Geobios 52 (2019) 1–25

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Research paper

Late Moscovian (mid-Pennsylvanian) rugose corals from Wadi Araba

(Egypt, Eastern Desert): Taxonomy, palaeoecology and
palaeobiogeography§
Mahmoud Kora a, Hans-Georg Herbig b, Heba El Desouky a,*
a
Mansoura University, Department of Geology, Faculty of Science, 35516 Mansoura, Egypt
b
Universität zu Köln, Institut für Geologie und Mineralogie, Zülpicher Straße 49a, 50674 Köln, Germany

A R T I C L E I N F O A B S T R A C T

Article history: An important late Moscovian rugose coral association is described from the Rod El Hamal Formation of
Received 17 December 2017 the Wadi Araba area (northern Eastern Desert, Egypt). The upper part of the formation yielded a
Accepted 15 November 2018 moderately abundant but poorly diversified coral fauna composed of large dissepimented solitary rugose
Available online 24 November 2018
corals. In total, six species belong to the Bothrophyllidae and the Geyerophyllidae, including three new
species: Bothrophyllum suezensis, Bothrophyllum cylindricum, and Amygdalophylloides omarai. Three
Keywords: undeterminable Bothrophyllidae taxa are left in open nomenclature. The corals were attached to the soft
Pennsylvanian
substrate by talons and rootlets, either rooted and growing upward as mud-stickers or as secondary
Rugosa
New species
recliners. They show frequent rejuvenations and constrictions. Encrustation and/or bioerosion are
Palaeoecology scarce. Carbonate microfacies indicate a habitat in littoral and inner neritic zones adjacent to a low
Palaeobiogeography terrigenous hinterland. Colonial taxa are missing and tabulate corals are rare. The corals from the Rod El
Northern Gondwana Hamal Formation are the only example of a late Moscovian coral fauna on the southern margin of the
Palaeotethys, that is to say along the northern margin of Gondwana. The association shows similarities
with northwestern Spain (Cantabrian Mountains), the Donets and Moscow basins, and thus a general
attribution to the Palaeotethys realm.
C 2018 Elsevier Masson SAS. All rights reserved.

1. Introduction
Carboniferous deposits are scarce in northern Egypt and only crop
out in the Gulf of Suez as discontinuous exposures bordering both sides
of the Gulf. East of the Gulf, they are exposed in the west-central Sinai
at Um Bogma, Wadi Feiran and Abu Durba, and west of the Gulf in the
Northern Galala, Abu Darag, Wadi Araba and at Wadi El-Dakhel in the
northern reaches of Wadi Qena (Kora, 1998;Fig. 1(A)). At the southern
cliffs of the Northern Galala plateau, northern Wadi Araba, the mixed
calcareous-siliciclastic Rod El Hamal Fm. of supposed Pennsylvanian
age was investigated for its rugose coral content.
Rugose corals from Wadi Araba were first noticed by Schweinfurth
(1885) who cited only one species (cf. Zaphrentis guerangeri Milne-
Edwards et Haime). Abdallah and El Adindani (1965) and El Adindani
(1968) identified few Pennsylvanian rugose corals as Lophophyllidium
Grabau, Caninia Michelin, Clisiophyllum Dana and Cyathaxonia Michelin,
and the tabulate corals Syringopora Goldfuss and Aulopora Goldfuss
§
Corresponding editor: Bertrand Lefebvre.
* Corresponding author.
E-mail address: hebaeldesoky@mans.edu.eg (H.E. Desouky).
from the Wadi Araba. All these identifications were based only on
external characters and are therefore mostly unreliable. Herbig and
Kuss (1988) described and illustrated Amygdalophylloides ivanovi
(Dobrolyubova, 1937), Bothrophyllum pseudoconicum Dobrolyubova,
1937, and Pseudozaphrentoides ex gr. juddi (Thomson, 1893) from
Pennsylvanian strata cropping out along the eastern side of the
Northern Galala and its southern rim bordering Wadi Araba. The
occurrence of the first two taxa is confirmed herein, but Pseudoza-
phrentoides ex gr. juddi (Thomson, 1893) is transferred to Bothrophyllum
cylindricum nov. sp. A brief summary of the coral association described
herein was already presented in El Desouky (2016).
2. Stratigraphic and geological setting
The Pennsylvanian succession of Wadi Araba is only repre-
sented by the Rod El Hamal Fm., exposed on both sides of the Wadi
Araba, mostly west of the junction with the Wadi Rod El Hamal
(Fig. 1(B)). Abdallah and El Adindani (1965) measured a ca. 350 m-
thick section of a mixed carbonate-siliciclastic succession. For the
present study, only the upper part of the formation was sampled
and studied at three sites, as the strata below are devoid of corals.

https://doi.org/10.1016/j.geobios.2018.11.004
0016-6995/
C 2018 Elsevier Masson SAS. All rights reserved.
2 M. Kora et al. / Geobios 52 (2019) 1–25
Fig. 1. Stratigraphy and geological setting. A. Carboniferous exposures in the Gulf of Suez region (after Kora, 1998). B. Geological map of the Wadi Araba area, modified from
Abdallah and El Adindani (1965).
The first site is located at Wadi Abu Kharuf, a small tributary of
Wadi Rod El Hamal (29811’01’’N, 32825’59’’E). The second and third
sites are located to the west of Wadi Abu Kharuf (29809’39.90’’N,
32824’36.50’’E, and 2989.20’41’’N, 32824’56.90’’E, respectively;
Fig. 1(B)).
The Rod El Hamal Fm. has nowhere an exposed base. In Wadi
Araba borehole n81, it overlies a fossiliferous black shale unit (‘‘B
member’’ of the oil company geologists’ classification; Said, 1971).
In the Wadi Araba area, the Rod El Hamal Fm. is unconformably
overlain by the Permo–Triassic red beds of the Qiseib Fm.
(Abdallah and El Adindani, 1965; Darwish, 1992; Issawi et al.,
2009).
The Rod El Hamal Fm. is composed of three lithostratigraphic
units (Fig. 1(B)). Unit 1 – equivalent to that of Abdallah and El
Adindani (1965) – comprises 70 m – thick, cross-bedded
sandstones alternating with multicolored shales and capped by
a thin sandy fossiliferous limestone bed as exposed at the western
side of the entrance of Wadi Rod El Hamal at its junction with
Wadi Araba (29808’51.36’’N, 32827’20.32’’E). This unit contains
the oldest sedimentary rocks exposed in Wadi Araba area. Unit 2
(ca. 130 m thick) is composed of repetitive sequences of green,
finely laminated fossiliferous marls interbedded with bluish grey,
highly fossiliferous crinoidal dolomitic limestone beds, and
capped by a thick cross-bedded sandstone bed. This unit is
well-exposed on the southern slope of Wadi Araba, facing Wadi
Rod El Hamal (29808’35.47’’N, 32827’32.01’’E). Abdallah and El
Adindani (1965) subdivided the beds of this unit into two mapped
units (namely ‘unit 2’ and ‘unit 3’ of these authors) not
differentiated herein.
Unit 3 (former ‘unit 4’ of Abdallah and El Adindani, 1965) is best
exposed to the North of Wadi Araba at Wadi Rod El Hamal, Wadi
Abu Kharuf, and Wadi Bekheit. A 47.5 m-thick succession
belonging to this unit has been investigated in the present study
at Wadi Abu Kharuf (29811’01’’N, 32825’59’’E; Fig. 2(A)). It is made
up of relatively high hills and cliffs of yellowish to reddish
fossiliferous sandstones at the base. These sandstones are followed
by a ca. 3 m-thick fossiliferous sandstone with many trace fossils
and ripple marks on the beds surfaces. Thick green marl layers
interbedded with thin beds of fossiliferous limestone cover the
previous beds in the form of weathered patches and isolated low
 M. Kora et al. / Geobios 52 (2019) 1–25 3

Fig. 2. Lithostratigraphic succession studied in the Wadi Araba area (Rod El Hamal Fm., Unit 3). A. General view of the studied section of Unit 3 in Wadi Abu Kharuf. B, C. Field
photographs showing weathered patches of relatively thick marl beds intercalated with thin fossiliferous limestone beds. D, E. Rugose corals (Bothrophyllum suezensis nov.
sp.) are weathered-out on the top of the limestone and marly beds. F. Fossiliferous limestone with well-preserved brachiopod shells. G, H. Fossiliferous limestone slabs with
gastropod and brachiopod shells, crinoid ossicles and echinoid spines.

