late carboniferous solitary rugose corals from the western side of the gulf of suez, egypt

N. Jb. Geol. Palaont. Mh. Stuttgart, Oktober 1991

Late Carboniferous solitary rugose corals from thewestern side of the Gulf of Suez, Egypt

By Mahmoud Kora and Yasser Mansour, Mansoura

With 6 figures and 1 table in the text

KORA, M. & MANSOUR, Y. (1991): Late Carboniferous solitary rugose corals from the

western side of the Gulf of Suez, Egypt. - N. Jb. Geol. Palaont., Mh. 1991 (10): 597 - 616;Stuttgart.

Abstract: Cornute rugose corals from the lower member of the Aheimer Formation in theeastern cliffs of the Northern Galala indicate a late Westphalian to Stephanian age and ashallow, quiet normal marine environment for the coral-bearing horizon. Striking faunistic

similarities suggest that connections betWeen North Africa, Spain and the Donetz Basinwith North America might have continued to the end of the Carboniferous. All indentifiedspecies belong to the order Stauriida; seventeen of them are new for Egypt.

Zusammenfassung: Eine reiche Hornkorallen-Fauna aus Tonschiefern der unteren Ahei-mer-Formation vom Ostrand des Nordlichen Galala spricht fur spates Westfal bis Stephanund einem ruhigen Flachmeer-Biotop. Enge faunistische Beziehungen lassen vermuten,daB die Wanderwege zwischen Nordafrika, Spanien und dem Donetzbecken, aber auchnach Nordamerika, bis zum Ende des Karbons bestehenblieben. Aile bestimmten Artengehoren zur Ordnung Stauriida, wobei Antiphylliden und Hapsiphylliden dominieren undLophotidiien und Verbeekielliden seltener sind. Siebzehn Arten sind neu fur Agypten.

Introduction

Works dealing with the paleontology and stratigraphic distribution of the

Carboniferous corals in Egypt are very few and the coral fauna is usually

described among fossil assemblages including other taxa. Few rugose corals

from the Gulf of Suez and Sinai had long been loosely identified as Zaphrenthis

RAFFINESQUE & CLIFFORD or Zaphrentoides STUCKENBERG (WALTER 1890,

KLEBELSBERG 1911 and BALL 1916). Also, Late Carboniferous rugose corals

including species provisionally allocated to Lophophyllidium GRABAU,

Caninia MICHELIN, Clisiophyllum DANA and Cyathaxonia MICHELIN were

identified from the Wadi Araba area in the Eastern Desert (ABDALLAH & EL

ADINDANI 1965 and EL ADINDANI 1968). All these identifications were based

only on external features; the more important and diagnostic internal charac-

ters were neglected. '

0028-3630/91/1991-0597 $ 4.50@

1991 E. Schwe;zerbm'sche Verlagsbuchhandlung, D-7000 Stuttgart 1

598 M. Kora and Y. Mansour

The only works on the Egyptian Carboniferous corals based on thinsections study were made by OMARA (1967 & 1971), KORA &Jux (1986) andKORA (1989) describing an Early Carboniferous fauna from Sinai, and byHERBIG & Kuss (1988) dealing with Late Carboniferous rugose corals from theNorthern Galala. In fact, the few fossils listed by ABDALLAH& EL ADINDANI(1965) and AWAD & SAID (1966) from the Abu Darag and Wadi Aheimersuccessions on the western side of the Gulf of Suez need to be reviewed indetail. For example, a study based only on five specimens (HERBIG & Kuss1988) resulted in the identification of three additional solitary rugose coralsnamely; Amygdallophylloides ivanovi (DOBROLYUBOVA), Bothrophyllumpseudoconicum DOBROLYUBOVAand Pseudozaphrentoides ex gr. juddi (THOM-SON). Similarly, the detailed study of more than one hundred and twentycornute solitary rugose coral specimens, thin sectioned in the present work,proved to be essential and certainly will bring additional data on the paleontol-ogy, paleoecology and paleobiogeography of the investigated Upper PaleozoicsuccessIOn.

