downwards flexed outer half, presence of
antero-lateral projections, pygidial border wide and of equal
width in posterior half of pygidium, then strongly narrowing
anteriorly.
Pygidium rather high and of strongly parabolic
outline, almost parallel-sided anteriorly in dorsal view
(Fig. 5a, b). Axis long (c. 90 % of the sagittal length of
the pygidium), gently narrowing (tr.) backwards, rounded
posteriorly, and reaching the posterior border; 21 axial rings
delimited by rather deep inter-ring furrows of sigmoidal
course from the sagittal axis to dorsal furrows, all but the very
last rings bearing a pair of flat and slightly depressed muscle
scars, the alignment of which leads to a transversal trilobation
of the axis. Pleural field rather narrow (e.g. representing less
than a third of the width of the pygidium anteriorly) and
composed of 13 ribs, which are delimited by deep pleural
furrows; most of these furrows are weakly sigmoidal in shape.
Pygidial border separated from pleural field by a strong break
in slope, increasing in width posteriorly at the expense of
this pleural field until the mid-length (sag.) of the pygidium,
then remaining of equal width; faint post-axial ridge bearing
an almost inconspicuous sagittal furrow. In lateral view
(Fig. 5c, d, g, h), axis rather low compared with pleural
field; it is strongly flexed downwards and lowers posteriorly
behind axial ring 3, but also to a lesser degree anteriorly
in front of this ring (‘sagittally arched’ axis); axial rings
prominent dorsally while pleural ribs are flat-topped. Smooth
and slightly depressed area occurs on the antero-lateral
region of the pleura and it extends anteriorly on a forward
projection of the latter. Border rather thick but flat dorsally.
In posterior view (Fig. 5e, f), axis elliptical in transverse
section. Fulcrum subdivides the pleural field into a roughly
horizontal inner half and a strongly downwardly flexing
(more than 60◦ ) outer half. Faint sagittal furrow occurs on
the posterior tip of the axis. Ventral morphology unknown.
Sculpture: numerous thin terrace ridges occur on the outer
part of the dorsal side of the pygidial border; they run
backwards following an almost exsagittal course, therefore
rapidly meeting the border margin, and then they continue
their course on the lateral side and possibly further on the
doublure.
Description.
Although only known from pygidia, the new
species already exhibits very distinctive characters compared
with all the other representatives of Pseudophillipsia.
However, with its wide pygidial border, P. (s.l.) parvizii sp.
nov. vaguely resembles Pseudophillipsia (Carniphillipsia)
rotunda Hahn & Hahn in Hahn, Hahn & Ramovš, 1990
from the Sakmarian of Slovenia. The two taxa also share
similar numbers of axial rings (21 v. 20) and pleural ribs
(13 in both), strongly downwardly flexed outer pleurae and
prominent dorsal axial rings. However, the pygidium of P.
(C.) rotunda is semi-circular in outline and devoid of large
antero-lateral expansions. It also exhibits a fine granulation
on the pleurae and, at least in P. (C.) rotunda noricana, its
axis is trapezoidal in transverse section (Hahn et al. 2002),
all characters that prevent any possible confusion with the
new species from Iran. P. (C.) pocivalensis Hahn et al. 2002,
another taxon from the Permian of Slovenia, exhibits a wide
pygidial border as in P. (s.l.) parvizii sp. nov. and P. (C.)
rotunda, but as in the latter species, it lacks antero-lateral
expansions and bears small granules on the axis and the
pleurae (Hahn et al. 2002, pl. 1, fig. 11). Compared with
the new species, it also has fewer axial rings (c. 18) and
pleural ribs (10), an axis that is much more strongly flexed
downwards posteriorly and a dorsally concave rather than flat
pygidial border. In the Permian of Iran, P. (s.l.) armenica from
the Capitanian of the Abadeh region (Fig. 4b, d, e) and P. (C.)
sp. A (Fig. 6i, j, l, n, o) from the Guadalupian of Chahriseh
have in common with the new species a similarly strongly
vaulted pygidium resulting from strongly downwardly flexed
outer pleurae, well-incised pleural furrows and antero-lateral
expansions. However, the pygidia of these taxa differ from
that of P. (s.l.) parvizii sp. nov. by the following traits: a
less parabolic outline, a narrower border, a more backwards
tapering axis, straight inter-ring furrows, more prominent
pleural ribs, and the presence of granules on both the axis
and the pleurae.
Two important characteristics of the new species are clear
adaptations to enhanced capacities for volvation: the narrow
pygidial border anteriorly that abruptly widens at about the
mid-length (exs.; Fig. 5a, b) and the antero-lateral expansions
of the pygidium (Fig. 5d, g, h). The first feature is known in
trilobites such as Ameura missouriensis (e.g. Hahn & Hahn,
1987, pl. 5, fig. 1) or Pseudophillipsia (Nodiphillipsia) aff.
obtusicauda (Lerosey-Aubril & Angiolini, 2009, fig. 2h). In
these forms, it permits the anterior part of the pygidium to
Discussion.
1036
R . L E R O S E Y- AU B R I L
be enclosed within the cephalic capsule during enrollment,
which prevents lateral shearing (Lerosey-Aubril & Angiolini,
2009), and it seems reasonable to assume that P. (s.l.) parvizii
sp. nov. enrolled its body in a comparable way. A similar mode
of enrollment has been described in some rare Ordovician
pterygometopids (Ludvigsen & Chatterton, 1982) and more
recently in the Devonian scutelluid Paralejurus rehamnanus
(Feist, Lerosey-Aubril & Johnson, 2010). However, it is
within the proetoids, particularly the Late Palaeozoic forms,
that it has been most commonly observed (Lerosey-Aubril
& Angiolini, 2009). This atypical mode of volvation might
have been especially efficient for protective purposes, since
it combined the resistance against dorso-ventral pressures
of the sphaeroidal mode with the resistance against lateral
shearing of the spiral type (Lerosey-Aubril & Angiolini,
2009). The other morphological particularity related to
enrollment of the new species is the presence of antero-lateral
expansions of the pygidium. Similar features are observed in
other Late Palaeozoic proetoids: Anisopyge perannulata (e.g.
Brezinski, 1992, figs 10.13–24), Hentigia bulbops (Haas,
Hahn & Hahn, 1980, pl. 3, figs 1–7) and H. planops (Haas,
Hahn & Hahn, 1980, pl. 5, figs 7, 8), Pseudophillipsia (s.l.)
armenica (e.g. Kobayashi & Hamada, 1978, fig. 4a–c), P.
(C.) paffenholzi (Weber, 1944, pl. 2, fig. 6) and possibly also
in P. (Pseudophillipsia) binodosa (Kobayashi & Hamada,
1984, pl. 6, fig. 10). These antero-lateral expansions of the
pygidium have also been described recently in the scutelluid
Paralejurus rehamnanus (Feist, Lerosey-Aubril & Johnson,
2010). Here, they received dorsally the outer portions of
the pleurae of at least the four posterior thoracic segments
during volvation and they abutted the cephalic doublure. This
probably permitted a reinforcement of the body capsule and
a similar role can be hypothesized for the proetoids listed
above as well as for P. (s.l.) parvizii sp. nov. These pygidial
particularities of the new species exemplify particularly
well how important volvation has remained for trilobites in the Late Palaeozoic (Lerosey-Aubril & Angiolini,
2009).
Subgenus Pseudophillipsia (Carniphillipsia) Hahn &
Brauckmann, 1975
Type species. Pseudophillipsia ogivalis Gauri, 1965, from
the Lower Kasimovian (Pennsylvanian) of the Zoellner Ridge
near Waidegger-Alm, Carnic Alps, Austria.
Diagnosis.
See Hahn & Hahn, 1987, p. 588.
Pseudophillipsia (Carniphillipsia) paffenholzi (Weber,
1944)
Figure 6a–c, e, f
1939 Pseudophillipsia paffenholzi; Weber, p. 199, pl. 46,
fig. 16a, b.
1944 Cyphinium (?) paffenholzi n. sp.; Weber, pp. 10–11,
21–2, pl. 2, figs 5–7, 10.
1965 Pseudophillipsia armenica Weber; Arkhipova,
pp. 82–3, pl. 45, fig. 1a, b.