hills (Fig. 2(B, C)). The rugose corals studied here were collected
from these marl and limestone beds. Besides the rugose corals, it
yielded many well-preserved fossils, such as brachiopod shells,
gastropods, tabulate corals, echinoid spines and plates, crinoid
ossicles, bryozoans, and trilobites (Fig. 2(D–H)). The microfacies
examination of the carbonate rocks indicates that they were
deposited in an environment ranging from restricted lagoonal to
open lagoonal environment (El-Shazly, 2011). About 20 m-thick
unfossiliferous sandstone beds overly the previous marl and
limestone beds, mostly covered with talus and topped by a thin
fossiliferous sandstone bed with trace fossils as well as brachiopod
and molluscan moulds.
The age of the Rod El Hamal Fm. is controversial and different
ages were proposed. Walther (1890) recorded and illustrated some
4 M. Kora et al. / Geobios 52 (2019) 1–25
foraminifers, bryozoans, brachiopods, molluscs, echinoderms and
a single rugose coral species, and assigned the limestone and the
marl beds to the classical West-European Early Carboniferous.
Schellwien (1894) restudied Walther’s material, revised his
identifications and concluded a Late Carboniferous age. El-
Nakkady (1958) placed the Rod El Hamal Fm. at the base of the
Spirifer mosquensis horizon C1/2 of the Russian nomenclature, i.e.,
at the base of the late Moscovian, that is to say into the
Westphalian. Abdallah and El Adindani (1965) recorded several
bryozoans, brachiopods and rugose and tabulate corals in the
different units of the Rod El Hamal Fm. and attributed it an
uppermost Pennsylvanian or younger age. Awad and Said (1966)
assigned a Namurian–Westphalian age to the formation. Omara
and Gramann (1966) and Omara and Kenawy (1966) correlated the
marine Carboniferous deposits in Wadi Araba with the American
upper Missourian–lower Virgilian (equivalents of the Kasimovian–
Gzhelian interval, or lower Stephanian in Western Europe). This
was based on only externally studied agglutinated and calcareous
foraminifers, ostracods and few conodonts (‘‘Streptognathodus
oppletus Ellison, 1941’’, and fragments of idiognathids and
hindeodellids), which were collected from two shale beds in unit
2 of the Rod El Hamal Fm. Said and Eissa (1969) studied an
assemblage of agglutinated foraminifers from unit 2 and postulat-
ed an upper Pennsylvanian age. Based on rugose corals, Herbig and
Kuss (1988) attributed a middle Westphalian C to middle
Stephanian B (Moscovian–Kasimovian) age. Recently, Ernst et al.
(2017) recorded a bryozoan assemblage from the uppermost coral-
bearing horizons of the formation that shows connections to the
Pennsylvanian (Moscovian–Kasimovian) of Russia. Refined study
of the bryozoans indicates a Moscovian age, one species being only
known from the upper Moscovian (A. Ernst, pers. comm.). During
the present study, an epimastoporellacean alga somewhat similar
to Paraepimastopora irregularis or P. jewetti was found. Both species
are known from the Desmoinesian (late Moscovian) of North
America (D. Vachard, pers. comm.). This means that according to
brachiopods (El-Nakkady, 1958), corals (Herbig and Kuss, 1988;
this work), bryozoans and algae, a late Moscovian age is emerging
for the coral-bearing upper unit of the formation, whereas the
earlier stratigraphic assignments based on externally studied
foraminifers and rare conodonts remained unspecific.
3. Material and methods
About 220 rugose coral specimens were collected from the marl
and limestone beds of unit 3 of the Rod El Hamal Fm. (Fig. 2). These
lithologies are easily eroded and remain as isolated outcrop
patches and small escarpments (Fig. 2(B, C)), so that the corals
were washed and mostly isolated from their matrix (Fig. 2(D, E)).
Hence, calices and apexes are fairly well preserved, but in some
specimens, the calices are partly eroded or still filled with matrix.
Most of the corallites were sectioned transversally and
longitudinally and studied in thin-sections using a Leica M420
microscope equipped with a Leica digital camera DFC 320. Serial
transverse sections were made from well-preserved specimens to
record the ontogenetic development. All the collected materials
are deposited at the Geology Department, Faculty of Science,
Mansoura University, Mansoura, Egypt. Nomenclature follows Hill
(1981) with some additions by Fedorowski (2016).
4. Systematic palaeontology
Subclass Rugosa Milne-Edwards et Haime, 1850
Order Stauriida Verrill, 1865
Suborder Caniniina Wang, 1950
Family Bothrophyllidae Fomichev, 1953
Genus Bothrophyllum Trautschold, 1879
Type species: Turbinolia conica Fischer von Waldheim, 1837.
Included species: Fedorowski (2016) listed and discussed
thoroughly the species included in the genus
Bothrophyllum. Recently, he added B. kalmyussi Fedorowski, 2017
and B. gorbachevensis Fedorowski, 2017 to his previous list.
Furthermore, the herein newly described B. suezensis nov. sp.
and B. cylindricum nov. sp. have to be added.
Age and distribution: Bothrophyllum is known from the Viséan
to the lower Permian, being typical of the Pennsylvanian
(Chwieduk, 2013). It is recorded from the British Isles, Cantabrian
Mts., North Africa, Donets and Moscow basins, Ural Mts., China,
Australia, and North America.
Diagnosis (Hill, 1981, emended by Fedorowski, 2016): Solitary
bothrophyllid coral with axial septum present during entire neanic
growth stage. Inner margins of several major septa approach
corallite axis up to early maturity, whereas in late maturity axial
area may be free of septa. Cardinal septum equal to average major
septa in length or longer, shorter in some species. Thickened part of
cardinal septum may be short. Counter septum is long up to early
mature growth stage, may be equal to other major septa in
advanced maturity. Septa may be thickened in tabularium,
thinning first in counter quadrants. They are thin in dissepimen-
tarium. Cardinal fossula is open. Loose indistinct axial structure is
composed of inner margins of major septa and axial tabellae; may
be gradually reduced during growth. Biform tabularium and/or
disstabularium may develop or not.
Bothrophyllum suezensis nov. sp.
Figs. 3–7
Derivation of the name: From the Gulf of Suez, type area of this
species.
Holotype: Specimen RAr 9 (Fig. 3); six transverse and one
longitudinal thin-sections available.
Type locality: Wadi Rod El Hamal (29811’01’’N, 32825’59’’E),
Wadi Araba area, northern Eastern Desert, western side of the Gulf
of Suez, Egypt.
Type horizon: Rod El Hamal Fm., unit 3, late Moscovian.
Material: In addition to the holotype (RAr 9), paratypes are
represented by 16 almost completely preserved corallites, more or
less retaining their tips (RAr 1, 4-8, 10, 19, 21, 25-27, 30, 33, 40,
153), with 82 transverse and 15 longitudinal thin-sections; an
additional 47 incomplete corallites (RAr 11-13, 15-18, 20, 32, 34,
42, 44, 82, 146, 149, 152, 190–220) and many fragments, with
48 transverse and 13 longitudinal thin-sections available. Nineteen
additional corallites were not sectioned and only examined from
their external morphology and polished surfaces (RAr 31, 39, 42,
47, 50, 54, 56, 57, 63, 66, 69, 78, 154, 151, 155-158, 160, 162), plus
12 small sized, complete corallites (RAr 28, 37, 43, 47, 48, 52, 62,
70, 169, 170, 171).
Diagnosis: Corallites are trochoid, 3.5 – 6.5 cm in height, and
maximum 43 major septa in 44.8 mm diameter. Major septa are
long up to the late mature stage. Minor septa are short and mostly
restricted to the outer dissepimentarium, forming a pseudonaotic
structure at the peripheral part. The cardinal fossula is distinct and
variably placed on concave or lateral sides of corallites. The
dissepimentarium is made of up to 9 rows of concentric, angulo-
concentric, herringbone, pseudo-herringbone, and occasional
lonsdaleoid dissepiments. The tabularium is normal, ca. 2/3 of
the corallite diameter; it consists of complete to incomplete mesa-
shaped to domal axial tabellae with downturned peripheries.
Description:
External features. Corallites are trochoid, 3.5–6.5 cm high, ca.
5 cm in maximum diameter near the calice of largest corallites.
Calices are deep, partly eroded and still filled with sediment in
some specimens. Apexes are partly preserved. Talons are common
 M. Kora et al. / Geobios 52 (2019) 1–25 5

Fig. 3. Bothrophyllum suezensis nov. sp., holotype (RAr 9). A. External view of the concave side of the corallite; black arrow points to a talon. B. External view of the corallite
(lateral side), showing the positions of the transverse and longitudinal sections. C. Last mature stage at the base of the calyx with wide dissepimentarium and sediment-filled
calice. D–F. Successive ontogenetic stages show reduction in the thickness of major septa in the tabularium with maturity in D. G, H. Successive early immature stages show
the connection of the long cardinal and counter septa and the development of the dissepimentarium in the early mature stage in H. I. Longitudinal thin-section. Black dots
indicate the positions of cardinal septum (below), counter septum (above) and the two alar septa. Scale bars: 5 mm.

at the flanks of the apex in the convex side (Fig. 3(A)), Attachment
scars are also well-developed near the apex of some corallites
(Fig. 7(A)). Growth lines, rugae, septal ridges and furrows are well-
developed on the outer wall of the corallites. Rejuvenation and
geniculation are common (Fig. 5(A1–4, B1)). Almost all specimens
reach the adult stage below their calices, even in the small
corallites that are up to 2 cm in their maximum diameter at the
calice. They show morphological features that are not different
from the larger specimens with maximum calical diameter of 5 cm.
Internal features. Major septa are long up to the late mature
stage, almost reaching the axial part of the tabularium, especially
in the immature stages and withdrawn near the calice. They are
strongly dilated in the tabularium throughout ontogeny, but thin
in the dissepimentarium. A stereoplasmatic thickening of major
septa in the cardinal quadrants is more developed than in the
counter quadrants in the last mature stages (Figs. 3(D, E), 4(A3, A4),
5(A5, A6)). Minor septa are mostly restricted to the outer
dissepimentarium and are continuous or not, forming a pseudo-
naotic structure near the wall. In the adult, stout stage, the number
of septa reaches ca. 43  2 for a 44.8 mm maximum corallite
diameter. Cardinal and counter septa join together throughout
early ontogeny. At maturity; the cardinal septum thins adaxially,
becoming discontinuous as it withdraws (Figs. 4(A2, A3), 7(D)). The
counter septum is commonly as long as the adjacent major septa,
especially in the late mature stages, but is clearly longer in the
early mature stages (Figs. 3(G), 4(A4), 5(A9)). In most growth
stages, major septa are thinner in the dissepimentarium than in the
tabularium (Fig. 3(D-G)). Cardinal fossula is commonly well
marked, invading the dissepimentarium, edged by the underde-
veloped cardinal-lateral major septa and accentuated by the
curvature of sections of tabulae. Its position varies among
individuals of this species. It can be traced in the convex side in
6 M. Kora et al. / Geobios 52 (2019) 1–25

Fig. 4. Bothrophyllum suezensis nov. sp., specimens RAr 21 (A), 33 (B). A1. External view of RAr 21, showing the position of the thin-sections. A2–A6. Serial transverse sections
from the last mature stage below the calice in A2 with thin septa and well-developed pseudonaotic structure, to A6 that shows the last sectioned immature stage near the
apex with almost all the septa joined in the center and the connected cardinal and counter septa. A7. Longitudinal thin-section showing mesa-shaped axial tabellae with
downturned peripheries. B1. External view of RAr 33, showing an Aulopora colony encrusting the upper mature part of the corallite; positions of the successive thin-sections
are marked. B2. External calicinal view of the abraded calice, illustrating the position of the cardinal fossula on the lateral side. B3–B8. Successive transverse thin-sections
from the last mature to the immature stages, all showing the long cardinal septum. Black dots indicate the positions of cardinal septum (below; except for B2, where the
cardinal is located on the lateral side), counter septum (above) and the two alar septa. Scale bars: 5 mm.

some corallites (Figs. 5(A1), 6(A4)) or in the lateral side (Figs. 4(B2),
5(C–F), 7(A)). Dissepimentarium is well-developed in the mature
stages and is made of up to 9 rows of dissepiments in the largest
corallite (Fig. 3(C)). It is composed of concentric, angulo-concen-
tric, herringbone, pseudo-herringbone, and some rare lonsdaleoid
dissepiments only developed in the rejuvenated sites (Figs. 4(A2,
B3, B5), 5(A5, B2), 6(A5, C1)). Lateral dissepiments occur
sporadically in some corallites (Figs. 5(B4–5), 6(C1–3)). Incipient
pseudonaotic structures are poorly developed in late
mature stages; they occur only in the peripheral parts of the
dissepimentarium (Figs. 3(D), 4(A2), 5(A5, A6)). A biform
tabularium is not commonly developed because minor septa are
short. The tabularium and the dissepimentarium are clearly
separated by stereoplasmatic thickening of the innermost row
of dissepimentarium and the septa itself (Figs. 3(E), 4(A4), 5(A6),
7(D, E)), and thus the disstabularium is absent.
In longitudinal section the dissepimentarium consists of regular
globose dissepiments declined towards the tabularium with
angle > 608, the innermost row being almost vertical (Figs. 3(I),
4(A7), 5(A11), 6(B), 7(J)). The tabularium is more or less ca. 2/3 as
 M. Kora et al. / Geobios 52 (2019) 1–25 7

Fig. 5. Bothrophyllum suezensis nov. sp., specimens RAr 7 (A), 8 (B), 18 (C), 19 (D), 32 (E), and 1 (F). A1–A4. External views of the concave, lateral and convex sides of the corallite
RAr 7, showing rejuvenation and geniculation phenomena and the cardinal fossula (arrow) on the convex side (A1). A5. Uppermost mature transverse-section at the base of
the calice, illustrating the effect of rejuvenation on dissepimentarium width and the formation of lonsdaleoid dissepiments. A6–A10. Successive transverse thin-sections from
the adult stage, near the calice with amplexoid septa, to the immature stage near the apex with most septa close to the center. A11. Longitudinal thin-section showing the
effect of rejuvenation on the dissepimentarium and almost horizontal tabulae. B1. External lateral view of RAr 8 showing successive rejuvenations; positions of thin-sections
are marked. B2–B4, B6. Ontogenetic series of transverse sections. B5. Detail of the transverse section B4, showing lateral dissepiments on the major septa. C–F. External views
of the specimens RAr 18, 19, 32 and 1, with position of the cardinal fossula on the lateral side (black arrows). Black dots indicate the positions of cardinal septum (below),
counter septum (above) and the two alar septa. Scale bars: 5 mm.