The Paleozoic succession exposed on the western side of the Gulf of Suezranges in thickness from about 60 m in the area around Abu Darag Lighthouse

32 30'E150 7'.".':.m ~:=:~

0~

t.D

c:~12 «.~

~ 0 .".."...~ :; ~.:: § ~9 ....

0lL

>- ...

~~ II ~X:~

y~lIow sandston~

with siltstone

Interbeds<"v

<.-<:-

crinoidal dolostone

intercalating brown

sandst one

300N

II. .c. grey shales with

~~« .1 ::~-:. crinoids.brachiopods~:; :c~ , COR 5u 2 E ~~~- I!

and AL

J'v

<"~

4km.

~ Qu.aternary

EI::9 loweof Tertiary32°2(1

E3 Upper Cretaceous ~ Jurassic

E:] Lower Cretaceous E;J Upper Permian

2 20'[[I] Lower Permian

~ Upper Carboniferous

32°l,()'

Fig. 1. Geological sketch map of the study area on the western side of the Gulf of Suez(modified from ABDALLAH&EL ADINDANI1965), and the lithostratigraphy of the Aheimer

Formation.

Late Carboniferous solitary rugose corals 599

to more than 200 m near Ain Sukhna in the north. They form a narrow strip ofdark coloured sandstones alternating with a number of fossil-bearing shale,marl and dolostone beds. In this sequence, Upper Carboniferous, LowerPermian and Upper Permian exposures could be differentiated (Fig. 1). Eachoutcrop is surrounded mainly by Cretaceous - Early Tertiary strata in a faultedregion. The studied succession in the eastern cliffs of the Northern Galala,around Bir Aheimer, is about 162 m thick. It was assigned to the UpperCarboniferous- Permian Aheimer Formation of ABDALLAH& EL ADINDANI(1965). It could be subdivided into three members as briefly described in Fig. 1.The rugose coral fauna described here was collected from a 13.1 m thick darkgrey shale horizon at the base of the lower member. All studied materials aredeposited at the Geology Department, Faculty of Science, Mansoura Univer-sity, Mansoura, Egypt.

Taxonomic notes

The terminology and systematic classification of the Subclass RugosaMILNE-EDWARDS& HAIME outlined in the Treatise on Invertebrate Paleontol-ogy (HILL 1981) is adopted. The illustrations given by POTY (1981) for theinternal structural elements, which were subsequently used by BOLL (1983)and RODRIGUEZ (1984a), have been found useful. The present work has alsogained much from the studies of SANDO&BAMBER(1985), FEDOROWSKI(1987),WILSON(1990) and RODRIGUEZ& KULLMANN(1990). All the studied corals areincluded in Order Stauriida VERRILL, of which thirteen genera belonging tofive families are identified.

Family Antiphyllidae lUNA 1970

Six species belonging to four genera are encountered (Fig. 2). The genusActi/nophrentis, introduced by FOMICHEV (1953), remained invalid untilIVANOVSKIY(1967) designated A. donetziana FOMICHEV as its type species.Moreover, Actinophrentis IVANOVSKIYwas synonymized by WEYER (1975)with Rotiphyllum HUDSON. This synonymy is not accepted here, since theholotype of A. donetziana retains a pinnate arrangement of septa up to itstopmost part (FEDOROWSKI 1987), in contrast to Rotiphyllum rushianum(VAUGHAN)and other unambiguous rotiphylla.

Bradyphyllum d. bellicostatum (Fig. 2b) differs from B. bellicostatumGRABAUin that the major septa are not withdrawn from the axis, but tend to beconjoint due to rhopaloid thickening. In this respect, it approaches the septalarrangement in B. oppositum FOMICHEV described by RODRIGUEZ (1984a)from the Upper Carboniferous of Spain. Moreover, a single specimen ofActinophrentis nikitovkensis nana FOMICHEVfrom the Upper Gzelian of theDonetz Basin was included by FEDOROWSKI(1987) with his Lytvolasma aucta.