1975 Pseudophillipsia (Pseudophillipsia) paffenholzi
Weber; Hahn & Brauckmann, p. 119.
1978 Pseudophillipsia paffenholzi (Weber); Kobayashi &
Hamada, p. 160.
1983 Pseudophillipsia paffenholzi (Weber); Owens, p. 28.
1984 Pseudophillipsia paffenholzi (Weber); Kobayashi &
Hamada, p. 69.
A cephalo-thorax with
anterior border broken, librigenae slightly tilted downwards
below the cranidium and only the five anteriormost thoracic
Material, locality and horizon.
segments (VSEGEI 49–5217; Fig. 6a), which I herein
designate as the lectotype; a thoraco-pygidium, apparently
complementary to the cephalo-thorax, with left half of
the five posteriormost thoracic segments and most of the
pygidium preserved (VSEGEI 50–5217; Fig. 6e), which
should be considered as a paratype; both specimens are from
an unknown outcrop of probable Wordian age along the Vedi
River, ‘some 2 km above Dagnas village’, Vedi area (Ararat
Province), southern Armenia (label 6 in Fig 1). A pygidium
(VSEGEI 51–5217; Fig. 6f), slightly broken anteriorly on
the left side, and a partial right librigena (VSEGEI 52–
5217; not refigured herein); both specimens are from a
second unknown outcrop of probable Wordian age along
the Vedi River, ‘some 1.5 km above Dagnas village’, Vedi
area (Ararat Province), southern Armenia (label 6 in Fig. 1).
A cephalon (PIN 2321–18, missing specimen; Arkhipova,
1965, pl. 45, fig. 1a, b; Fig. 6b, c herein), with the medial part
of the glabella exfoliated anteriorly and broken posteriorly, is
also tentatively assigned to this species; Gnishik Formation,
Wordian, Vedi area (Ararat Province), southern Armenia
(label 6 in Fig. 1).
Diagnosis. A species of Pseudophillipsia (Carniphillipsia)
displaying the following combination of characters: glabella
elongate, with narrow (tr.), parallel-sided and flat-topped
posterior two-fifths; median preoccipital lobe wide (tr.) and
trapezoid; L1 long (exs.); two pairs (S2–S3) of faint and
short anterior glabellar furrows; anterior border particularly
narrow (sag. and exs.); pygidium with 25 + 1 flat-topped
axial rings; 12 flat-topped pleural ribs; pleural furrows
sigmoidal; tiny tubercles occur medially along posterior
margin of preoccipital lobe, all segments from occipital ring
to last pygidial axial ring, and abaxial four-fifths of pleural
ribs.
The following description aims to complete
Weber’s original one (1944) or to emphasize some features
that are of interest for the recognition of the species and its
affinities with other taxa.
Glabella elongate, narrow (tr.) and parallel-sided along
posterior two-fifths, then moderately widening forwards;
posterior margin of occipital ring convex (Fig. 6a); SO gently
and moderately curved forwards; rather deep S1 defining
trapezoid and wide (tr.) median preoccipital lobe and long
(exs.), rather narrow and somewhat triangular L1; S2–3 short
and faint; anterior border particularly narrow; fixigenal field
narrow (tr.) in post-ocular region, not visible in pre-ocular
region. Eye kidney-shaped and long (exs.), extending from
the mid-length (exs.) of L1 to mid-length (exs.) of glabella;
librigenal border rather narrow; posterior border strongly
widening owing to abrupt backward deflection of cephalic
posterior margin abaxially. In lateral view (Weber, 1944, pl. 2,
fig. 5b), glabella roughly flat-topped posteriorly, then strongly
and increasingly sloping forwards; lateral border rather high.
In the thorax, axial ring wide (sag. and exs.), composed of
a postannulus of even width all along (sag. and exs.) and
a preannulus and an articulating half-ring both about half
the length (sag.) of postannulus (Weber, 1944, pl. 2, figs 5b,
6b; Fig. 6a, e herein); pleural furrows confined to fulcral
region. Pygidium of strongly parabolic outline (Fig. 6e, f),
subdivided into axial and pleural lobes of sub-equal width
(tr.); axis subdivided into a median part and two abaxial
parts about a third width (tr.) of it by two rows of muscle
scars; 25 + 1 axial rings delimited by deep inter-ring furrows
slightly curved backwards from the sagittal line to muscle
scar and then running backwards until the axial furrow;
antero-lateral projection on pleura receiving outer portion
of two posteriormost thoracic pleurae; 12 flat-topped pleural
ribs (not 11 as claimed by Weber); pleural furrows deep
Description.
Late Palaeozoic trilobites of Iran and Armenia
1037
Figure 6. (a–c, e, f) Pseudophillipsia (Carniphillipsia) paffenholzi (Weber, 1944). (a, e) Wordian (?), unknown outcrop along the Vedi
River, Vedi area, southern Armenia. (a) VSEGEI 49-5217, lectotype cephalo-thorax in dorsal view. (e) VSEGEI 50-5217, paratype
thoraco-pygidium probably complementary to VSEGEI 49-5217 in dorsal view. (b, c) PIN 2321-18 (specimen missing), partial
1038
and sigmoidal; no interpleural furrows discernible; pygidial
border regularly and significantly widening from front to rear.
Laterally (Weber, 1944, pl. 2, fig. 6b), pygidium rather high
mainly owing to the pleural field, which is twice as high as
the axis; axis is highest at axial ring 3, then it slightly and
gently flexes downwards posteriorly and anteriorly.
Sculpture: tiny granules along posterior margin of median
preoccipital lobe, occipital lobe, thoracic and pygidial axial
rings, and the abaxial four-fifths of pygidial pleural ribs; thin
terrace ridges with exsagittal course occur near lateral margin
of pygidial border.
Remarks. I have chosen to follow Kobayashi & Hamada
(1978, 1984) and Owens (1983) instead of Hahn &
Brauckmann (1975) in considering that this species was
described by Weber in 1944, not 1939. Indeed, even if the
name of the species was proposed and two specimens were
figured in 1939, it seems Weber himself considered he was
describing the taxon for the first time in 1944 as shown by
the mention of ‘n. sp.’ after the name of the species in his
text and figure captions. Moreover, I did not follow Hahn
& Brauckmann (1975) in their attribution of the species to
the subgenus Pseudophillipsia (Pseudophillipsia), in so far
as the cranidium of this species (Fig. 6a) exhibits faint and
short glabellar furrows S2 and S3, indicating it belongs to
the subgenus Pseudophillipsia (Carniphillipsia).
Discussion. At first sight, the cephalon figured by Arkhipova (1965, pl. 45, fig. 1a, b; Fig. 6b, c herein) may look rather
different to the lectotype in its general proportions (Fig. 6a),
but this stems from the fact that in the lectotype the anterior
fixigenae are broken and the librigenae are tilted downwards.
Moreover, not only were the two specimens discovered in
the same geographical area, but they both exhibit a narrow
(tr.) and parallel-sided posterior portion of the glabella with
elongate L1 (exs.), a particularly narrow anterior border,
a similar librigenal morphology and a comparable profile
of the anterior portion of the glabella. The only noticeable
difference between the two specimens is the slightly greater
widening (tr.) anteriorly of the anterior portion of the glabella
in Arkhipova’s cephalon, which might be simply suggested
by the different preservations of the two specimens or related
to intra-specific variation.
Otherwise, P. (C.) paffenholzi is best compared with P. (C.)
sp. A from the Middle–Late Permian of the Chahriseh area
(Fig. 6i, j, l, n, o) and P. (C.) chongqingensis Lu, 1974 from
the Lopingian of SW China (Fig. 6d, g, h, k, m, p). These three
taxa share the following characters: an elongate glabella with
parallel-sided posterior portion, a rather wide (tr.) and long
(sag. and exs.) median preoccipital lobe, elongate (exs.) L1,
a narrow and high cephalic anterior border, and a pygidium
provided with flat-topped pleural ribs. This combination of
characters may indicate these taxa constitute a distinct clade
within the subgenus Pseudophillipsia (Carniphillipsia),
which I informally call the ‘paffenholzi-group’. However,
P. (C.) paffenholzi differs from these two forms in having a
shorter parallel-sided portion of the glabella (posterior twofifths instead of posterior half), a convex posterior margin of
R . L E R O S E Y- AU B R I L
the occipital ring, a gently and moderately curved forwards
SO, a slightly triangular L1, only two (S2–3) anterior
glabellar furrows discernible, a more complex course of interring and pleural furrows, a significant but regular widening of
the pygidial border rearwards, and the particular distribution
on each tagmata of the tiny granules.