wide as the corallite diameter. It consists of complete to
incomplete mesa-shaped to domal axial tabellae with downturned
peripheries. There are 8 to 13 tabulae per 5 mm.
Remarks: This new species is different from B. pseudoconicum
Dobrolyubova, 1937 in the following features:
 B. pseudoconicum has more septa for the same diameters
compared to B. suezensis nov. sp. (Fig. 8);
 The pseudonaotic structure is strongly expressed in B. pseudo-
conicum (e.g., Dobrolyubova, 1937: pl. 12, fig. 7; pl.14, figs. 1,2;
pl. 16, figs. 1–3; pl. 17, fig. 1), but poorly developed in the
Egyptian species;
 Minor septa are long and continuous in B. pseudoconicum, but
discontinuous and mostly confined to the outer part of the
dissepimentarium in the present species.
Rodrı́guez (1984b) figured B. pseudoconicum that has a short
cardinal septum and long minor septa, but as described above, the
cardinal septum is longer in the present corals, and the minor septa
are shorter. Comparisons with B. conicum and other Bothrophyllum
species from the Moscovian and Kasimovian of Moscow Basin are
shown in Table 1.
Fedorowski (1975) introduced B. timanioides timanioides from
the lower Kasimovian of Bjørnøya, which has larger number of
8 M. Kora et al. / Geobios 52 (2019) 1–25

Fig. 6. Bothrophyllum suezensis nov. sp., specimens RAr 5 (A), 44 (B), 149 (C). A1, A2. External lateral views of RAr 5 showing the positions of the thin-sections in A2.
A3. External view of the convex side of RAr 5. A4. Calicular view of RAr 5 illustrating the cardinal fossula on the convex side (arrow). A5–A9. Serial transverse thin-sections. B.
Longitudinal thin-section of specimen RAr 44, showing a very weak axial structure in the immature stage and mesa-shaped tabellae with downturned peripheries in the
mature stage. C1. Transverse section at the floor of the calice of specimen RAr 149. C2, C3. Details of C1, showing the presence of lateral dissepiments on the major septa. Black
dots indicate the positions of cardinal septum (below), counter septum (above) and the two alar septa. Scale bars: 5 mm.

septa for the same corallite diameter compared to B. suezensis nov.
sp. He also described B. timanioides nanum from the same horizon,
which differs from B. suezensis nov. sp. by having a short cardinal
septum and by its small corallites. In addition, he described two
additional unidentified species of Bothrophyllum which are also
different from B. suezensis nov. sp., namely Bothrophyllum sp. 1 and
?Bothrophyllum sp. 2. The first species differs in the structure of the
cardinal fossula with a short cardinal septum, larger corallite
diameters, larger number of septa, and longer minor septa than in
the present new species. The second species differs by its short
cardinal septum, the generally short major septa, and strong
thickening of dissepiments. Finally, B. suezensis nov. sp. differs from
B. tenuiseptatum Wang, 1987 from the upper Carboniferous of
central Hunan (China) in having a larger number of septa, and a
wide and complex dissepimentarium.
Bothrophyllum kalmyussi Fedorowski, 2017 and B. gorbache-
vensis Fedorowski, 2017 from the Early Bashkirian of the Donets
Basin are considerably different from B. suezensis nov. sp. B.
kalmyussi is smaller in dimensions, and has a smaller n:d ratio in
mature part (31:11.5 mm vs. 43:44.8 mm in B. suezensis nov. sp.).
 M. Kora et al. / Geobios 52 (2019) 1–25 9

Fig. 7. Bothrophyllum suezensis nov. sp., specimen RAr 6. A–C. External concave, convex and lateral views, respectively, with positions of transverse and longitudinal thin-
sections indicated in C. D–I. Ontogenetic series of transverse sections, from adult to immature stages, showing regular thick septa in all stages. J. Longitudinal sections as
marked in C. Black dots indicate the positions of cardinal septum (below), counter septum (above) and the two alar septa. Scale bars: 5 mm.

The cardinal fossula of the former species is closed, while open in B.
suezensis nov. sp. It also differs in the arrangement of the major
septa in both early and advanced maturity. B. gorbachevensis
Fedorowski, 2017, differs in the n:d ratio at maturity (35:15.5 mm,
much smaller than in B. suezensis nov. sp.) and in the underdevel-
opment of the cardinal fossula.
B. proteum Semenoff-Tian-Chansky, 1974 from the early
Bashkirian of the Béchar Basin in Algeria is protocolonial. This
feature is totally unknown in other species of Bothrophyllum
allowing Fedorowski (2016) to exclude the Algerian specimens
from the genus Bothrophyllum. Flügel (1993) described a new
Viséan B. baculonodosum and a new Sakmarian (lower Permian) B.
asseretoi from North Iran, but Fedorowski (2016) excluded both
species from Bothrophyllum and included them into the Hetero-
caniniidae Hill, 1981. In fact, both species are morphologically
different from B. suezensis nov. sp., in means of the more complex
axial structure, their long minor septa, the numerous septa they
possess and the dissepimentarium. Corals similar to B. suezensis
nov. sp. were previously included into the cyathopsid genus
Pseudozaphrentoides by de Groot (1963), Rodrı́guez (1984b) and
Rodrı́guez et al. (1997), as discussed by Fedorowski (1975, 1987,
2016). B. suezensis nov. sp. differs from Pseudozaphrentoides
suevensis Rodrı́guez et al., 1997 from the lower Moscovian of
Eastern Asturias (Spain) by the smaller length of the corallites, the
10 M. Kora et al. / Geobios 52 (2019) 1–25
Fig. 8. Relationship between the number of major septa and corallite alar diameter
of Bothrophyllum suezensis nov. sp., B. cylindricum nov. sp., and B. pseudoconicum
Dobrolyubova,1937. Corallites with largely eroded dissepimentarium are excluded.
smaller number of septa at the same diameter, and its broader and
more complex dissepimentarium. Also P. melendezi Rodrı́guez,
1984b has smaller number of septa at the same diameter. It is also
different in having long minor septa, whereas they are short in B.
suezensis nov. sp. B. suezensis is smaller and has a more complex
dissepimentarium than P. rabanaliensis de Groot, 1963 from the
lower Moscovian of Northern Palencia (Spain).
Small sized, fairly immature corallites are commonly repre-
sented in the studied material. They are included in B. suezensis
nov. sp. as they display all the features of the immature stages of
the species. This group of corals is here included in the family
Bothrophyllidae and in the genus Bothrophyllum because of the
morphology of their immature stages that are characterized by
long major septa and the axial connection of cardinal and counter
septa. Fedorowski (2016) suggested that B. pseudoconicum
Dobrolyubova, 1937, with both types of tabularia (normal and
biform) and with the cardinal septum temporarily shortened,
forms an intermediate evolutionary step, leading towards bothro-
phyllids lacking a biform tabularium. This is also the case for B.
suezensis nov. sp.
Bothrophyllum cylindricum nov. sp.
Fig. 9
1988. Pseudozaphrentoides ex. gr. juddi (Thomson, 1893) –
Herbig and Kuss, p. 9–11, figs. 6e–k, 7c.
Derivation of the name: From the cylindrical, nearly straight
shape of the corallites.
Holotype: Specimen RAr 22 (Fig. 9); seven transverse and one
longitudinal thin-sections available.
Type locality: Wadi Rod El Hamal (29811’01’’N, 32825’59’’E),
Wadi Araba area, northern Eastern Desert, western side of the Gulf
of Suez, Egypt.
Type horizon: Rod El Hamal Fm., Unit 3, late Moscovian.
Material: Four well-preserved corallites, including the holotype
(RAr 22) and three paratypes (RAr 23, 46, 150), with 10 transverse
and 2 longitudinal thin-sections.
Diagnosis: Bothrophyllum with 35  2 septa for a diameter of
19.5 mm at the calice floor; long cardinal and counter septa that
join in the early neanic stage to form an axial septum; rudimentary
minor septa; open cardinal fossula on the convex side; up to 5 rows
of concentric and herringbone dissepiments; complete and/or
incomplete mesa-shaped tabulae.
Description:
External features. Corallites are cylindrical to trochoid, up to
6 cm in height. Calice is deep. Growth lines, rugae and rejuvenation
are very marked. Talons and attachment scars are visible near the
apex (Fig. 9(A1–3)).
Internal features. Major septa are long, with 35 septa for a
maximum diameter of 19.5 mm at the calice floor, quite strongly
thickened in the tabularium at mature stage (Fig. 9(A6, B4–B5)),
but thin in the dissepimentarium at any stage. Almost all the major
septa meet in the centre of the corallite in early immature stage,
then gradually withdraw in late mature stages. The cardinal and
counter septa are long and join in the early immature stage,
forming an axial septum (Fig. 9(A10–11, B9)). In advanced stages,
the cardinal septum thins adaxially in the tabularium and becomes
discontinuous (Fig. 9(A9, B3–8)). The counter septum is as long as
the other major septa in the counter quadrants, or slightly longer.
Minor septa are variable in size; long and crossing 3-4 rows of
dissepiments (some discontinuously) to reach the inner wall and
extend into outer tabularium as dilated septal pegs in the cardinal
quadrants (Fig. 9(A5, A9)), short and only crossing the outer row of
dissepiments (Fig. 9(B3–8)). Cardinal fossula open, located always
on the convex side. Dissepimentarium starts to develop during the
early immature stages (Fig. 9(A11)), at a diameter of 7 mm. The
dissepimentarium is made of up to 5 rows of concentric and
herringbone dissepiments. They are regular globose in longitudinal
section. Some rare lonsdaleoid dissepiments occur during rejuve-
nescence phases (Fig. 9(B3)). Tabularium is complete with mesa-
shaped tabulae or incomplete, showing up to 9 tabulae / 5 mm
length (Fig. 9(A12, B10)).
Remarks: The Egyptian new species differs from Pseudoza-
phrentoides species by its long major septa in the early immature
stage, whereas those of Pseudozaphrentoides are short and
amplexoid (Fedorowski, 1975, 2016). Specimens described from
Bir Madsus in the Wadi Araba area by Herbig and Kuss (1988) as
Pseudozaphrentoides ex. gr. juddi (Thomson, 1893) are included in
Bothrophyllum cylindricum nov. sp. Note that Herbig and Kuss
(1988) described the cardinal to be short in the immature stage,
where all the septa are long and joined at the axis, but from the
illustrations the cardinal septum is indeed long. The specimens are
also comparable in the general cylindrical shape, the corallite
diameter and number of septa, the long cardinal and counter septa
that are joined in the centre, and all tabularium and dissepimen-
tarium characters.
Bothrophyllum cylindricum nov. sp. differs from B. suezensis nov.
sp. in the following characters:
 Although reaching nearly the same corallite length as the
holotype of B. suezensis nov. sp., corallites of B. cylindricum nov.
sp. are more slender;
 Regarding the relation of corallite diameter vs. number of septa,
they reach up to 35 septa for 19.5 mm maximum corallite
diameter, whereas in B. suezensis nov. sp. there are 43 septa for a
maximum 44.8 mm diameter (Fig. 8);
 At same diameters, B. suezensis nov. sp. displays thickened septa;
 The species is also different from B. suezensis nov. sp. by its
cylindrical corallite shape and the less complex and narrower
dissepimentarium.
The other Bothrophyllum species all differ from B. cylindricum
nov. sp., as can be seen in their comparison to B. suezensis nov. sp.
and in Table 1.
Bothrophyllid gen. indet. A sp. indet. 1
Fig. 10(A1–4, B1–4)
Material: Two corallites (RAr 77, 93) collected from unit 3 of the
Rod El Hamal Fm. (late Moscovian), west of Wadi Rod El Hamal
(29809’39.90’’N, 32824’36.50’’E), Wadi Araba area, northern
Eastern Desert, western side of the Gulf of Suez, Egypt. Two
transverse and two longitudinal thin-sections available.
Description:
External features. Corallites are small, solitary, slightly curved,
and ca. 2–2.5 cm long. They are moderately preserved with partly
eroded calices. Calices are deep. External wall partly eroded to
Table 1
Comparison of the Moscovian and Kasimovian Bothrophyllum spp. from the type region of the Moscow Basin with Bothrophyllum suezensis nov. sp., B. cylindricum nov. sp. and further undetermined Egyptian Bothrophyllidae.