Also, a great similarity to the encountered L. aucta FEDOROWSKI(Fig.2c) isseen in some specimens of Allotropiophyllum irregulare FOMICHEV,which aregeologically older "Late Moscovian" and show a larger number of septa.

The present work agrees with many authors (e. g. DE GROOT 1963, WEYER1975 and HILL 1981) who included Monophyllum FOMICHEV in synonymywith Rotiphyllum HUDSON. Meanwhile, FEDOROWSKI(1987) concluded thatthe combination of characters shown by Monophyllum sokolovi FOMICHEVpermits consideration of Monophyllum as a separate genus very close toRotiphyllum and Actinophrentis. However, neither one of the three importantdifferences between Rotiphyllum and Monophyllum (cited by FEDOROWSKI1987: 67-68) is observed in the present material, and all the studied specimens(Fig. 2d-g) satisfy the diagnosis of the genus Rotiphyllum HUDSON and itssynonym Monophyllum FOMICHEVas given by HILL (1981).

Family Hapsiphyllidae GRABAU1928

The Hapsiphyllidae is one of the most controversial families in the rugosecoral literature. The solution of HILL (1981) seems unsatisfactory to manyworkers, because it places together some unrelated genera. Nevertheless, thepresent study follows HILL'S (1981) classification until the taxonomy of theHapsiphyllidae is clarified and more taxa are thoroughly investigated. Fivespecies belonging to four genera could be identified.

Allotropiophyllum sp. (Fig. 3a) differs from A. sinense (GRABAU),from theLower Permian of China, by the absence of scattered hollow spines, probablydue to abrasion; in having less developed minor septa; in the absence of theaxial space; and in having more major septa. In this respect, it approachesA. sniatkovi FOMICHEVas described from the Upper Carboniferous of Spain(RODRIGUEZ 1984a) which is smaller, has the cardinal septUm not distinctly

Fig. 2. Antiphyllid rugose corals from the Aheimer Formation, Gulf of Suez. Scale bar =1 mm.

a: Actinophrentis donetziana FOMICHEV1953, RAH134; a1,2: transverse and longitudinalsections.

b: Bradyphyl/um d. bel/icostatum GRABAU 1928. RAH142; b1,2: transverse and longitud-inal sections.

c: Lytvolasma aucta FEDOROWSKI 1987. RAH158; c1,2: transverse and longitudinalsections.

d: Rotiphyl/um aequabile DE GROOT 1963. RAH14; dl,2: transverse and longitudinalsections.

e, f: Rotiphyl/um sokolovi (FOMICHEV).RAH143; el,2: transverse and longitudinal sec-tions. RAH146; f: transverse section.

g: Rotiphyl/um exile DE GROOT 1963. RAH43; gl,2: transverse and longitudinal sections.


shortened and the septal arrangement around cardinal fossula somewhatdifferent from the present material.

The studied specimen of Allotropiochisma (Alligia) d. flabellumFEDOROWSKI(Fig. 3b) has in general a smaller size and a smaller number ofmajor septa than the type material described by FEDOROWSKI(1987) from theWolfcampian of Texas. It is partly similar to ?£uryphyllum robustumFEDOROWSKI,which differs by having more robust major septa, less welldeveloped alar fossulae and a stronger stereocolumn. The specimen assignedhere to the genus Meniscophyllum SIMPSON(Fig. 3c) is close to Meniscophyl-lum sp. described by RODRIGUEZ (1984a: Fig. 94, pI. VI, figs. 5,6) from theUpper Carboniferous of East Asturias, Spain, which has 22 major septa at adiameter of 10 mm in the adult stage.

On the other hand, the sectioned specimen of Amplexizaphrentis paral-le/oides (DE GROOT) is partly compressed (Fig. 3d) so that the cardinal fossulais not typically parallel-sided as the type material described from the Westpha-lian D of Spain (DE GROOT 1963). The encountered coral shows someresemblance with several sections assigned to Zaphrentites parallelus(CARRUTHERS)from the Lower Carboniferous of South China (KULLMANN&LIAO 1985: 138). The present material, however, differs in the stronger dilationof major septa, and in the greater length of the counter septum. Also, thespecimen (Fig. 3e) assigned here to A. clithria (DE GROOT) has more majorsepta than the type material described from the Westphalian D of Spain (DEGROOT 1963: 42), having 22 major septa at a diameter of 8-10 mm.