As this work refigures Lu’s original material, it should
be mentioned that both the cephalon and the thoracopygidium assigned to P. (C.) chongqingensis display unusual
features. Indeed, the main glabellar lobe of the holotype
cephalon exhibits poorly defined furrows that shallow and
meet anteriorly to isolate a slightly inflated median area
(Fig. 6d, h). Similarly, poorly defined furrows associated with
a somewhat locally deformed cuticle are sometimes observed
in asteropygine trilobites (e.g. Morzadec, 1981, pl. 35, fig. 1a;
pers. obs.). The course of these unusual furrows in P. (C.)
chongqingensis strikingly recalls the distribution pattern
of the frontal auxiliary impressions (FAIs) in Phillipsia
(Lerosey-Aubril, Hegna & Olive, 2011). Accordingly, I
interpret these furrows as resulting from the attachments of
extrinsic muscles of a pouch-like modified portion (‘crop’)
of the foregut. If Lerosey-Aubril, Hegna & Olive (2011) are
right in suggesting a relationship might have existed between
the distribution of FAIs and the shape/size of this putative
crop, the latter might have been particularly large in P. (C.)
chongqingensis.
The pygidium assigned to P. (C.) chongqingensis is also
unique in displaying a malformed pleural rib. Indeed, the
second rib of the left pleura is short (tr.) and does not reach
the lateral border (Fig. 6k, m). This is compensated by wider
(exs.) abaxial parts of ribs 1 and 3 and has apparently no
consequence on the general shape of the pleura. As the border
is normal in this area, the malformation might correspond
to either the result of a developmental dysfunction or to a
completely, though imperfectly healed, marginal injury.
Pseudophillipsia (Carniphillipsia) sagittalis (Kobayashi &
Hamada, 1978)
Figure 4k, n
1978 Iranaspidion sagittalis Kobayashi & Hamada,
pp. 157–60, figs 1–3.
1981 Iranaspidion sagittalis Kobayashi & Hamada; Hahn
& Hahn, p. 223.
1981 Iranaspidion sagittalis Kobayashi & Hamada;
Kobayashi & Hamada, p. 56, pl. 1, figs 1–3.
1983 Iranaspidion sagittalis Kobayashi & Hamada;
Owens, p. 29.
1984 Iranaspidion sagittalis Kobayashi & Hamada;
Kobayashi & Hamada, pp. 68–9, text-fig. 5k, pl. 14,
figs 1–3.
2003 Iranaspidion sagittalis Kobayashi & Hamada;
Owens, p. 382.
2009 Pseudophillipsia
(Carniphillipsia)
sagittalis
(Kobayashi & Hamada); Lerosey-Aubril & Angiolini,
p. 438.
cephalon in (b) dorsal and (c) right lateral views, reproduced from Arkhipova (1965, pl. 45, fig. 1) with the permission of the
Palaeontological Institute, Moscow; Wordian, Gnishik Formation, Vedi area, southern Armenia. (f) VSEGEI 51-5217, pygidium in
dorsal view; Wordian (?), unknown outcrop along the Vedi River, Vedi area, southern Armenia. (d, g, h, k, m, p) Pseudophillipsia
(Carniphillipsia) chongqingensis Lu, 1974, Lopingian, SW China. (d, g, h) NIGPAS 21975, broken holotype cephalon in (d) dorsal,
(g) right lateral and (h) frontal views. (k, m, p) NIGPAS 21976, partial thoraco-pygidium in (k) dorsal, (m) posterior and (p) left lateral
views. (i, j, l, n, o) Pseudophillipsia (Carniphillipsia) sp. A, SMF 90843, strongly weathered, complete enrolled specimen; precise
horizon unknown, Wordian to Wuchiapingian, Chahriseh area, Central Iran. (i) cephalon, (j) frontal view, (l) thorax, (n) pygidium and
(o) left lateral view. Scale bars = 5 mm.
Late Palaeozoic trilobites of Iran and Armenia
Material, locality and horizon. Complete enrolled holotype
specimen (PA 16764, specimen missing; Kobayashi &
Hamada, 1978, fig. 3a–c; Fig. 4k, n herein), complete
enrolled specimen (PA 16762, specimen missing; Kobayashi
& Hamada, 1978, fig. 1a, b), cephalon (PA 16763, specimen
missing; Kobayashi & Hamada, 1978, fig. 2) and possibly
four other specimens (missing) mentioned but not described
or figured by Kobayashi & Hamada (1978); uppermost part
of Unit 1 of Taraz (1971), Surmaq Formation, Capitanian
(Midian; Kobayashi & Ishii, 2003), southwestern part of the
Kuh-e-hambast Range, Abadeh area (Fars Province), Central
Iran (label 1 in Fig. 1).
Diagnosis (emended). A species of Pseudophillipsia (Carniphillipsia) displaying the following combination of characters: cranidium particularly vaulted; glabella short (sag.)
and almost parallel-sided all along; frontal lobe short (sag.),
inflated, covered with coarse tubercles along postero-lateral
margin, and bearing faint sagittal sulcus postero-medially;
median occipital lobe trapezoid; pygidium with 11 pleural
ribs.
Remarks. Following Owens (1983), Lerosey-Aubril &
Angiolini (2009) have considered the genus Iranaspidion
to be a junior synonym of Pseudophillipsia. This view
was supported by the demonstration that all the diagnostic
characters of Iranaspidion could in fact be observed in
different representatives of Pseudophillipsia and therefore,
that their combination in a taxon could justify its distinction
at the species level, but hardly at a generic one. Accordingly,
the type (and only) species of Iranaspidion, I. sagittalis, was
maintained but reassigned to the subgenus Pseudophillipsia
(Carniphillipsia). As mentioned above, I consider that the
isolated pygidium described by Kobayashi & Hamada (1978;
Fig. 4b, d, e herein) belongs to P. (s.l.) armenica instead
of P. (C.) sagittalis. Considering this and the remarks of
Lerosey-Aubril & Angiolini (2009) on the characters used
in the original diagnosis of Iranaspidion, the diagnosis
of P. (C.) sagittalis needed significant emendations. Most
characters proposed in this emended diagnosis focus on
the main glabellar lobe. The fact that the glabella is
almost parallel-sided along its entire length (sag.) is also
particularly notable. In fact, the features highlighted in
this diagnosis seem so original to me that I believe they
could be used for the definition of a new concept of the
genus Iranaspidion. However, because the holotype, the two
paratypes and the four non-figured specimens mentioned
by Kobayashi & Hamada (1978) are presently missing, this
would be inadvisable. The isolated cephalon figured by the
same authors (their fig. 2) does not match perfectly with
the diagnosis proposed above (e.g. parallel-sided glabella),
but this is likely owing to differences in preservation (or
preparation) between the holotype and this specimen. Lastly,
as discussed above, the similarities in pygidial morphology
between P. (s.l.) armenica and P. (C.) sagittalis, if confirmed
by direct observations of the specimens from Abadeh,
might ultimately lead to these two species being placed in
synonymy.
Pseudophillipsia (Carniphillipsia) sp. A
Figure 6i, j, l, n, o
2000 Pseudophillipsia (?Carniphillipsia) sp.; Feist et al.
in Mistiaen et al. p. 99, pl. 8, figs 2, 3.
2009 Pseudophillipsia (Carniphillipsia) sp.; LeroseyAubril & Angiolini, p. 433, table 1.
Strongly weathered, complete enrolled specimen (SMF 90843; Fig. 6i, j, l, n,
Material, locality and horizon.
1039
o). Precise horizon unknown; Wordian to Wuchiapingian
(middle–late Murghabian to early Djulfian, see Mistiaen
et al. 2000), Chahriseh area (32◦ 54 N, 52◦ 3 W; Esfahan
Province), Central Iran (label 2 in Fig. 1).