Species n:d of last Cardinal septum Cardinal fossula Counter septum Major septa Minor septa Dissepimentarium Tabularium Age and locality
mature stage

Bothrophyllum Variable, Long up to late Well developed, Elongated Long up to late Penetrating deeply Wide, max. 12 rows Biform peripheral Moscovian,
conicum 44:30  33 for mature stage closed throughout mature stage, a little into the tabularium of dissepiments in the tabularium Moscow Basin
Trautschold, largest corallites growth withdrawn in most with their inner most advanced stage,
1879, advanced growth sages, margins thickened, mostly simple concentric,
emend. joined in the centre to thinner in herring-bone, angulo-
Dobrolyubova, form a loose axial dissepimentarium concentric dissepiments
1937 structure, commonly
thickened in the cardinal
quadrants
Bothrophyllum 40:25  27 Long; strongly Well developed, Long in early Strongly thick in the Very long, attain half Wide, up to 11 rows Up to 14 tabulae per Kasimovian,
conicum thickened at the open stages of growth tabularium in all growth of the length of major of irregular, concentric, 5 mm length, complete Moscow Basin
robustum periphery and and equal to stages, in mature stages septa late in maturity, angulo-concentric, and/or incomplete,
Dobrolyubova, thread-like thin in adjacent major thicker in cardinal contrajunct, some herringbone interseptal, with flat surfaces and
1940 the inner part septa in mature quadrants than in contratingent, thick arched dissepiments; downturned edges
stage counter quadrants, thin in tabularium, thin in pseudonaotic structure (mesa-shaped)
in the dissepimentarium the dissepimentarium occurs in mature stages;
like the major septa in longitudinal sections
more or less regular
globose dissepiments

M. Kora et al. / Geobios 52 (2019) 1–25

inclined towards the
tabularium with
angle >608, the
innermost row being
almost vertical
Bothrophyllum 45:36 Variable in length Indistinct Elongated, as Long up to late mature Variable in length; Regular to irregular Normal and/or biform Moscovian,
pseudoconicum in the advanced long as the stage, a little withdrawn penetrate the concentric, herringbone tabularium, tabulae Moscow Basin
Dobrolyubova, growth stages adjacent major in most advanced growth tabularium when dissepiments dominate slightly convex and
1937 septa stages, joined in the dissepimentarium is the inner part which is split into vesicles
centre to form a loose narrow, but do not not intersected by minor
axial structure, enter when wide, septa, pseudonaotic
commonly thickened in often reach 1/4 and structure occurs in the
the cardinal quadrants never exceed 1/3 outermost part
of the major septal
length
Bothrophyllum 26:9 Slightly shorter Not marked Long Long, joined in the centre Do not exceed 1/3 of Small, irregular Steeply inclined axial Kasimovian,
complexum than the adjacent to form a complex axial the major septal length, dissepiments forming a tabellae border the Moscow Basin
Dobrolyubova, cardinal-lateral structure thickened in the ring not exceeding central median lamella,
1937 septa tabularium 1 mm width, separated flat and moderately
from the tabularium inclined peri-axial
by a thick stereozone tabellae
Bothrophyllum 40:17 Short from the Narrow, closed Long in early Long up to the late Vary in length; Very narrow, simple, Small, irregular, Moscovian,
kashiricum early mature stage, stages of growth, mature stage, withdrawn protruding in the 1–2 rows of small vesicular axial tabellae; Moscow Basin
Kabakovich, hardly does not differ near the calice tabularium for a very concentric dissepiments, horizontal in the
1937 distinguished in the from the adjacent short distance in the not developed in places centre, larger, more
last mature stage major septa in youngest stages, reach regular obliquely
mature stages about 1/4 of the major downwards near the
septal length in the last periphery, peri-axial
mature stage tabellae steeply rising
towards the
dissepimentarium
Bothrophyllum 35:12 Long up to the last Not marked Long, may reach Long, some reach 2/3 of Very short, mostly Narrow, simple, 2–3 rows Divided into two parts; Moscovian,
okense Kossovaya, mature stage the centre and the radius and confined to the of simple inter-septal peri-axial vesicular Moscow Basin
2001 join to the intersecting with the dissepimentarium in all dissepiments tabellae and flat to
cardinal septum tabulae, forming a non- growth stages, some cone-shaped axial
permanent axial enter the tabularium as tabellae

11
structure, thickened in thick pegs in the
the cardinal quadrants cardinal part
 12
Table 1 (Continued )

Species n:d of last Cardinal septum Cardinal fossula Counter septum Major septa Minor septa Dissepimentarium Tabularium Age and locality
mature stage

Bothrophyllum 43:44.8 Long, joins with Well marked, As long as the Long up to the late Short, mostly re- Wide, up to 9 rows of About 2/3 as wide as Late Moscovian,
suezensis nov. sp. the counter invading the adjacent major mature stage, reaching stricted to the outer concentric, angulo- the diameter of the Egypt
septum dissepimentarium, septa in the late the axial part of the dissepimentarium, concentric, herringbone, corallite, 8–13 tabulae
throughout early edged by the under- mature stages, tabularium especially in forming pseudonaotic pseudo-herringbone, per 5 mm, complete to
ontogeny, thins developed cardinal- longer in the the immature stages; structure at the occasional lonsdaleoid incomplete mesa-
and becomes lateral major septa, early mature withdrawn near and in peripheral part, and lateral dissepiments, shaped to domal axial
discontinuous variably placed on stages the calice, dilated in the continuous to pseudonaotic structure tabellae with
adaxially or concave or lateral tabularium throughout discontinuous occurs in the peripheral downturned
withdraws at sides of corallites ontogeny, but thin in the part in late stages; in peripheries
maturity dissepimentarium longitudinal sections
regular globose
dissepiments declined
towards the tabularium
with angle >608,
innermost row being
almost vertical
Bothrophyllum 35:19.5 Long, joins with Distinct, open, located As long as the Long, quite strongly Variable in size; either Five series of rows of Complete and/or Late Moscovian,
cylindricum the counter on the convex side of other major septa thickened in the short and restricted to concentric and incomplete mesa- Egypt
nov. sp. septum in the the corallite in the counter tabularium at mature the outer row of the herringbone shaped tabulae, up to
early immature quadrants, or stage, but thin in the dissepimentarium, or dissepiments, regular 9 tabulae per 5 mm

M. Kora et al. / Geobios 52 (2019) 1–25

stage forming an slightly longer dissepimentarium at long crossing 3–4 rows globose in longitudinal length
axial septum, thins any stage, almost all of dissepiments section
adaxially in the septa join in the centre in
tabularium and early immature stage,
becomes gradually withdraw in
discontinuous in late mature stages
advanced mature
stages
Bothrophyllid 29:12–15 Slightly shorter Indistinct Same length as Long, thin in the About 1/4–1/2 of the Narrow, with max. 3 Biform, flat axial Late Moscovian,
gen. indet. than the other the adjacent dissepimentarium, major septa in length, rows of concentric, tabellae, peri-axial Egypt
A sp. indet. 1 adjacent major major septa slightly thickened in the continuous in the small herringbone to tabellae steeply
septa tabularium, join in the dissepimentarium irregular interseptal declined towards the
centre forming a loose where they have the dissepiments, globose dissepimentarium
and irregular network same thickness of the or elongated in where they become flat
together with the extra- major septa, longitudinal section
septal lamellae of the discontinuous in the
minor septa tabularium, can be
traced near to the
centre forming extra-
lamella
?Bothrophyllid 31:15 Long Distinct, edged by Same length as Long, reach the centre Nearly confined to the Narrow, with max. 3 Nine axial tabellae per Late Moscovian,
gen. indet. bent cardinal-lateral the adjacent of the tabularium dissepimenta-rium in rows of regular 5 mm, dome-shaped or Egypt
B sp. indet. 1 major septa, supports major septa without connecting the counter quadrants, dissepiments, the mesa-shaped axial
a long and curved and leaving a narrow enter the tabularium innermost row being tabulae with almost
cardinal septum free tabular area for a very short distance thickened, globose, vertical shoulders, flat
in the cardinal almost vertically peri-axial tabellae
quadrants, reach 1/3 inclined in the
of the major septal longitudinal section
length in the fossular
breaks
?Bothrophyllid 31:12 Short in the fossula Located on the convex Same length as Long, meet in the About 1/3 of the Narrow, consists of small, Biform tabularium, Late Moscovian,
gen. indet. at the last mature side of the corallite, the adjacent centre at all growth major septa in length irregular dissepiments, steeply inclined axial Egypt
C sp. indet. 1 stage marked only on the major septa stages, forming a separated from the tabellae, flat and
last mature stage, near complex columella tabularium by a thick moderately inclined
the calice floor stereozone formed by peri-axial tabellae
thickening of the minor
and major septa
 M. Kora et al. / Geobios 52 (2019) 1–25 13

Fig. 9. Bothrophyllum cylindricum nov. sp., specimens RAr 22 (Holotype, A), 23 (B). A1–A3. External views; lateral, concave, convex sides of the corallite RAr 22 (holotype)
showing the weak constrictions, rejuvenescences, and marked rugae on the wall; positions of thin-sections marked in A1. A4. Calicinal view showing the fossula on the convex
side. A5–A11. Successive transverse thin-sections illustrating the ontogenetic development. A12. Longitudinal section. B1–B2. Lateral and concave views of specimen RAr 23,
respectively, showing repeated rejuvenescence. B3–B9. Successive transverse sections according to the positions marked in B1, starting from the late mature stage at the
calice floor in B3. B10. Longitudinal thin-section as marked in B1. Black dots indicate the positions of cardinal septum (below), counter septum (above) and the two alar septa.
Scale bars: 5 mm.

well-preserved. Concentric wrinkles and very delicate septal ridges
are visible on the well-preserved parts of the wall. Attachment
scars and a talon are seen at the apex flank (Fig. 10(B1, 2)).
Internal features. Major septa are long, thin in the dissepimen-
tarium, slightly thickened in the tabularium, and then again thinning
axially. They join in the centre, forming a loose and irregular network
together with the extraseptal lamellae of the minor septa. There are
29 septa at a diameter of 12–15 mm, just below the calice. Minor
septa are ca. 1/4 of the major septa in length and enter the tabularium
for a short distance. They have the same thickness as the major septa
in the dissepimentarium where they are continuous. In the counter
quadrants (convex side of the corallite), the dissepimentarium is
wider and the minor septa are ca. 1/2 of the major septa in length
(Fig. 10(A3)). The minor septa can be traced again in the tabularium
near the centre, forming extralamella. Major and minor septa are
undulated in the tabularium (Fig. 10(A3, B3)). The cardinal fossula is
indistinct. The cardinal septum is slightly shorter than the other
adjacent major septa in the last mature stage (Fig. 10(A3)), whereas
the counter septum has the same length as the other major septa. The
dissepimentarium is narrow, with maximum 3 rows of dissepiments.
Dissepiments are concentric, small herringbone to irregular inter-
septal, with stereoplasmatic thickening, appearing globose or
elongated in longitudinal section. The tabularium is biform
(Fig. 10(B4)). It is divided; the axial tabellae are rather flat, the
14 M. Kora et al. / Geobios 52 (2019) 1–25