Family Verbeekiellidae SCHOUPPE& STACUL1955

A specimen assigned to the genus Verbeekiella PENECKE(Fig. 3f) exhibits afabric that is mostly destroyed due to weathering and poor preservation, sothat specific identification is difficult. HILL (1981), althoug1¥>he left Pseudo-

Fig. 3. Hapsiphyllid and verbeekiellid rugose corals from the Aheimer Formation. Scalebar = 1 mm

a: Allotropiophyllum sp. RAH156; al,2: transverse and longitudinal sections.

b: Allotropiochisma (Alligia) d. flabellum FEDOROWSKI1987. RAH138; bl,2: transverseand longitudinal sections.

c: Meniscophyllum sp. RAH145; transverse section.

d: Amplexizaphrentis paralleloides (DE GROOT). RAH42; dl,2: transverse and longitudi-nal sections.

e: Amplexizaphrentis clithria (DE GROOT). RAH136; el,2: transverse and longitudinalsections.

f: Verbeekiella sp. RAHI64; fl,2: transverse and longitudinal sections.g: Pseudowannerophyllum differens FLUGEL 1975. RAHI6; gl,2: transverse and longitud-

inal sections.


wannerophyllum FLUGELto the family Verbeekiellidae SCHOUPPE& STACULinwhich it has been originally included by FLUGEL(1975), remarked that "densepacking of elements in axial structure may indicate relationship to Lophophyl-lidae". This has led FEDOROWSKI(1987) to introduce an emended diagnosis forthis coral which places it with the family Lophophyllidiidae MOORE & JEF-FORDS. In fact, the features described by FEDOROWSKI (1987: 137, 138),including the loose columella starting its development from the counterseptum, and the secondary lamellae appearing by means of peripheral split, arenot documented in the studied specimen of Pseudowannerophyllum differensFLUGEL (Fig. 3g). Therefore, the genus is described here under its originalfamily and not in the Lophophyllidiidae.

Family Bothr'ophyllidae FOMICHEV 1953

The transverse section of Bothrophyllum pseudoconicum DOBROLYUBOVA(Fig. 4a) exhibits some similarities in form and septal arrangement to some

sections assigned to Pseudozaphrentoides rabanaliensis DE GROOT from the

Westphalian D of Northern Spain. However, the encountered specimen is

more or less identical to B. pseudoconicum DOBROLYUBOVA described in detailby HERBIG & Kuss (1988) from Bir Qiseib in the study area. Both specimens

are close to similar material known from the Westphalian D and Stephanian of

the Cantabrian Mountains in Spain (DE GROOT 1963 and RODRIGUEZ 1984a).

Family Lophotichiidae WEYER 1972

HILL (1981) considered Lophotichium MOORE & JEFFORDS a member of

Cyathaxoniidae MILNE-EDWARDS & HAIME and included Lophotichiinae in

synonymy with the latter. The present authors agree with FEDOROWSKY (1987)

who pointed out that the relation of Lophotichium and Cyathaxonia is not

closer than on the family level. Consequently, FEDOROWSKI (1987) has trans-

Fig. 4. Bothrophyllid and lophotichiid rugose corals from the Aheimer Formation. Scale

bar = 1 mm.

a: Bothrophyllum pseudoconicum DOBROLYUBOVA 1937. RAH15; a1,2; transverse andlongitudinal sections.

b: Paraduplophyllum (Paraduplophyllum) oppositum FEDOROWSKI 1987. RAH169; b1,2:transverse and longitudinal sections.

c: Paraduplophyllum (P.) multiplicatum FEDOROWSKI 1987. RAH140; cl,2: transverse andlongitudinal sections.

d: Paraduplophyllum (Vacoea) aff. nealranchense FEDOROWSKI 1987. RAH127; d1,2:transverse and longitudinal sections.

e, f, g: Assimulia (Ericina) fracta FEDOROWSKI 1987. RAH159; e: transverse section.