Cephalon apparently of parabolic outline
(Fig. 6a). Glabella long (maximum width/maximum length
ratio: 0.6); dorsal furrows broad and rather shallow, subparallel along posterior half of glabella, then gently and
moderately diverging forwards (c. 25◦ ), associated with
deep fossulae opposite the mid-distance between β and
γ abaxially; occipital ring with straight posterior margin,
strongly widening (exs. and sag.) medially owing to forward
curvature of the median three-fifths of SO; S1 broad and deep
near axial furrow, splitting into shallow anterior and posterior
branches rearwards; L1 ovoid, long (exs.), rather narrow (tr.)
and inflated; median occipital lobe rectangular, wide (tr.) but
rather short (exs. and sag.); main glabellar lobe somewhat
eroded postero-sagittally, and bearing three short and shallow
anterior glabellar furrows (S2–S4). Anterior border mostly
broken, but where present rather narrow (exs.) and separated
from glabella by a narrow preglabellar furrow. Posterior
fixigenal field narrow (tr.); palpebral area rather wide (tr.)
but with a narrow (tr.) and rather long (exs.) palpebral lobe;
anterior fixigenal field particularly narrow (tr.) posteriorly,
progressively doubling in width (tr.) anteriorly. α located
opposite the posterior section of the dorsal furrow anteriorly;
α–β rather long and divergent (c. 40◦ ); β rather sharp and
exceeding δ abaxially; β–γ converging (c. 35◦ ) then subparallel near fossulae; γ–δ short and divergent (c. 35◦ ); δ–
ε short and convergent (c. 30◦ ); ε–ζ rather long and subparallel; ζ–ω rather long, broadly curved anteriorly, thus
poorly defining η inflexion point; ω exceeding β abaxially.
Eye kidney-shaped, rather broad (tr.), extending from the
mid-length (exs.) of L1 to mid-length (exs.) of glabella.
Librigenal field apparently narrow (tr.), subdivided into an
inner sub-horizontal platform, which is narrow (tr.) anterolaterally but strongly widens posterior to eye, and an outer
part, which strongly slopes downwards abaxially and only
slightly widens (tr.) posteriorly. Librigenal border broken.
Posterior border furrow only slightly curving backwards
abaxially. Posterior border rather wide (exs.) adaxially and
strongly widening (exs.) abaxially. In lateral view (Fig. 6o),
posterior portion of glabella abraded, but main glabellar lobe
rather high posteriorly and increasingly sloping forwards
although without overhanging the anterior border; L1 rather
inflated. Eye rather high and apparently bounded abaxially
by a subocular groove. In frontal view (Fig. 6j), cephalon
rather strongly arched (tr.) with posterior part of the glabella
and palpebral lobe rather high; anterior border particularly
high. Doublure unknown.
Thorax composed of nine segments. Dorsal furrows
gently converging backwards and delimiting a wide (tr.)
axial lobe (c. 40 % of maximal width anteriorly). Only the
posteriormost segment can be studied entirely, although most
characters seem common to the others. Axial ring wide (sag.
and exs.) and composed of a postannulus of even width all
along (sag. and exs.) and a preannulus and an articulating
half-ring both about twice narrower (sag.) than the postannulus (Fig. 6l). Pleura moderately flexed backwards abaxial
to fulcrum, where it significantly widens (exs.) and bears a
broad (up to two-thirds of maximum width) and petaloid
articulating facet; pleural furrow apparently restricted to
vicinity of fulcrum. In lateral view (Fig. 6o), axis and pleura
of similar heights. In posterior view, axis rather high; pleura
strongly flexed downwards (c. 45◦ ) abaxial to fulcrum.
Pygidium apparently of strongly parabolic outline
(Fig. 6n), subdivided into axial and pleural lobes of equal
Description.
1040
R . L E R O S E Y- AU B R I L
Figure 7. Stratigraphical distribution of the Late Palaeozoic trilobites from Iran and Armenia. Abbreviations: A. – Acropyge; C. –
Carniphillipsia; Pe. – Persia; Ps. – Pseudophillipsia; AA – Araxian–Azerbaijanian Zone; ALB – Alborz block; AR – Arabian Plate;
CI – Central Iran; SSZ – Sanandaj–Sirjan Zone; Ar. – Armenia, Ir. – Iran; Guadalup. – Guadalupian; Loping. – Lopingian; TOU –
Tournaisian; VIS – Visean; SER – Serpukhovian; BAS – Bashkirian; MOS – Moscovian; KAS – Kasimovian; GZH – Gzhelian;
ASS – Asselian; SAK – Sakmarian; ART – Artinskian; KUN – Kungurian; ROA – Roadian; WOR – Wordian; CAP – Capitanian;
WUC – Wuchiapingian; CHA – Changhsingian.
width (tr.). Axis delimited by deep axial furrows, increasingly
tapering backwards, subdivided into a median part and two
abaxial parts about three times narrower (tr.) by two rows of
muscle scars, and possibly bearing a faint sagittal sulcus
posteriorly; 20 + 1 axial rings delimited by rather deep
axial furrows that are straight medially but backwardly
curved abaxial to muscle scars. Pleural region provided
with antero-lateral projection that receives the outer portion
of the two posteriormost thoracic pleurae; 12 pleural ribs
defined by broad and rather deep pleural furrows that
regularly but increasingly curve backwards abaxially; no
interpleural furrows discernible. Pygidial border broken.
Laterally (Fig. 6o), pygidium rather high mainly owing to
pleural field, which is twice as high as axis. Axis highest
at axial ring 3, then it strongly and increasingly flexes
downwards posteriorly. According to the partially visible
internal mould, doublure was wide and mainly orientated
in a sub-vertical plan. In posterior view (Fig. 6j), axis low
and broadly elliptical in section. Fulcrum rather abaxially
positioned, with outer part of pleural field strongly flexed
downwards (45◦ ).
Sculpture: few thin terrace ridges apparently occurred
on pygidial doublure as indicated by its internal mould.
Otherwise, no sculptural features can be observed owing to
the weathering undergone by the specimen.
Discussion. As mentioned above, P. (C.) sp. A belongs to
a group of morphologically close taxa, which also includes
P. (C.) paffenholzi and P. (C.) chongqingensis. In addition
to the characters common to the three species, it resembles
P. (C.) paffenholzi (Fig. 6a–c, e, f) in exhibiting a narrow
(sag.) anterior border furrow, a posterior cephalic border that
strongly widens abaxially, 12 flat pleural ribs and a pygidial
pleural field about as twice as high as the axis. However, the
parallel-sided portion of its glabella is longer compared with
that of the Armenian species, its median preoccipital lobe is
rectangular instead of trapezoid and it has fewer axial rings
(20 + 1). Like P. (C.) chongqingensis, it also differs from
P. (C.) paffenholzi in having a straight posterior margin of
the occipital ring, a medially strongly forwardly curved SO,
three short anterior glabellar furrows (S2–S4) instead of two
and simpler courses of inter-ring and pleural furrows. On the
other hand, P. (C.) chongqingensis (Fig. 6d, g, h, k, m, p)
displays a trapezoidal (not rectangular) median preoccipital
lobe, broad cephalic border furrow, a greater number of axial
rings (25 + 1) and pleural ribs (14), and a pygidial axis
triangular in section, all traits which cannot be observed in
P. (C.) sp. A.
The specimen found in the Middle–Late Permian of the
Chahriseh area might represent a new species with close
phylogenetic relationships with P. (C.) chongqingensis and P.
(C.) paffenholzi. Awaiting the discovery of better preserved
specimens, however, it seems more appropriate to leave this
taxon in open nomenclature.
4. Affinities of Permian trilobites from Iran:
palaeogeographical implications
Although rare, the Permian trilobites of Iran and Armenia are of similar ages (Wordian to Wuchiapingian;
Fig. 7) and they have been found in localities
representative of three different tectonic units (Alborz,
the SSZ, Transcaucasia), the positions of which relative
to one another remains poorly known for that time.
The trilobite taxa found in the Permian of the two
countries can be separated into four groups with regard
to their taxonomic affinities (Table 2). The first one
comprises the representatives of the genus Acropyge,
which strongly suggest the Alborz, Central Iran (SSZ)
and Transcaucasia microplates represented a single
biochore in late Guadalupian/early Lopingian times.