Fig. 10. Indeterminate bothrophyllid genera and species. A, B. Bothrophyllid gen. indet. A sp. indet. 1, specimens Rar 93 (A), 77 (B). A1, A2: External lateral and concave views
of specimen RAr 93; position of thin-sections indicated; A3, A4: Transverse and longitudinal sections; B1, B2: External views of specimen RAr 77 showing the attachment scar
and a small talon at the flank of the apex and weak lateral constrictions in the adult stage in B2; B3: Transverse section, showing the extra-septal lamellae in the tabularium
center and the biform tabularium; B4: Longitudinal section, oblique in its lower part, showing the biform tabularium (arrow). C. ?Bothrophyllid gen. indet. B sp. indet. 1,
specimen RAr 106. C1: External convex side with partly eroded wall; C2: External view of the calice with eroded margin; C3: Transverse section below the calice with thick
long major septa and short minors; C4: Longitudinal thin-section showing a wide tabularium and a narrow dissepimentarium. D. ?Bothrophyllid gen. indet. C sp. indet. 1,
specimen RAr 92. D1, D2: External lateral and concave views showing an attachment scar and talon in the juvenile part in D1; position of thin-sections indicated; D3, D5:
Successive transverse sections below the calice; D4: Detail of the axial structure in D3 with a thick median lamella; D6: Longitudinal section showing the axial structure and
the axial lamella; D7: Detail of the outer part of the dissepimentarium of transverse section D3, showing the undulating minor and major septa, the stereoplasmatic
thickening in the narrow dissepimentarium and the biform tabularium. Black dots indicate the positions of cardinal septum (below), counter septum (above) and the two alar
septa. Scale bars: 5 mm, except D4 and D7: 1 mm.

peri-axial tabellae are steeply declined towards the dissepimenta- extra septal lamellae. However, B. conicum var.1 has numerous
rium where they become flat (Fig. 10 (A4, B4)). septa (36  2 for 10–12 mm diameter in the adult stage) compared
Remarks: The species shows some similarities to Bothrophyl- to the present Egyptian coral that has less septa at larger diameter
lum conicum var.1 Dobrolyubova, 1940 in its axial structure with (29  2 septa for a diameter of 15 mm in the adult stage).
 M. Kora et al. / Geobios 52 (2019) 1–25
?Bothrophyllid gen. indet. B sp. indet. 1
Fig. 10(C1–4)
Material: One corallite (RAr 106) with one transverse and one
longitudinal thin- section, collected from the late Moscovian unit
3 of the Rod El Hamal Fm. (29811’01’’N, 32825’59’’E), Wadi Araba
area, northern Eastern Desert, northern Eastern Desert, western
side of the Gulf of Suez, Egypt.
Description:
External features. Fragment of a solitary cylindrical (?) coral,
moderately preserved with worn wall. Calicinal rim and apical part
are largely eroded. Septa in the calice are long, thick and meet in
the centre. Two alar fossulae are visible (Fig. 10(C2)). Tabular floor
slightly raised in the middle of the calice.
Internal features. There are 31  2 septa at a diameter of
15 mm. Major septa are long and reach the centre of the
tabularium without connecting, leaving a narrow free tabular
area. Minor septa are short, nearly confined to the dissepimenta-
rium in the counter quadrants. In the cardinal quadrants, they
enter the tabularium for a very short distance, except for those
located in the cardinal and the alar fossulae; they are long and
reach more or less ca. 1/3 of the major septal length. Cardinal and
counter septa are long. All septa are thin in the dissepimentarium
and thickened in the tabularium. The cardinal fossula is edged by
bent cardinal-lateral major septa and supports a long and curved
cardinal septum. The two alar fossulae are distinct (Fig. 10(C3)).
The narrow dissepimentarium consists of maximum 3 rows of
regular dissepiments, the innermost row being thickened.
In longitudinal section (Fig. 10(C4)), the dissepiments are
globose, almost vertically inclined. The axial tabulae are dome-
shaped or mesa-shaped with almost vertical shoulders. Peri-axial
tabellae are flat and fill a peripheral gutter. There are 9 axial
tabellae per 5 mm, some of them being thickened. In the upper part
of the corallite, they are piled up and form an incipient
columnotheca (sensu Fedorowski, 2009).
Remarks: The long cardinal and counter protosepta of this
specimen would plead for its inclusion into the Bothrophyllidae;
however, its tabularium differs considerably from that of all
bothrophyllid species. This suggests a position of this coral
specimen in a new genus. Nevertheless, more material in a better
preservation stage would be necessary to exclude open nomen-
clature.
?Bothrophyllid gen. indet. C sp. indet. 1
Fig. 10(D1–7)
Cf. 1937. Bothrophyllum complexum Dobrolyubova – p. 37–39,
75, pl. 10, figs. 12–19.
Material: Two small corallites (RAr 92, 100) from unit 3 of the
Rod El Hamal Fm., late Moscovian, west of Wadi Rod El Hamal
(29809’39.90’’N, 32824’36.50’’E), Wadi Araba area, northern
Eastern Desert, western side of the Gulf of Suez, Egypt. One
transverse and one longitudinal thin-section, and 3 acetate peels
available.
Description:
External features. Small, solitary, slightly curved corallites,
maximum 2 cm high. Moderately well-preserved with deep, partly
eroded calices. Constrictions are common. Wall partly eroded, with
fine striations where better preserved. Talon and attachment scar
are visible at the apical flank. The attachment was to the small
branched tabulate coral Aulopora sp. (Fig. 10(D1)).
Internal features. Long major septa meet in the centre at all
growth stages, forming a complex columella. There are 31 major
septa for a diameter of 12 mm; they are undulated, commonly by
diagenesis (Fig. 10(D3, 7)). Septa are thick in the dissepimentarium
and the outer tabularium and thin towards the axis. Cardinal
septum short in the fossula at the last mature stage (Fig. 10(D3));
the fossula is located on the convex side of the corallite. Counter
15
septum has the same length as the adjacent major septa. Minor
septa are ca. 1/3 of the majors in length. The narrow dissepi-
mentarium consists of small, irregular dissepiments. It is better
developed in the cardinal quadrants. It is separated from the
tabularium by a thick stereozone formed by thickening of the
minor and major septa (Fig. 10(D7)). A network of thick tabulae
and thick septal lamellae with a medial plate composes a complex
columella (Fig. 10(D3–5)). In longitudinal section, the axial
structure appears bracketed by steeply inclined axial tabellae.
The tabularium is biform. The peri-axial tabellae are flat and
moderately inclined, forming a peripheral gutter (Fig. 10(D6)).
Remarks: This species is closely similar to Bothrophyllum
complexum Dobrolyubova, 1937; it differs in having larger n:d ratio
in the adult (31:12 in the present species vs. 26:9 in the former
species). Dobrolyubova (1937) doubtfully assigned her species to
the genus Bothrophyllum in spite of the differences mentioned in
the original description. These are occurrences of a complex axial
structure in mature stages and of a thick stereozone produced by
the thickening of the septa. These characters are also present in the
current material. Dobrolyubova (1937) finally suggested to
separate B. complexum as a new genus and to include it in the
subfamily Clisiophyllinae, in spite of some differences in longitu-
dinal sections. Similarly, Fedorowski (2016) listed B. complexum
Dobrolyubova, 1937 as a potential member of a different genus or
subgenus. The complex axial structure and the thick stereozone
and the irregular dissepiments may suggest affinity with the
Geyerophyllidae, thus its position in the Bothrophyllidae is also
questionable.
Suborder Lonsdaleiina Spasskiy, 1974
Family Geyerophyllidae Minato, 1955
Genus Amygdalophylloides Dobrolyubova et Kabakovich, 1948
Type species: Amygdalophyllum ivanovi (Dobrolyubova, 1937).
Included species: see Minato and Kato (1975), plus A. uzurense
(Yamagiwa et Ota, 1963) of Minato and Kato, 1975; A. gracilis
(Hayasaka, 1924) of Minato and Kato, 1975; A. (?) degrootae
Rodrı́guez, 1984b; A. liebanensis Rodrı́guez, 1984b; A. densus
Yoshida et Okimura, 1992; A. denticulatus Yoshida et Okimura,
1992; A. omiensis Yoshida et Okimura, 1992; A. longiseptatus
Yoshida et Okimura, 1992; A. pravus Yoshida et Okimura, 1992; A.
wuwangshie Igo et Kamikawa, 1998; A. anticuum Rodrı́guez et Said,
2009; A. omarai nov. sp.
Age and distribution: Amygdalophylloides was previously
recorded from the Moscow basin, Spitsbergen, Northern Spain,
Carnic Alps, Yugoslavia, Japan, China, North Vietnam, and Egypt.
Ranging in age from the late Viséan (‘‘Namurian’’) to the lowermost
Permian and reaching its maximum distribution during the late
Moscovian (Minato and Kato, 1975).
Diagnosis (after Hill, 1981, and Minato and Kato, 1975): Small
solitary corallites; ceratoid to almost straight; major septa
reaching or approaching the thick columella; minor septa
commonly short or not developed; calice deep; axial structure
made of a solid columella, oval in section, with or without
irregularly arranged and poorly developed axial tabellae and a
median lamella originating from the cardinal septum; septa
thickened peripherally to form a narrow stereozone; dissepiments
regular to somewhat irregular; lonsdaleoid dissepiments sporadic
and steeply inclined; clinotabulae, wide and merging with narrow
horizontal peri-axial tabulae.
Remarks: Differences between Amygdalophylloides and other
geyerophyillid genera were discussed in Rodrı́guez (1985), and
with a special focus on gregarious growing genera by Rodrı́guez
and Bamber (2012).
Amygdalophylloides omarai nov. sp.
Figs. 11, 12
16 M. Kora et al. / Geobios 52 (2019) 1–25

Fig. 11. Amygdalophylloides omarai nov. sp., specimens RAr 159 (holotype, A), 165 (B), 3 (C). A1–A3. External views of the concave, convex and lateral sides of the specimen RAr
159 (holotype), respectively, with several rejuvenescences; position of thin-sections indicated. A4. Uppermost mature transverse section below the calice, showing irregular
columella, short counter septum (above), and partly eroded outer dissepimentarium. A5. Details of the irregular columella in A4. A6, A8, A10. Successive transverse sections at
the lower immature part of the corallite, with cardinal septum connected to the columella and lonsdaleoid dissepiments in A6. A7, A9. Detail of the columella in A6 and A8,
respectively, to show the oval-shaped columella, formed of septal lamellae and axial tabellae. A11. Longitudinal thin-section showing the thick columella with irregular
margins and internal voids. B1, B2. External views of the corallite RAr 165 with partly eroded wall; note the presence of rootlets on the lateral side of the corallite and the
tabulate Aulopora sp. encrusted near the calice. B3–B5. Successive transverse sections from the upper adult to the lower immature stage near the tip (sections marked in B2),
showing also the rootlets affecting the outer dissepimentarium and the septa in B4 and B5. B6. Two longitudinal thin-sections showing the thick columella. C1, C2. External
views of the corallite RAr 3 illustrating broken rootlets at the flanks of the tip in C2 (arrow). C3. Longitudinal section illustrating the connection of the columella to the cardinal
septum, and the deep calyx filled with micritic peloidal grains. C4. Drawn transverse section in the calical part, showing outer lonsdaleoid dissepiments. Black dots indicate
the positions of cardinal septum (below), counter septum (above) and the two alar septa. Scale bars: 5 mm, except A7 and A9: 2.5 mm.