RAH120; £1,2: transverse and longitudinal sections. RAH147; g: transverse section.


ferred the genus Paraduplophyllum Wu & ZHOU from the family Metriophyl-lidae HILL to Lophotichiidae WEYER.The strongly biform tabularium presentin Paraduplophyllum and lacking in Metriophyllum MILNE-EDWARDS& HAIMEspeaks adequately against its original placement. On the other hand,Duplophyllum sp. described from the Middle and Upper Carboniferous ofChina (Guo 1983: pI. II, fig. 3) appears fairly similar to the studied Paradu-plophyllum (P.) oppositum FEDOROWSKI (Fig. 4b). Meanwhile, the speciesidentified here as Paraduplophyllum (Vacoea) aff. nealranchense (Fig. 4d) islarger than the type material of FEDOROWSKI(1987). Actually not all thecharacteristic features of the species are clear in the sections made, so thatspecific assignment is only tentative.

In spite of the fact that all these corals were derived from the so calledLophophyllidium-bearing shale series (ABDALLAH& EL ADINDANI 1965), nosingle specimen could be assigned to the genus Lophophyllidium GRABAU.It isprobable that the preliminary allocation for some of these corals to that genuswas not accurate, since the identification was based only on morphologicalfeatures (d. EL ADINDANI 1968). Even if Lophophyllidium GRABAUdoes occurin this horizon, it should be very rare. Therefore, it is suggested here to replacethe term Lophophyllidium-bearing shale by the informal name "lowermember" of the Aheimer Formation. Moreover, these corals and their hostrocks are neither homotaxial nor coeval with those described from the car-bonates of the Urn Bogma Formation, and the correlation recently made bysome workers could not be supported.

Age assignment

The age ranges of the identified corals compiled from their distribution inthe well known Carboniferous-Permian basins in N. America, W. Europe,NW. Africa, the U.S.S.R., SE Asia, etc. are shown in Fig. 5. The stratigraphicranges given indicate that the majority of these fossils have a Late Carbonifer-ous affinity. The presence of Rotiphyllum sokolovi (FOMICHEV) and Both-TOphylium pseudoconicum DOBROLYUBOVA(unknown below Westphalian D)indicates that the lower limit of the age of this coral-bearing horizon is at thebase of Westphalian D or Late Moscovian. Meanwhile, this lower member ofthe Aheimer Formation is characterized by the presence of corals belonging tothe genera Paraduplophyllum Wu & ZHOU, Lytvolasma SOSHKINA, Allot-ropiochisma FEDOROWSKI,Verbeekiella PENECKEand Assimulia FEDOROWSKIwhich are known to cross the Carboniferous/Permian boundary (Fig. 5).However, these corals are associated with characteristic Pennsylvanianbrachiopods like Anthracospirifer sp. and Rugosochonetes califomicus WAT-KINS.This suggests that the upper limit for the age of this interval is at the topof Stephanian C or the Gzelian. The Late Westphalian to Stephanian agesuggested here for the lower member of Aheimer Formation is in agreement

: Pennsvlvanian L. PermiarI~

Desma ines. Missour. Virgilian Wolfcampian N. Am~rican EpochsI=-I".

. IIW~stphalian St~phanian Epochs. Rotlo~g~nd~sl W. Europun

."'1n 10 !l ~I

'"In I

G> ~(flI lit

"N lit X"

I Moscovi..n !. !.~Koiislmovian 3 U.S.S.R. Epochs

"ij"

~I ~::JIi"::J

I I

IMeniscophyllum, Sl?