Indeed, Acropyge is a rare but very distinctive taxon
and the presence of A. encrinuroides in Abadeh and
in Ogbin on the one hand, and of A. weggeni in Yush
and a similar form (Acropyge sp. A) in Vedi on the
Late Palaeozoic trilobites of Iran and Armenia
1041
Table 2 Permian trilobite faunules associated with some Iranian tectonic units and Transcaucasia and their affinities
Arabian Plate
Alborz
SSZ
Transcaucasia
(Dena – Ir.)
(Yush – Ir.)
(Abadeh, Chahriseh, Ir.)
(Ogbin, Vedi – Ar.; Tananam – Az.)
South China
North Caucasus
–
P. (s.l.) caucasica
–
–
A. sp. A
A. encrinuroides
P. (s.l.) armenica
P. (s.l.) caucasica
P. (C.) paffenholzi
–
(A. brevica,
A. multisegmenta)
–
–
P. (s.l.) parvizi
–
A. encrinuroides
P. (s.l.) armenica
P. (C.) sagittalis
P. (C.) sp. A
–
–
–
A. weggeni
–
P. (s.l.) aff. caucasica
P. (C.) chongqingensis
–
–
–
Representatives of the genus Acropyge suggest special palaeobiogeographical affinities between the Alborz and the SSZ (Iran),
Transcaucasia and South China. Faunal relationships between the SSZ, Transcaucasia and South China are also supported by the presence of
similar species of Pseudophillipsia (Carniphillipsia) (‘paffenholzi-group’). That the Alborz, the SSZ and Transcaucasia might have been
associated with a single biochore is also suggested by the occurrence in the Permian faunules of these areas of another suite of particularly
similar species of Pseudophillipsia (‘armenica-group’). Pseudophillipsia parvizi is only known from the Arabian Plate (Dena Mountain) and
shows no clear affinities with any representatives of the genus Pseudophillipsia. Abbreviations: A. – Acropyge, C. – Carniphillipsia, P. –
Pseudophillipsia; Ar. – Armenia, Az. – Azerbaijan, Ir. – Iran.
other, should be regarded as indicative of close biogeographical affinities between the three tectonic units.
Interestingly, the remaining two species belonging to
this genus, A. brevica and A. multisegmenta, are known
from the late Lopingian of South China, a continent that
is generally located much further to the east on Permian
palaeogeographical reconstructions (e.g. Stampfli &
Borel, 2002; Torsvik & Cocks, 2004; Muttoni et al.
2009a). Biotic affinities in the Middle and Late Permian
between South China, Transcaucasia and Central Iran
(SSZ) are also suggested by the occurrence of the
three similar forms composing the paffenholzi-group
(i.e. P. (C.) chongqingensis, P. (C.) paffenholzi and P.
(C.) sp. A). With their flat-topped pleural ribs in the
pygidium, these taxa are easily differentiated from the
representatives of the armenica-group, which exhibit
prominent pleural ribs along with prominent axial
rings, a strongly downwardly flexed axis posteriorly and
a rather strong curvature of the pleural field in posterior
view. Taxa belonging to the armenica-group have
been found in Alborz (P. (s.l.) aff. caucasica), Central
Iran (SSZ; P. (s.l.) armenica and P. (C.) sagittalis)
and Transcaucasia (P. (s.l.) caucasica), confirming
that close palaeobiogeographical relationships existed
between these three terranes in the Permian period.
Outside Iran and Armenia, P. (s.l.) caucasica is also
known from the North Caucasus (Weber, 1944). Lastly,
P. parvizi sp. nov., the only Permian trilobite occurring
on the Iranian part of the Arabian Plate (Dena Mountains), is unique within the genus Pseudophillipsia in
exhibiting a pygidium with a strongly parabolic outline
and a border that significantly, and abruptly, widens
rearwards. With its flat-topped pleural ribs, however, it
might be regarded rather close morphologically to the
paffenholzi-group, but testing this assumption would
require the cephalic morphology of this new species to
be known. Until then, no particular affinities between
the Zagros area and other localities in Iran or on the
Arabian Plate can be deduced from Permian trilobites.
In summary, the comparison of the Permian trilobite
faunules from Iran and Armenia with contemporaneous
faunas from other regions leads to two main conclusions. Firstly, there were apparently no notable biotic
barriers between the Alborz, Central Iran (SSZ) and
Transcaucasia microplates in Middle and Late Permian
times. In contrast, some typical trilobite taxa are almost
restricted in their palaeogeographical distribution to
these three terranes, suggesting that they represented a
distinct biochore at that time. No obvious relationships
between these faunules and contemporaneous trilobite
faunas from southern Laurussia (e.g. Slovenia: Hahn,
Hahn & Ramovš, 1990; Hahn et al. 2002) or northern
Gondwana (Oman: Goldring, 1957; Pakistan: Grant,
1966) could be demonstrated. Secondly, affinities of
several of these Iranian and Armenian taxa point
towards faunal exchanges with South China. It is worth
noting that the palaeobiogeographical signal extracted
from these Permian trilobites from Iran and Armenia
is in accordance with other palaeontological and
sedimentological data. It has been shown for example
that the Alborz and the Central Iran microplates share
a similar stratigraphic evolution during Carboniferous
and Permian times (Leven, 1998; Leven & Gorgij,
2006; Gaetani et al. 2009). Sedimentological and
stratigraphical data also support the view that the
Alborz and Central Iran microplates were close to
Transcaucasia during the Permian period (Leven, 1998;
Gaetani et al. 2009). Likewise, the co-occurrence
of many algal (Théry, Vachard & Dransart, 2007),
brachiopod (Shen & Shi, 2000), bryozoan (Sakagami,
1980; Ernst, Senowbari-Daryan & Hamedani, 2006),
coral (Ezaki, 1991) and foraminiferal (Leven, 1998;
Théry, Vachard & Dransart, 2007) taxa in Permian
sections of these three terranes suggests they might
have constituted a distinct palaeobiogeographical unit.
The faunal affinities between the Iranian blocks and
Transcaucasia on the one hand and South China on the
other (e.g. in corals, see Wu & Zhao, 1983 and Ezaki,
1991), herein illustrated by Permian trilobites, have
been recently discussed by Gaillot & Vachard (2007).
These latter authors argued that these affinities would
be best explained if South China might have had a
more western location than the one frequently proposed
in palaeogeographical reconstructions (e.g. Stampfli &
Borel, 2002; Torsvik & Cocks, 2004; Muttoni et al.
2009a). An alternative view was defended by Shen &
Shi (2000) in their study of brachiopod palaeobiogeography in the Wuchiapingian (early Lopingian) period.
1042
This revealed affinities between brachiopod faunas
from the South China, Southeast China and Central
Thailand terranes and that of Alborz and Transcaucasia
(‘Southern Armenia’). Similar Cathaysian affinities
for Transcaucasia are also shown by Capitanian (late
Guadalupian) brachiopods (Shen & Shi, 2004), while
in Changhsingian (late Lopingian) time, brachiopod
faunal affinities suggest the inclusion of Iran and
southern Armenia into a ‘Western Tethyan Province’
that also comprises Laurussian localities, such as
Hungary and Slovenia (Shen, Archibald & Shi, 2000).
According to Shen & Shi (2000), these Cathaysian
affinities of Iranian/Armenian brachiopod faunas during late Guadalupian and early Lopingian times may
be explained by the distribution of the more eastern
Cimmerian blocks, which constituted bridges between
the Alborz/Central Iran/Transcaucasia assemblage and
Cathaysian terranes. In addition to the particular
configuration of these continental blocks, they also
emphasized the fact that all of them were located at
similar palaeolatitudes, which might have facilitated
faunal exchanges. The integration of Iran and southern
Armenia into a ‘Western Tethyan Province’ in late
Lopingian time, not shared with eastern Cimmerian
blocks, would have resulted from the proximity of
these tectonic units to the southern Laurussian margin.