1988. Amygdalophylloides ivanovi (Dobrolyubova) – Herbig and
Kuss, p. 15–16, 18–19, figs. 6a–d, 7a–b.
Derivation of the name: The species is named in the memory of
the late Egyptian palaeontologist Prof. Sayed Omara who
contributed to the stratigraphy and palaeontology of Wadi Araba.
Holotype: Specimen RAr 159 (Fig. 11(A1–A11)); four transverse
and one longitudinal thin-sections available.
Type locality: West of Wadi Rod El Hamal (29809’39.90’’N,
32824’36.50’’E), Wadi Araba area, northern Eastern Desert, western
side of the Gulf of Suez, Egypt.
Type horizon: Unit 3 of the Rod El Hamal Fm., late Moscovian.
Material: Paratypes: 8 moderately to well-preserved small to
middle-sized solitary curved corallites (RAr 3, 96, 109, 135, 137,
164–166), with 19 transverse and 10 longitudinal thin-sections.
 M. Kora et al. / Geobios 52 (2019) 1–25 17

Fig. 12. Amygdalophylloides omarai nov. sp. (small-sized corallites), specimens RAr 135 (A), 164 (B), 96 (C). A1. External view of specimen RAr 135; position of thin-sections
indicated. A2, A3. Two transverse sections in the adult stage below the calice show the oval columella connected to the cardinal septum. A4. Detail of the columella in A3.
A5. Close-up of the cardinal part in transverse section A3, showing the cystoporate bryozoa Eridopora sp. encrusting the external wall of the corallite. A6. Longitudinal section
illustrating thick columella, wide tabularium and narrow dissepimentarium. A7. Transverse section near the apex at the early immature stage, illustrating the cardinal
septum which forms the incipient columella. B1. External view of the corallite RAr 164 with lateral small rootlets; position of thin-sections indicated. B2. Transverse section at
the last mature stage below the calice, which is partly filled with fossiliferous peloidal micrite; note irregular columella, tabulae-like dissepiments and lonsdaleoid
dissepiments at the partly eroded outermost dissepimentarium. B3, B4. Successive transverse sections at the immature stages, showing the connection to the cardinal septum
in B4. B5. Longitudinal thin-section, marked in B1, showing the thick columella and in part collapsed tabulae. C1. External view of the corallite RAr 96 with a deformed calice
due to growth constriction. C2, C3. Successive transverse thin-sections marked in C1, showing the irregular columella in C2 and the more regular, oval columella in C3.
C4. Longitudinal thin-section illustrating the thin columella at the immature stage and thickening with maturity. Black dots indicate the positions of cardinal septum (below),
counter septum (above) and the two alar septa. Scale bars: 5 mm, except A4 and A5: 1 mm.

Thirty-one small corallites were studied only for their external
morphology (RAr 101, 110, 129–132, 134, 136, 138, 139, 140–143,
169, 171–176, 178–187). All were collected from the late
Moscovian unit 3 of the Rod El Hamal Fm. at Wadi Araba.
Diagnosis: Solitary, ceratoid to conical corallites, 15–35 mm
high with maximum diameter of 17.5 mm, having maximum of
32 major septa. The columella is thick and composed of a median
lamella and thick septal lamellae.
Description:
External features. Small to middle-sized ceratoid to straight
conical solitary corallites with heights from 15–35 mm. The
ceratoid corallites are strongly curved at the young stage and
nearly straight at the adult stage. Fairly well-preserved specimens;
all show a partly eroded wall and therefore the peripheral margins
of the dissepimentarium and septa are exposed. Calices are
abraded, deep, with a slightly projected axial structure. Apexes are
often preserved. Growth lines, rugae, septal furrows and ridges can
be traced on the well-preserved parts of the wall. Attachment scars
occur at the apex flanks in almost all specimens. Constrictions and
rejuvenescence are common.
Internal features. Corallite diameter in last adult stage, below
the calice floor, varies from 13–16 mm; number of major septa is
27–32. Major septa are long and almost reach the columella, but
are not attached at the adult stages. They are generally straight
18 M. Kora et al. / Geobios 52 (2019) 1–25

Table 2
Comparison of Amygdalophylloides omarai nov. sp. with some Amygdalophylloides spp. from Spain, Moscow, and Japan.

Description A. ivanovi A. (?) degrootae A. liebanensis A. anticuum A. omarai nov. sp.

(Dobrolyubova, 1937) Rodrı́guez, 1984 Rodrı́guez, 1984 Rodrı́guez et Said,
2009

Shape Small, almost straight, Small, ceratoid Small, ceratoid, Small, ceratoid Seratoid, straight or
cone-shaped erect or slightly slightly curved
curved
Max. corallite diameter 10–14 mm 10–15 mm 8–11 mm 5.5–8 mm 17–17.5 mm
Maximum nb. of major septa 26 24–30 22–24 25–28 28–32
Length of minor septa compared to 2/3 About 1/2 1/3–1/2 About 1/2–2/3 About 1/2 or less
major septa
Dimensions of axial structure with About 1/6 Less than 1/10 Do not exceed 1/5 Less than 1/4 1/6–1/3
respect to the corallite diameter in
longitudinal section
Dissepiments Small interseptal 3–4 rows of regular 2–3 rows of regular 1–2 rows of 1st and 2–3 rows of angular
dissepiments interseptal interseptal 2nd order or v-shaped
dissepiments dissepiments lonsdaleoid interseptal and
lonsdaleoid
Age and locality Upper Moscovian Upper Moscovian Upper Moscovian Upper Viséan, S Upper Moscovian,
(Myachkovian)– (Podolskian), N (Podolskian), N Spain NE Egypt
Kasimovian, Moscow Spain Spain
Basin
Upper Carboniferous–
lower Permian, Carnic
Alps, Bosnia and
Herzegovina
Upper Moscovian
(Podolskian), N Spain

with axial ends bent towards the cardinal septum. Septa are longer (Dobrolyubova, 1937). This character approaches it to repre-
in the cardinal quadrants but withdrawn in counter quadrants; sentatives of the genus Kionophyllum Chi, 1931, but the latter
counter septum is shortest (Fig. 11(A4, A6)). The cardinal septum is display a complete and well-developed lonsdaleoid dissepi-
long and connected to the solid columella in the early immature mentarium (Rodrı́guez, 1985);
stages. In some corallites it becomes detached in the adult stages  Amygdalophylloides omarai nov. sp. has larger corallites (up to
(Figs. 11(A4, B3, C4), 12(B2, B3)), but in others it is still attached up 3.5 cm: holotype, Fig. 11(A1–3)), more septa (max. 32  2) and a
to the last mature stage (Fig. 12(A2, A3)). Minor septa are ca. 1/2 as larger diameter (max. 16 mm) than A. ivanovi. In the latter,
long as the major septa, or less; they either reach the edge of the maximum 26  2 septa are developed for a diameter ranging
tabularium or enter for a short distance. A septal stereozone is between 10 and 14 mm (Fig. 13);
well-developed in the outermost tabularium. The columella is oval  Amygdalophylloides omarai nov. sp. has a short counter septum,
to irregular in shape and formed of septal lamella and few axial whereas in A. ivanovi it has the same length as the other major
tabellae. The wall and parts of the dissepimentarium are partly septa;
eroded. At the early immature stage, up to a diameter of 6 mm, the  The minor septa in A. ivanovi reach 2/3 of the length of the major
dissepimentarium is yet undeveloped (Fig. 12(B4)); it starts to septa, but in A. omarai nov. sp. they are less than 1/2 as long as
develop at a diameter of 8 mm (Fig. 11(B5)). At the adult stages, it is the major septa.
fairly wide, where better preserved consisting of up to 7 rows of
dissepiments. Interseptal dissepiments are angular or v-shaped. Differences of A. omarai nov. sp. with other Amygdalophylloides
Two to three rows of lonsdaleoid dissepiments occur at the outer species from northern Spain, the Carnic Alps, the Moscow Basin
dissepimentarium, but do not form a complete lonsdaleoid
dissepimentarium (Figs. 11(A6, C4), 12(B2)).
In longitudinal sections dissepiments are elongate and strongly
inclined towards the axis; they become nearly vertical near the
tabularium, where they are thickened and form a thin stereozone
(Figs. 11(A11, C3), 12(A6)). The wide tabularium occupies more
than 3/4 of the diameter of the corallite. It is divided into two parts:
the outer part consists of steeply inclined to almost vertical tabellae
(clinotabulae) followed by axial tabellae which are almost
horizontal or slightly inclined towards the columella; the inner
part consists of a thick columella of irregular outline which contains
some internal voids (Fig. 11(A11, B6). It occupies ca. 1/6 to 1/3 of the
diameter of the corallite and shows the median plate which is
continuous with the cardinal septum (Figs. 11(C3), 12(A6, B4, C4)).
Remarks: The individuals of this new species show remarkable
variability in size. Young specimens are common (Fig. 12).
Amygdalophylloides omarai nov. sp. differs from A. ivanovi
(Dobrolyubova, 1937) in the following aspects (Table 2):
Fig. 13. Relation between the number of major septa and coralla alar diameters for
Amygdalophylloides omarai nov. sp., Amygdalophyllum ivanovi Dobrolyubova, 1937,
 It is characterized by the presence of two to three rows of and A. ivanovi (Dobrolyubova, 1937) described by Herbig and Kuss (1988). Reduced
lonsdaleoid dissepiments which are not common in A. ivanovi diameter of the holotype in the most mature stage due to rejuvenation.
 M. Kora et al. / Geobios 52 (2019) 1–25
and Japan are compiled in Table 2. A differentiated stock of mostly
late Viséan to early Bashkirian Amygdalophylloides species includ-
ing a single late Moscovian species from Japan (Sugiyama and
Ezaki, 2002; see species list above) is not further discussed, as it has
a clearly separated stratigraphic range and belongs to a different
palaeobiogeographic region.
Amygdalophylloides ivanovi is most widespread in the Mosco-
vian Mediterranean coral Subprovince of Fedorowski (1978,
1981). Besides its representatives from the type region (Moscow
Basin; see compilation in Kossovaya, 2001) and Northern Spain
(Rodrı́guez, 1984b), Lophophylloides carnicum and Lophophylloides
profundum (Milne Edwards et Haime) described by Heritsch (1936)
from the upper Carboniferous and lower Permian of the Carnic Alps
were included in this species (Dobrolyubova, 1940). Both taxa
were also mentioned from northwestern Bosnia and Herzegovina
(part of former Yugoslavia; Heritsch, 1941). Very close in
dimensions is Amygdalophylloides(?) degrootae Rodrı́guez, 1984b,
originally based on a single specimen from De Groot (1963). How-
ever, the axial structure is less massive and more irregular with a
somewhat screwed central lamellae connected to the cardinal
septum, which led to a questionable assignment. The very small,
dense septa-bearing Amygdalophylloides anticuum Rodrı́guez et
Said, 2009 from southern Spain might be the oldest representative
and ancestor of the genus.
The geyerophyllid Geyeronaotia hispanica Rodrı́guez, 1984b
(from the Kasimovian of Cantabrian Mts., NW Spain) is superfi-
cially quite similar to the current species. Nevertheless, the very
complex, wide dissepimentarium with frequent naotic and lateral
dissepiments and the length of minor septa relative to major septa
in that species makes it different from the present species.
5. Discussion
5.1. Taphonomy
The studied corals of the Rod El Hamal Fm. have been reworked
to some degree after death, reoriented and concentrated by
physical processes. This resulted in partial or complete erosion of
calices, absence of apexes and partial erosion of walls in some
corallites. However, in general reworking appears to be minor as
demonstrated by the presence of completely preserved corals.
Hence they were still deposited close to their original habitats. The
corals are randomly distributed in the sediment, but mainly
oriented parallel to the bed surfaces, often in fossil pavements.
Most fracturing resulted from later diagenetic processes, probably
from compaction. This is evidenced because fractures do not
contain the sediment surrounding the corals, and also affect both
the coral skeleton and the cement filling the internal cavities.
5.2. Palaeoecology
The small- to large-sized, dissepimented solitary rugose corals
from the Rod El Hamal Fm., which moreover often bear complex
axial structures, are part of the Caniniid-Clisiophyllid fauna of Hill
(1938). More generalized to include the entire Palaeozoic era, it
was termed ‘‘the second gradational facies fauna’’ by Hill (1981). Its
members lived in relative shallow water, and occur mainly in
argillaceous limestones. This low diversity assemblage can be
compared with the near-shore inner shelf association RCA1 in
argillaceous thin-bedded limestones from the Upper Viséan of
Ireland (Somerville and Rodrı́guez, 2007) that was characterized
by the large-sized solitary genera Aulophyllum and Dibunophyllum.
In the late Moscovian of Egypt the niche is now occupied by
Bothrophyllum and Amygdalophylloides. Rejuvenescence, incrusta-
tions and borings are common in the studied fossil association.
19
Rejuvenescences. They follow on constrictions of coral growth
that caused a reduction in the diameter of the polyp, respectively
the corallite, and are commonly considered as responses to
environmental change (Scrutton, 1998; Rodrı́guez, 2004). Many
corals in the present assemblage suffered from repeated cons-
trictions and rejuvenations suggesting instability of the environ-
ment. They display mostly rejuvenescence of the lateral type
(Berkowski, 2012), in which the polyp withdrew only from a part of
the calicular periphery on one side of the calice, whereas the other
side remained unaffected (Fig. 14(A1–L2)). One specimen of
Amygdalophylloides omarai nov. sp. shows a marked axial
rejuvenescence (Fig. 14(M1–2)).
Berkowski (2012) attributed the reasons for the lateral
contraction and rejuvenescence to several external factors. The
most prominent are partial burial of the calicular part of the
corallite, toppling and resulting recumbent position of the
corallite, injury caused by other organisms, and competition
between coral and encrusting organisms. The resulting decrease in
diameter affected the dissepimentarium by the loss of peripheral
rows of dissepiments. The tabularium is commonly unaffected
during the rejuvenescence (Figs. 5(A11), 6(B)).
Encrustations. Encrusting and attached organisms are scarce
and do not cover large parts of the external walls of the Rugosa of
the Rod El Hamal Fm. Only three types of encrusters were
observed: cystoporate bryozoans (Fig. 12(A5)), small rugose corals,
and the tabulate coral Aulopora (Fig. 15(A1–C)).
Bioerosion. Borings are rare. Only one corallite of Bothrophyl-
lum suezensis nov. sp. (RAr 19) shows five rounded to oval cavities
of ca. 2 mm in largest diameter (Fig. 15(D2)). They occur at the
adult stage below the calyx in the dissepimentarium and in the
inner part of the tabularium between the septa. Due to the
stereoplasmatic thickening delineating the cavities, they most
probably were formed during the lifetime of the coral. Analogous
stereoplasmatic thickening is observed in rare examples from
upper Viséan–Serpukhovian corals from China (Wei Lin, pers.
comm.). Rodrı́guez (2004) attributed the scarcity of borings either
to a high rate of sedimentation or an unfavourable environment for
the boring organisms, or their rarity at this time.
Damage and skeletal repair. This feature is not frequent. Two
corals show re-cementation of fragments of the calicinal wall,
approximately into the earlier, primary position. In one of the
corallites, the calice has been broken and re-cemented during the
lifetime of the polyp. This is evidenced by the growth of an outer
epithecal wall on the broken part (Fig. 15(D1, D3)). In the second
corallite, probably the whole corallite was broken and parts were
lost, the remaining upper part of the corallite being re-cemented
slightly obliquely (Fig. 15(E1–E3)).
Attachment to the substrate. Rodrı́guez (2004) identified two
possibilities for solitary rugose corals to attach their coralla to the
substrate: radiciform processes (talons and rootlets) and/or
excrescences. Talons are small lateral extensions of the wall,
forming a flat or irregular base in the outline of the coral that fits
with irregularities of the hard substrate or hard substrate particles,
that were used as attachment sites during settlement. These are
common in specimens with well-preserved tips (Figs. 3(A), 10(B1–
B2, D1–D2), 14(F, L1–L2), 15(F1–F2)). In most of the specimens
studied here, attachment scars are found without further
modifications of the wall to form talons (Figs. 14(M1–M2),
15(G1–G2)).
Rootlets are long tubules that expand radially from the coral to
fix it in soft or firm substrates (Fig. 15(H–K)); they are not in direct
contact with hard particles in the substrate. Therefore, their
presence in rugose corals is a criterion for identification of soft
substrates (Rodrı́guez, 2004). Excrescences are irregular lateral
extensions, composed mainly of dense masses of stereoplasm. This
type is rare in the present coral fossil association. These lateral
20 M. Kora et al. / Geobios 52 (2019) 1–25

Fig. 14. Palaeoecology (Rejuvenescences). A1–I. External photographs of Bothrophyllum suezensis nov. sp. illustrating the lateral type constrictions and rejuvenescences
phenomena (arrows). J1–L2. External photographs of Bothrophyllum cylindricum nov. sp., specimens RAr 22 and 23, showing repeated lateral rejuvenescences and
constrictions only affecting the convex side of the corallite in J1 and L1; additionally two successive rejuvenescences in the calice affect the alar side of the corallite J2. M1,
M2. External photographs of Amygdalophylloides omarai nov. sp., specimen RAr 137, showing a marked axial rejuvenescence (arrow). Scale bars: 5 mm.

extensions affect the walls and the outer part of the septa
(Fig. 11(B4, B5).
Rugose and tabulate coral skeletons and brachiopod shells are
the most common skeletal objects that provide hard substrates for
larval attachment (Fig. 15(B1, B2)). Talons are mostly common
within the bothrophyllid corals from Rod El Hamal Fm., whereas
rootlets are more common within the geyerophyllids. The
occurrence of attachment scars in the earliest growth stages, i.e.,
near the apex but not along the skeleton during later growth,
suggests that the corallites were initially attached to some hard
 M. Kora et al. / Geobios 52 (2019) 1–25 21

Fig. 15. Palaeoecology (cont.): A1. External lateral photograph of Amygdalophylloides omarai nov. sp., specimen RAr 135, showing an encrusting cystoporate bryozoan in the
mature part of the corallite. A2. Enlargement of the encrustation marked in A1. B1, B2. External photograph of a calice fragment, encrusted by a small columellate rugose
coral; enlargement in B2. C. Broken corallite encrusted by the tabulate Aulopora. D1. External photograph of Bothrophyllum suezensis nov. sp., specimen RAr 19, showing a
partly broken calice; note the large attachment scar in the apex flank. D2. Transverse thin-section marked in D1, showing cavities in the dissepimentarium in the counter
(upper) side resulting from syn-vivo living soft-bodied organism. D3. Detail of the square marked in D1 to illustrate the growing epithecal wall on the broken part of the calice
(arrow). E1–E3. External photograph of B. suezensis nov. sp., specimen RAr 15, showing a lateral constriction in the lower part of the corallite (arrow) in E1 and broken and re-
cemented parts of the calice in E2. F1, F2. External photographs of the immature, broken part of B. suezensis nov. sp., specimen RAr 90, showing large attachment scar and
accompanying talon. G1, G2. External photograph of B. suezensis nov. sp., specimen RAr 35, showing a large attachment scar on the lateral side of the corallite; note the
position of the cardinal fossula (arrow) on the same side. H–K. External photographs of Amygdalophylloides omarai nov. sp., specimens RAr 131, 134, 140, and 165,
respectively, illustrating rootlets occurring from the immature to the mature parts of the corallites; note the small attachment scar near the apex in I1 and I2. Scale bars:
5 mm.
22 M. Kora et al. / Geobios 52 (2019) 1–25
substrate, subsequently became detached and lived free in a
recumbent mode of life. Attachment scars and talons are usually
proximolateral and situated on the cardinal side of the corallite
(Scrutton, 1998). However, in the present coral association they do
not systematically coincide with the convex or concave side. In
some corallites of Bothrophyllum suezensis nov. sp., the cardinal
quadrants and the attachment scars are located on the lateral side
(Figs. 5(D), 7(A)).
To sum up, the presence of talons and rootlets indicates that the
corals from the Rod El Hamal Fm. either grew upward rooted in a
soft sediment or were recumbent on it as secondary recliners.
According to Berkowski (2012), both types of settlement and
growth are typical for soft, argillaceous sediments with embedded
hard particles of mostly biogenic origin forming an attachment
base for sessile organisms.
Orientation of the cardinal fossula. Most corals in the present
assemblage possess a remarkably obvious cardinal fossula. It
clearly delineates the plane of bilateral symmetry, as the enclosed
cardinal septum differs from the other major septa in thickness,
length and/or continuity. According to Hill (1981), the position of
the fossula in the calice is commonly on the convex side of the
corallite and possibly related to the preference of the larva to settle
on its cardinal side. In several Ordovician genera, Elias (1984) and
Elias et al. (1988) indicated that the cardinal side is oriented in the
down-current direction and occurs on the convex side of the
corallite. In the studied corals, however, the position of the fossula
is variable with respect to the curvature of the corallite and not
always confined to the convex side of the corallite. Some of the
bothrophyllid corallites studied here show a cardinal fossula on the
convex side (Figs. 5(A1), 6(A4), 9(A4)), whereas others have their
cardinal fossula on the lateral side (Figs. 4(B2), 5(C–F), 7(A),
15(G1)). It has to be noted that the attachment scars and the
cardinal fossulae are found on the same side of the corallite.
Therefore, the assumption of the preferential larval settlement and
substrate attachment on the cardinal side is probably correct, in
spite of the changing position of the cardinal septum/cardinal
fossula from the convex to the lateral sides.
5.3. Depositional environment
The coral-bearing rocks are bioclastic rudstone with peloidal
grainstone matrix. The bioclasts are often fragmented. They are
embedded in densely packed grainstone, consisting of peloids and
silt-sized detrital quartz grains. The peloids are the main
allochems. They were derived from reworking of sandy carbonate
mud. Relicts of that mud are still preserved inside the fossil
cavities. Bioclasts include large-sized rugose corals, auloporid
tabulate corals, punctate and inpunctate brachiopod shells and
spines, gastropod and bivalve shells, crinoid and echinoid
fragments, calcareous foraminifera and calcareous algae, trilobite
shell fragments, and bryozoans (Fig. 16(A, B)). Bioturbation occurs.
In some cases, the rudstone grades into calcareous sandstone
(Fig. 16(C–E)). The carbonate microfacies indicates deposition
adjacent to a low terrigenous hinterland in littoral and inner neritic
zones, which were inhabited by a diverse, predominantly filter-
feeding faunal association. The feeding type indicates an elevated
availability of nutrients, which primarily originated from the
hinterland.
According to microfacies interpretation, as well as coral
morphology and their position in the sediment, they lived in a
soft muddy substrate in inner ramp environments. They are seen in
the greenish marl packages of the studied succession. After
winnowing of the mud by tempestites and concurrent reworking
and concentration of the previously randomly settling organisms,
the biota were redeposited in thin fossiliferous limestone beds
(Fig. 16(F)), mostly without much transportation or fragmentation
of the larger fossils.
5.4. Palaeobiogeography
Except for Egypt, post-Bashkirian corals are not known in the
extended Carboniferous epicontinental Saharan basins along the
Gondwanan margin in northern Africa (Semenoff-Tian-Chansky,
1985). This causes difficulties to elucidate the palaeobiogeographic
affinities of the corals described herein.
The Bashkirian corals from the Béchar Basin in Algeria
(Semenoff-Tian-Chansky, 1974) and further Algerian basins
evidence a faunal turnover at the mid-Carboniferous boundary
(Semenoff-Tian-Chansky, 1985; Aretz, 2011), but clearly differ
from the Egyptian corals described herein. They are part of the
Mediterranean Province as defined by Fedorowski (1978, 1981)
and can be grouped in a southern subprovince (Fedorowski, 1981),
more precisely a West-Mediterranean Subprovince (Dubatolov
and Vassiljuk, 1981) which includes Northern Africa, Spain
(Cantabrian Mts.), the former Yugoslavia, Donets Basin, and Asia
Minor. According to Semenoff-Tian-Chansky (1985), the Bashki-
rian corals from the Béchar Basin show more affinities to the
Donets and the Russian platform (Voronezh anticline) than to the
Cantabrian Mts. In addition, Aretz (2011) advocated affinities to
the Donets, but did not rule out connections with western North
America due to the problematic affinities of Bothrophyllum proetum
Semenoff-Tian-Chansky, 1974, as discussed by Semenoff-Tian-
Chansky and Guillaume (1994).
In contrast, the Bashkirian corals from the northern flank of the
Tindouf Basin differ, especially from the Béchar Basin, and are
regarded as late Mississippian survivors (Somerville et al., 2013;
Cózar et al., 2014) which inhabited a partly isolated basin in the
western North Gondwana. A strongly similar assemblage from
Adarouch (central Morocco) additionally yielded the first Cysto-
lonsdaleia Fomichev, 1953 from Northern Africa, which again
indicates communications with the Donets and Moscow basins
(Rodrı́guez et al., 2015).
The general palaeobiogeographic coral patterns of the Bashki-
rian continued through the Moscovian Stage. According to
Fedorowski (1978, 1981), the southern subprovince of the
Mediterranean Province again included the Cantabrian Mts.,
former Yugoslavia, Moscow and Donets basins; now are also
included the Carnic Alps. An abundance and diversity peak was
reached during the late Moscovian (Myachkovian). A very similar
grouping was proposed by Dubatolov and Vassiljuk (1981), who
placed the Donets outside the western Mediterranean Subprovince
into a central Mediterranean Subprovince, and attributed the
Moscow Basin to a separate Uralo-Arctic Province. However, these
authors also stressed a late Moscovian phase of diversification and
prosperity.
Besides the late Moscovian coral fauna from the upper part of
the Rod El Hamal Fm. described herein, only one other Egyptian
rugose coral assemblage consisting of small cornute taxa –
including Actinophrentis, Bradyphyllum, Lytvolasma, Rotiphyllum,
Allotropiophyllum, Allotropiochisma, Meniscophyllum, Amplexiza-
phrentis, Verbeekiella, Pseudowannerophyllum, and Assimulia – has
been recorded (Kora and Mansour, 1991). This fauna is from dark
grey shales in the lower part of the Aheimer Fm. from the eastern
slopes of the Northern Galala plateau (Fig. 1). This formation,
which is missing in the Wadi Araba area, was considered to be
slightly younger (Westphalian D–Stephanian C = late Moscovian–
Gzhelian; Kora and Mansour, 1991). At a generic level, the corals of
the Rod El Hamal Fm. and from the lower Aheimer Fm. show strong
similarities to the Donets Basin, the Cantabrian Mts. and North
America, thus indicating marine connections between these
 M. Kora et al. / Geobios 52 (2019) 1–25 23