On the other hand, Shen & Shi (2000) illustrated the
palaeobiogeography of Wuchiapingian brachiopods by
using a map (modified from Ziegler, Hulver & Roeley,
1997), which shows South China and other Cathaysian
blocks located rather close to Iranian microplates and
Transcaucasia, compared with other palaeogeographical models of that period (e.g. Stampfli & Borel,
2002; Torsvik & Cocks, 2004). Also it might be argued
that the palaeobiogeographical affinities between these
different tectonic units might have resulted from direct
faunal exchanges, instead of requiring that eastern
Cimmerian blocks had played the role of migration
bridges. The fact that these eastern Cimmerian blocks
and Cathaysian terranes also exhibit similarities in
faunal contents might have simply been the result of the
somewhat central position of the Cathaysian terranes,
though more in the north and close to almost all the
Cimmerian blocks as illustrated by Shen & Shi’s (2000)
figure 6. In conclusion, the data presented by these
authors do not appear to me to be in conflict with the
proposition of Gaillot & Vachard (2007) that South
China (and possibly other Cathaysian terranes) might
have occupied a rather eastern location in Lopingian
time. To some extent, the particular affinities of
Guadalupian to Lopingian trilobites from Iran/Armenia
and South China provide support for this assumption.
5. Conclusion
The Late Palaeozoic trilobites of Iran and southern
Armenia illustrate that though rare in the Late
Palaeozoic, these fossil organisms might still contribute
to the recognition of biochores and to the elaboration
of proper and robust definitions for them. Of course,
R . L E R O S E Y- AU B R I L
with such low diversity and abundance, it would
be hazardous to consider the palaeobiogeographical
signal extracted from post-Devonian trilobites alone
for constraining palaeogeographical models. However,
this should not mask the fact that trilobites were once
(i.e. in Early–Middle Palaeozoic times) critical to test
palaeogeographical reconstructions and that the ability
of these fossils to accurately reflect movements of
continents relative to one another certainly stemmed
from some particular traits of their ecology and their
biology. Indeed, trilobites were chiefly benthic and
depth-sensitive inhabitants (Fortey & Cocks, 2003),
the dispersal capacities of which were rather limited
for most of them despite the frequent occurrence of
planktonic stages in their early ontogeny. Moreover,
it is also likely that other as yet unknown biological
or ecological characteristics of trilobites might have
conferred them a particular sensitivity to geographical and climatic barriers. This has been recently
demonstrated by the investigations of Hendricks &
Lieberman (2007) on the biogeography of Cambrian
arachnomorphs. These authors depicted very different palaeobiogeographical patterns for trilobites and
non-trilobite arachnomorphs in the Cambrian period.
While tectonic events have likely had a strong influence
on the palaeogeographical pattern of Cambrian trilobites, non-trilobite arachnomorphs exhibit much more
important dispersal abilities, and their geographical
distribution and its evolution through time are better
explained by environmental changes at a supracontinental scale (their ‘cyclic events’), like sea-level
changes. Although the diversity and abundance of
trilobites are much less in the Permian period than
in the Cambrian period, it seems Permian trilobites
still had these intrinsic capacities for diversification
when geographically or climatically isolated. This is
illustrated by the original trilobite fauna of Japan during
Capitanian time, which mirrors the so-called SinoMongolian Province defined from other taxonomic
groups (Owens & Hahn, 1993; Lerosey-Aubril, 2008)
and probably generated by climatic barriers. Another
example is the appearance of a clear separation between
trilobite faunas from Western America and the periTethyan regions (i.e. the North American and Tethyan
parts of the Palaeo-equatorial Realm of Grunt &
Shi, 1997) during Guadalupian time, which reflects
the closure of the Uralian seaway. Thus, even if we
cannot reasonably rely on trilobites to the same extent
in the Permian period as earlier in the Palaeozoic
for constraining palaeogeographical models, it seems
they remained sufficiently sensitive to geographical
or climatic barriers to permit light to be shed on
some specific palaeogeographical issues, such as the
Late Palaeozoic evolution of tectonic units of Iran and
neighbouring areas.
Acknowledgements. I am particularly grateful to M.
Ghobadi Pour (National Museum of Wales), M. Parvizi
(University of Payam-e Noor) and M. Yazdi (Esfahan
University) who made available for study the specimens
of, respectively, Pseudophillipsia (Carniphillipsia) sp. A,
Late Palaeozoic trilobites of Iran and Armenia
Pseudophillipsia (s.l.) parvizii sp. nov. and Persia praecox
gen. nov. sp nov. I am also indebted to D. Vachard (University
Lille 1) who studied the thin-sections made from the trilobitebearing rock samples from the Yush area and the Dena
Range. E. Naimark (Palaeontological Institute, Moscow)
provided me with photographs of the original material of
Weber (1944) housed at the ‘A. P. Karpinsky’ Russian
Geological Research Institute (Saint Petersburg) and helped
me with the Russian part of Weber’s work. X.-J. Zhu (Nanjing
Institute of Geology and Palaeontology) did the same with
the type specimens of Pseudophillipsia (Carniphillipsia)
chongqingensis hosted by Zhu’s institution. Both of them
deserve my sincere gratitude for their help. I am also grateful
to K. J. McNamara (University of Cambridge), who patiently
corrected an earlier draft of the manuscript and to Raimund
Feist (University of Montpellier II) and an anonymous referee
for their helpful comments. Finally, the Japan Academy
and the Palaeontological Institute (Moscow) are thanked for
permitting the reproduction of figures. This research received
no specific grant from any funding agency, commercial or
not-for-profit sectors.
References
ANGIOLINI, L., GAETANI, M., MUTTONI, G., STEPHENSON,
M. H. & ZANCHI, A. 2007. Tethyan oceanic currents
and climate gradients 300 m.y. ago. Geology 35, 1071–
4.
ARKHIPOVA, O. I. 1965. Trilobites. In Razvitie i Smena Morskikh Organizmov na Rubezhe Paleozoya i Mezozoya
(eds V. E. Ruzhentsev & T. G. Sarycheva), pp. 82–3.
Moscow: Nauka (in Russian).
BREZINSKI, D. K. 1992. Permian trilobites from West Texas.
Journal of Paleontology 66, 924–43.
EBBESTAD, J. O. R. & BUDD, G. E. 2003. Burlingiid trilobites
from Norway, with a discussion of their affinities and
relationships. Palaeontology 45, 1171–95.
ERNST, A., SENOWBARI-DARYAN, B. & HAMEDANI, A. 2006.
Middle Permian Bryozoa from the Lakaftari area,
northeast of Esfahan (central Iran).Geodiversitas 28,
543–90.
EZAKI, Y. 1991. Permian corals from Abadeh and Julfa, Iran,
West Tethys. Journal of the Faculty of Science, Hokkaido
University. Series 4, Geology and Mineralogy 23, 53–
146.
EZAKI, Y. 1993. Sequential disappearance of Permian Rugosa
in Iran and Transcaucasus, West Tethys. Bulletin of the
Geological Survey of Japan 44, 447–53.
FEIST, R., LEROSEY-AUBRIL, R. & JOHNSON, R. 2010.
Coaptative devices, enrollment, and life habits in
Paralejurus, a particular case in scutelluid trilobites.
Palaeobiodiversity & Palaeoenvironments 90, 125–37.
FORTEY, R. A. & COCKS, L. R. M. 2003. Palaeontological
evidence bearing on global Ordovician–Silurian continental reconstructions. Earth-Science Reviews 61, 245–
307.
FORTEY, R. A. & OWENS, R. M. 1975. Proetida – a new order
of trilobites. Fossils and Strata 4, 227–39.
GAETANI, M., ANGIOLINI, L., UENO, K., NICORA, A.,
STEPHENSON, M. H., SCIUNNACH, D., RETTORI, R.,
PRICE, G. D. & SABOURI, J. 2009. Pennsylvanian Early
Triassic stratigraphy in the Alborz Mountains (Iran). In
South Caspian to Central Iran Basins (eds M.-F. Brunet,
M. Wilmsen & J. W. Granath), pp. 79–128. Geological
Society of London, Special Publication no. 312.
GAILLOT, J. & VACHARD, D. 2007. The Khuff Formation
(Middle East) and time-equivalents in Turkey and
1043
South China: Biostratigraphy from the Capitanian to
Changhsingian times (Permian), new foraminiferal taxa,
and palaeogeographical implications. Coloquios de
Paleontología 57, 37–223.
GAURI, K. L. 1965. Uralian stratigraphy. Trilobites and
brachiopods of the Western Carnic Alps (Austria).
Jahrbuch der Geologischen Bundesanstalt 11, 1–26.
GEMMELLARO, G. G. 1892. I crostacei dei calcari con
Fusulina della valle del Fiume Sosio nella Provincia
di Palermo in Sicilia. Memorie della Società ltaliana
delle Scienze, Serie 3 8, 1–40.
GOLDRING, R. 1957. Pseudophillipsia (Tril.) from the
Permian (or Uralian) of Oman, Arabia. Senckenbergiana
Lethaea 38, 195–210.
GRANT, R. E. 1966. Late Permian trilobites from the Salt
Range, West Pakistan. Palaeontology 9, 64–73.
GRUNT, T. A. & SHI, G. R. 1997. A hierarchical biogeographical classification of the Permian global marine
biogeography. In Proceedings of the 30th International
Geological Congress (eds Y. G. Jin & D. Dineley).
Palaeontology and Historical Geology 12, 2–17.
HAAS, W., HAHN, G. & HAHN, R. 1980. Perm-Trilobiten
aus Afghanistan. Palaeontographica Abteilung A 169,
73–127.
HAHN, G. & BRAUCKMANN, C. 1975. Revision zweier
Trilobiten-Arten aus dem Perm Asiens. Geologica et
Palaeontologica 9, 117–24.
HAHN, G. & HAHN, R. 1967. Zur phylogenie der Proetidae (Trilobita) des Karbons und Perms. Zoologische
Beiträge, Neue Folge 13, 303–49.
HAHN, G. & HAHN, R. 1981. Über Acropyge (Trilobitae;
Ober-Perm). Senckenbergiana Lethaea 61, 217–25.
HAHN, G. & HAHN, R. 1987. Trilobiten aus dem Karbon
von Notsch und aus den Karnischen Alpen Osterreichs.
Jahrbuch der Geologischen Bundesanstalt 129, 567–
619.
HAHN, G., HAHN, R., MÜLLER, P. & RAMOVŠ, A. 2002. Neue
Trilobiten-Funde aus dem Unter-Perm Sloweniens.
Geologica et Palaeontologica 36, 99–113.
HAHN, G., HAHN, R. & RAMOVŠ, A. 1990. Trilobiten
aus dem Unter-Perm (Trogkofel-Kalk, Sakmarium) der
Karawanken in Slowenien. Geologica et Palaeontologica 24, 139–71.
HAHN, G., HAHN, R. & YUAN, J.-L. 1989. Trilobites from
the Upper Carboniferous (Westphalian A) of S-China
(N-Guangxi). Geologica et Palaeontologica 23, 113–
203.
HENDRICKS, J. R. & LIEBERMAN, B. S. 2007. Biogeography and the Cambrian radiation of arachnomorph
arthropods. Memoirs of the Association of Australasian
Palaeontologists 34, 461–71.
HUPÉ, P. 1953. Classe des trilobites. In Traité
de paléontologie, Tome III, les Formes Ultimes
d’Invertébrés, Morphologie et Evolution, Onychophores, Arthropodes, Echinodermes, Stomocordés (ed.
J. Piveteau), pp. 44–246. Paris: Masson.
KOBAYASHI, T. & HAMADA, T. 1978. Two new Upper Permian
trilobites from Central Iran. Proceedings of the Japan
Academy, Series B 54, 157–62.
KOBAYASHI, T. & HAMADA, T. 1981. A new trilobite from
the Upper Permian of Central Iran. Reports of the
Geological Survey of Iran 49, 55–8.
KOBAYASHI, T. & HAMADA, T. 1984. Permian trilobites
of Japan in comparison with Asian, Pacific and other
faunas. Palaeontological Society of Japan, Special
Papers 26, 1–92.
KOBAYASHI, F. & ISHII, K.-I. 2003. Palaeobiogeographic
analysis of Yahtashian to Midian fusulinacean faunas
1044
of the Surmaq Formation in the Abadeh region, Central
Iran. Journal of Foraminiferal Research 33, 155–65.
LEROSEY-AUBRIL, R. 2008. Trilobite biogeography and
Permian biochores. In Advances in Trilobite Research
(eds I. Rabano, R. Gozalo & D. Garcia-Bellido).
Cuadernos del Museo Geominero 9, 225–8.
LEROSEY-AUBRIL, R. & ANGIOLINI, L. 2009. Permian
trilobites from Antalya Province, Turkey, and enrollment
in Late Paleozoic trilobites. Turkish Journal of Earth
Sciences 18, 427–48.
LEROSEY-AUBRIL, R. & FEIST, R. 2012. Quantitative approach of diversity and decline in late Palaeozoic
trilobites. In Earth and Life: Global Biodiversity,
Extinction Intervals and Biogeographic Perturbations
Through Time (ed. J. A. Talent), pp. 535–55. Dordrecht:
Springer Science and Media.
LEROSEY-AUBRIL, R., HEGNA, T. A. & OLIVE, S. 2011. Inferring internal anatomy from the trilobite exoskeleton: the
relationship between the frontal auxiliary impressions
and the digestive system. Lethaia 44: 166–174.
LEVEN, E. J. 1998. Permian fusulinid assemblages and
stratigraphy of the Transcaucasia. Rivista Italiana di
Paleontologia e Stratigraphia 104, 299–328.
LEVEN, E. J. & GORGIJ, M. N. 2006. Upper Carboniferous–
Permian stratigraphy and fusulinids from the Anarak
region, Central Iran. Russian Journal of Earth Sciences
8(2), ES2002, doi: 10.2205/2006ES000200.
LIU, R.-Y. 1987. A new trilobite genus from Lower
Carboniferous of Xinshao, Hunan and its enrollment.
Acta Palaeontologica Sinica 26, 493–6.
LU, Y.-H. 1974. Permian trilobites. In Hand-book of Strata
and Fossils of Southwest China, p. 299. Nanjing Institute
of Geology and Palaeontology, Academia Sinica 236.
LUDVIGSEN, R. & CHATTERTON, B. D. E. 1982. Ordovician
Pterygometopidae (Trilobita) of North America. Canadian Journal of Earth Sciences 19, 2179–206.
MISTIAEN, B., GHOLAMALIAN, H., GOURVENNEC, R.,
PLUSQUELLEC, Y., BIGEY, F., BRICE, D., FEIST, M.,
FEIST, R., GHOBADI POUR, M., KEBRIA-EE, M., MILHAU,
B., NICOLLIN, J.-P., ROHART, J.-C., VACHARD, D. &
YAZDI, M. 2000. Preliminary data on the Upper
Devonian (Frasnian, Famennian) and Permian fauna
and flora from the Chahriseh Area (Esfahan Province,
Central Iran). Annales de la Société Géologique du Nord
(2ème série) 8, 93–102.
MOHTAT-AGHAI, P. & VACHARD, D. 2005. Late Permian
foraminiferal assemblages from the Hambast region
(Central Iran) and their extinctions. Revista Española
de Micropaleontologia 37, 205–27.
MORZADEC, P. 1981. Les trilobites. In La Tranchée de
La Lezais, Emsien Supérieur du Massif Armoricain
– Sédimentologie, Paléontologie, Stratigraphie (eds P.
Morzadec, F. Paris & P. Rachebœuf), pp. 279–87.
Mémoires de la Société Géologique et Minéralogique
de Bretagne 24.
MUTTONI, G., GAETANI, M., KENT, D. V., SCIUNNACH, D.,
ANGIOLINI, L., BERRA, F., GARZANTI, E., MATTEI, M.
& ZANCHI, A. 2009a. Opening of the Neo-Tethys Ocean
and the Pangea B to Pangea A transformation during the
Permian. GeoArabia 14, 17–48.
MUTTONI, G., MATTEI, M., BALINI, M., ZANCHI, A.,
GAETANI, M. & BERRA, F. 2009b. The drift history
of Iran from the Ordovician to the Triassic. In South
Caspian to Central Iran Basins (eds M.-F. Brunet, M.
Wilmsen & J. W. Granath), pp. 7–29. Geological Society
of London, Special Publication no. 312.
OEHLERT, M. D. 1886. Étude sur quelques trilobites du
groupe de Proetidae. Bulletin de la Société d’Étude des
Sciences d’Angers, Nouvelle Série 15, 121–43.