Fig. 16. Depositional environments. A, B. Microphotographs of bioclastic rudstone with densely packed peloidal grainstone matrix. Rugose coral (RC), abundant calcareous
algae (A) comparable to Paraepimastopora irregularis, brachiopods (Br), bivalves (B), gastropods (G), and unidentified fossil hash; small micritic intraclasts associated with the
peloid. C–E. Microphotographs of sandy limestone/calcareous sandstone cemented by sparry calcite; abundant fine-grained, relatively well-sorted, angular to sub-rounded,
quartz grains, bivalves (B), brachiopods (Br), ostracodes (Os), foraminifers (Fr), crinoids (Cr), echinoderm plates (Ec), and the tabulate coral Aulopora in picture E. Bioclasts in
part strongly fragmented. F. photograph of a rock slab showing a bioclastic rudstone with abundant echinoid spines, round and pentagonal crinoid ossicles and shells. Scale
bars: 2.5 mm (A, B), 1 mm (C–E), 1.5 cm (F).

regions. Besides a postulated but unknown direct East–West
seaway between northern Spain and North America (Fedorowski,
1978; Rodrı́guez et al., 1986; Garcı́a-Bellido and Rodrı́guez, 2005:
‘‘Midcontinent–Iberian-Seaway’’) along the suturing African and
Laurussian plates, only a connection via the Urals–Arctic seaway
would have been possible. It has to be stressed that Rodrı́guez
(1984a) earlier claimed the disappearance of American faunal
influences in the Cantabrian Mts. during the early Bashkirian and
gradual isolation during the late Moscovian, though faunal
elements still immigrated from eastern basins. Concerning
Northern Africa, Legrand-Blain et al. (1989) stated no more marine
connections with Northern America from the Bashkirian onwards,
and in spite of their confusing fig. 5, they noted that all Moscovian
calcareous algae, foraminifers, brachiopods and corals are typical
Palaeotethyan dwellers. Finally, palaeogeographic maps show the
sutured northwestern Africa and eastern North America (Torsvik
and Cocks, 2017) and a West–East marine gateway can be
definitely ruled out after the early Bashkirian.
In fact, the strong affinity of the Egyptian and North American
corals assumed by Kora and Mansour (1991) appears to be
problematic as it is based on mostly very simple, not facies
dependent, ubiquitous cyathaxonid-type coral genera which might
include homeomorphs. This also might be seen by genera listed
from these authors, that were assumed to occur in Egypt and
Northern America, but mostly not in the palaeogeographic closer
Donets Basin.
The Bothrophyllidae and the geyerophyllid Amygdalophylloides
from the Rod El Hamal Fm. clearly belong to the southern
subprovince of the Mediterranean Province of Fedorowski
(1981). Moscovian Bothrophyllum and allied taxa (see Fedorowski,
2016) are known from the Cantabrian Mts., the Carnic Alps, former
Yugoslavia, Moscow Basin, but also from the adjoining Uralian-
Arctic province. Likewise, the geyerophyllid genus Amygdalophyl-
loides is recorded from the late Moscovian of the Cantabrian Mts.
and from the late Moscovian and Kasimovian of the Moscow Basin.
In general, it is a Palaeotethyan genus known between Spain and
24 M. Kora et al. / Geobios 52 (2019) 1–25
Japan and recorded from the upper Viséan to the lowermost
Permian; it reached its maximum distribution during the
Moscovian (Minato and Kato, 1975; Rodrı́guez, 1985).
Opposed to the mentioned occurrences, the corals from the Rod
El Hamal Fm. are the only large Moscovian caninoid corals known
from North Africa and the Middle East, i.e., along the northern
margins of Gondwana. According to our knowledge, descriptions of
age-equivalent corals from the displaced North Gondwana
terranes in the Taurides (Turkey) or in the Zagros Belt (Iran) are
missing, which would close an important gap in knowledge. The
endemism of the fauna might be an artefact related to that missing
knowledge. Alternatively, it might be related to restricted facies in
an embayment in the present-day Gulf of Suez region shown on the
palaeogeographic map of Torsvik and Cocks (2017: fig. 9.3). In that
semi-enclosed embayment, the specially adapted, low diversity
coral fauna of the Rod El Hamal Fm. developed in a carbonate-poor
mixed siliciclastic-carbonate environment.
6. Conclusions
The coral-bearing limestones of the upper part of the Rod El
Hamal Fm. from the Wadi Araba area, western side of the Gulf of
Suez (Egypt) yielded a very low diversity, endemic association of
solitary dissepimented rugose corals. This resulted in the
introduction of three new species, in addition to some unidentified
bothrophyllid species and genera. A late Moscovian age is assigned
according to the associated brachiopods (El-Nakkady, 1958), the
currently studied bryozoan fauna and the Epimastoporacean
calcareous algae. The coral association also points itself to the
late Moscovian, as a general abundance and diversity peak is
reached during that time slice, where Bothrophyllids and
Amygdalophylloides are common.
Colonial Rugosa are missing, and except for small-scale
auloporid encrustations further tabulate corals (‘‘Syringopora’’:
Abdallah and El Adindani, 1965; El Adindani, 1968) are extremely
rare. The facies of the upper unit of the Rod El Hamal Fm. records a
depositional setting in turbid waters of an inner ramp, close to a
low terrigenous hinterland, probably in a gulf-like embayment
intruding from the Palaeotethys toward the South across the
present-day Carboniferous outcrops of the Gulf of Suez region. The
inner ramp was characterized by primarily muddy sediments
inhabited by a rich association of filter feeders. Episodical
reworking by tempestites caused the formation of thin bioclastic
limestone beds. Thus, a mixed siliciclastic-carbonate succession
was formed. Anatomical characters of the corals confirm this
interpretation. Rootlets and ceratoid to trochoid growth forms
prove life on a soft substrate; repeated rejuvenation indicates
interrupted growth in unstable environments. Erosional features
prove certain transport of the corals.
The endemic corals from the Rod El Hamal Fm. are the only
example of a Moscovian dissepimented coral fauna at the southern
margin of the Palaeotethys in North Africa and the Middle East,
along the northern margin of Gondwana. Palaeobiogeographic
affinities remain vague, but the big caninoids indicate similarities
to northwestern Spain (Cantabrian Mts.), the Donets and Moscow
basins, and thus, a general attribution to the Palaeotethys realm.
Acknowledgements
The authors wish to express their sincere appreciation to Prof.
Dr. Jerzy Fedorowski (Adam Mickiewicz University, Poznań,
Poland) for helpful discussions during a visit of the corresponding
author in Poznań. They also thank Dr. Julien Denayer (University of
Liège, Belgium) for discussions and valuable comments on a
previous version of the manuscript. The manuscript benefited from
constructive reviews of Prof. Dr. Ian Somerville (University College
Dublin, Ireland), Dr. Olga Kossovaya (Kazan Federal University,
Russia) and Dr. Markus Aretz (Paul Sabatier University, Toulouse,
France), and also from helpful comments of the editor in Chief Dr.
Gilles Escarguel and the associate-editor Dr. Bertrand Lefebvre. We
also acknowledge Dr. Wei Lin (Nanjing, Chinese Academy of
Science) for further comments on the corals, Dr. Andrej Ernst
(University of Hamburg, Germany) for determination of bryozoans,
and Dr. Daniel Vachard (University of Lille, France) for help with
the calcareous algae. Deepest gratitude also goes to Hanna
Cieszynski and Kathrin Jung (University of Cologne, Germany)
for their technical help. The authors express their gratitude to Prof.
Dr. Hesham Sallam and Dr. Hytham El-Atfy (Mansoura University,
Egypt) for their help during fieldworks. The corresponding author
acknowledges the Egyptian Ministry of Higher Education and
Scientific Research for providing financial support for a scholarship
at the University of Cologne, Germany.
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