R . L E R O S E Y- AU B R I L
OWENS, R. M. 1983. A review of Permian trilobite genera.
Palaeontology 30, 15–41.
OWENS, R. M. 2003. The stratigraphical distribution and
extinctions of Permian trilobites. Special Papers in
Palaeontology 70, 377–97.
OWENS, R. M. & HAHN, G. 1993. Biogeography
of Carboniferous and Permian trilobites. Geologica
et Palaeontologica 27, 165–80.
PARTOAZAR, H. 2002. Permian-Triassic boundary conodonts
from the Jolfa-Abadeh Belt along the Northwest and
Central Iran. Permophiles 41, 32–40.
QIAN, Y. 1977. Upper Permian trilobites from Qinglong and
Anshun of Guizhou. Acta Palaeontologica Sinica 16,
279–86 (in Chinese with English summary).
RIGBY, J. K., SENOWBARI-DARYAN, B. & HAMEDANI, A.
2005. First occurrence of wewokellid sponges (Calcarea,
Heteractinida) from the Permian of central Iran. Facies
51, 516–21.
ROEMER, F. 1880. Über eine Kohlenkalk fauna der Westküste
von Sumatra. Palaeontographica Abteilung A 27, 1–11.
RUBAN, D. A. 2007a. Major Paleozoic-Mesozoic unconformities in the Greater Caucasus and their tectonic reinterpretation: a synthesis. GeoActa 6, 91–102.
RUBAN, D. A. 2007b. Paleozoic palaeogeographic frameworks of the Greater Caucasus, a large Gondwanaderived terrane: consequences from the new tectonic
model. Natura Nascosta 34, 16–27.
RUBAN, D. A., AL-HUSSEINI, M. I. & IWASAKI, Y. 2007.
Review of Middle East Paleozoic plate tectonics.
GeoArabia 12, 35–56.
SAKAGAMI, S. 1980. Permian Ectoprocta (Bryozoa) from
the Abadeh region, Central Iran. Transactions and
Proceedings of the Palaeontological Society of Japan,
New Series 118, 269–89.
SENOWBARI-DARYAN, B. & HAMEDANI, A. 2002. First report
of the occurrence of Amblysiphonella (thalamid sponge)
in Permian of Iran and description of A. iranica n. sp.
from central Iran. Revue de Paléobiologie 21, 795–801.
SHEN, S. Z., ARCHBOLD, N. W. & SHI, G. R. 2000.
Changhsingian (Late Permian) brachiopod palaeobiogeography. Historical Biology 15, 121–34.
SHEN, S. Z. & SHI, G. R. 2000. Wuchiapingian (early
Lopingian, Permian) global brachiopod palaeobiogeography: a quantitative approach. Palaeogeography,
Palaeoclimatology, Palaeoecology 162, 299–318.
SHEN, S. Z. & SHI, G. R. 2004. Capitanian (Late Guadalupian,
Permian) global brachiopod palaeobiogeography and
latitudinal diversity pattern. Palaeogeography, Palaeoclimatology, Palaeoecology 208, 235–62.
STAMPFLI, G. M. & BOREL, G. D. 2002. A plate tectonic
model for the Paleozoic and Mesozoic constrained by
dynamic plate boundaries and restored synthetic oceanic
isochrons. Earth and Planetary Science Letters 196, 17–
33.
TARAZ, H. 1971. Uppermost Permian and Permo-Triassic
transition beds in Central Iran. Bulletin of the American
Association of Petroleum Geologists 55, 1280–94.
TARAZ, H., GOLSHANI, F., NAKAZAWA, K., SHIMIZU, D.,
BANDO, Y., ISHII, K.-I., MURATA, M., OKIMURA, Y.,
SAKAGAMI, S., NAKAMURA, K. & TOKUOKA, T. 1981.
The Permian and the Lower Triassic systems in Abadeh
region, central Iran. Memoirs of the Faculty of Science,
Kyoto University, Series Geology Mineralogy 47, 61–
133.
THÉRY, J. M., VACHARD, D. & DRANSART, E. 2007.
Late Permian limestones and the Permian-Triassic
boundary: new biostratigraphic, paleobiogeographical
and geochemical data in Caucasus and eastern Europe.
In Palaeozoic Reefs and Bioaccumulations: Climatic
Late Palaeozoic trilobites of Iran and Armenia
and Evolutionary Controls (eds J. J. Álvaro, M. Aretz,
F. Boulvain, A. Munnecke, D. Vachard & E. Vennin),
pp. 255–74. Geological Society of London, Special
Publication no. 275.
TORSVIK, T. H. & COCKS, L. R. M. 2004. Earth geography
from 400 to 250 Ma: a palaeomagnetic, faunal and facies
review. Journal of the Geological Society, London 161,
555–72.
WEBER, V. N. 1939. Trilobita. In The Atlas of the Leading
Forms of Fossil Faunas of USSR, Volume 6, Permian
(ed. B. Lisharew), pp. 196–200. Saint-Petersburg:
The Central Geological and Prospecting Institute (in
Russian).
WEBER, V. N. 1944. Trilobites of the Carboniferous and
Permian system of the USSR. Fascicule II. Permian
trilobites. Palaeontology of the USSR Monographs 71,
1–32 (in Russian, with abridged version in English).
WENDT, J., KAUFMANN, B., BELKA, Z., FARSAN, N. &
KARIMI BAVANDPUR, A. 2002. Devonian/Lower Carboniferous stratigraphy, facies patterns and palaeogeography of Iran. Part I. Southeastern Iran. Acta Geologica
Polonica 52, 129–68.
WENDT, J., KAUFMANN, B., BELKA, Z., FARSAN, N. &
KARIMI BAVANDPUR, A. 2005. Devonian/Lower Carboniferous stratigraphy, facies patterns and palaeogeography of Iran. Part II. Northern and central Iran. Acta
Geologica Polonica 55, 31–97.
WESTERMANN, G. E. G. 2000. Biochore classification and
nomenclature in paleobiogeography: an attempt at order.
Palaeogeography, Palaeoclimatology, Palaeoecology
158, 1–13.
WHITTINGTON, H. B., CHATTERTON, B. D. E., SPEYER, S.
E., FORTEY, R. A., OWENS, R. M., CHANG, W. T.,
1045
DEAN, R. A., JELL, P. A., LAURIE, J. R., PALMER, A.
R., REPINA, L. N., RUSHTON, A. W. A., SHERGOLD, J.
H., CLARKSON, E. N. K., WILMOT, N. V. & KELLY, S.
R. A. 1997. Treatise on Invertebrate Paleontology, Part
O, Arthropoda 1, Trilobita, Revised (ed. R. L. Kaesler).
Boulder: Geological Society of America and University
of Kansas Press.
WU, W. S. & ZHAO, J. M. 1983. Late Permian corals from
Zhejiang, Guangxi and Sichuan Provinces. Nanjing
Institute of Geology and Palaeontology, Academia
Sinica 6, 271–84 (in Chinese with English summary).
YAZDI, M. 1999. Late Devonian-Carboniferous conodonts
from eastern Iran. Rivista Italiana di Paleontologia e
Stratigraphia 105, 167–200.
YIN, K. 1978. Trilobita. In Atlas of Southwest Chinese Fossils
in Kueichou District. 2: Carboniferous–Quaternary (ed.
Bureau of Geology of Guizhou Province), pp. 440–
5, 628–9. Beijing: Geological Publishing House (in
Chinese).
ZANCHI, A., ZANCHETTA, S., GARZANTI, E., BALINI, M.,
BERRA, F., MATTEI, M. & MUTTONI, G. 2009. The
Cimmerian evolution of the Nakhlak-Anarak area,
Central Iran, and its bearing for the reconstruction of
the history of the Eurasian margin. In South Caspian to
Central Iran Basins (eds M.-F. Brunet, M. Wilmsen & J.
W. Granath), pp. 261–86. Geological Society of London,
Special Publication no. 312.
ZIEGLER, A. M., HULVER, M. L. & ROELEY, D. B. 1997.
Permian world topography and climate. In Late Glacial
and Postglacial Environmental Changes – Quaternary,
Carboniferous–Permian and Proterozoic (ed. I. P.
Martini), pp. 111–46. New York: Oxford University
Press.
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