Pseudowannerop,hyll um differensI

Bradyphyllum ct. bellicostatumI I- Rotiphyllum sokoloviI I- R. aequabile II I- R. exile I1- Amplexizaphrenfis clithriaI- A. parallel oidesI I

Bothrophyllum pseudoconicumI I

Actinophrentis donetzianaI I Paraduplophyllum

I I(P.) multiplicatumP.(A) oppositum

I I A(V.) aft. nealranchel15,I

Lytvolasma auctaI I

Allotropiochisma (A)

I 1 flabellumAssimulia (E.J fracta

I I

I IVerbeekiella sp,

Allotropiophyllum 51?

I I


Fig. 5. Stratigraphic ages of the identified corals from the western side of the Gulf of Suez.

with the Late Carboniferous age assignments recently given by HERBIG & Kuss

(1988) and ABD ELSHAFY & ABD ELAZEAM (1990).

Paleoecological aspects

The richly fossiliferous basal part of the lower member of Aheimer Forma-tion is composed of calcareous silty shale facies containing a varied fauna ofsolitary rugose corals, brachiopods, bryozoans, bivalves together with manycrinoids and rare trilobites. The association of faunas indicates a shallow,normal marine environment. The shales were accumulated under low energyconditions, as fines were deposited mainly from suspension. This is also


evidenced by the presence of delicate fenestrate bryozoans as well as thepreservation of most crinoids in growth position.

Thus, the studied corals correspond to the first gradational facies faunarecognized by HILL (1981) which is a fauna of small solitary nondissepimentedgenera, not of great diversity but accompanied by small brachiopods. It isregarded as adapted to the muddier, darker, quieter and therefore deeper partsof epicontinental or shelf sea floors. Deposition must have been slow lest thecorals would have smothered, since they cannot survive where large amountsof sediment are shifted over the bottom by waves or currents (HILL 1981);sediment in suspension is not inhibiting.

Similarly, SANDO(1980) and SANDO& BAMBER(1985) grouped the occur-rences of coral taxa into deep-water and shallow-water facies and derived ashallow-water habitat index (SWHI) that represented the percentage of occur-rences of each taxon in shallow water lithofacies. In the present study, theforms identified as Rotiphyllum are regarded as deep water biofacies (SWHI

= 33) whereas the forms identified as Amplexizaphrentis are regarded asshallow water biofacies (SWHI = 74). Among the lithofacies of the coralhabitats suggested by SANDO (1980), the shallow water terrigenous faciesrepresented by dark and light-coloured fissile to medium bedded mudstone,siltstone and silty micrite is the closest. The environment was that of shallowpoorly circulated water in basins marginal to shore line with a depth varyingfrom 0-100 m.

The biogenic structures "trace fossils" frequently encountered in the upperpart of the lower member of the Aheimer Formation include mostly simplevertical and deep burrows in a silty shale facies. These indicate a very shallow-marginal "intertidal" environment where ecological conditions were notfavourable for epibenthic organisms and they had to go deeper into substratefor protection. So, a sheltered shallow marine environment changed to interti-dal conditions during the deposition of the upper part of the lower member ofAheimer Formation.

At the beginning of the Moscovian, a transgression brought an epiconti-nental sea onto the northern and eastern Sahara including Wadi Araba, AbuDurba, Abu Darag and Wadi Aheimer areas around the Gulf of Suez in Egypt.The Carboniferous sea invaded Libya and extended far south to Chad andNiger passing to a more continental facies Qux 1986). That sea progressivelyleft North Africa, by the infilling of the Saharan basins from the south due tonortheastward tilting of the continent and from west to east, remaining only inthe Permo-Carboniferous troughs (LEGRAND-BLAINet al. 1987).

Since most of the studied rocks of the lower member of Aheimer Formationwere deposited mainly in an intertidal to shallow subtidal environment, it isconcluded that the study area remained, throughout most of the Late Car-boniferous time, near the shore of that sea with only few open marine


encroachments. At the same time, regional uplifting took place in other areas inthe Near East.

Paleo biogeographical implications

A paleozoogeographical subdivision of Carboniferous paleobasins by cor-als was carried out by a number of workers (e. g. HILL 1973, 1981;DUBATOLOV & VASSILJUK 1980; FEDOROWSKI 1981; 1986; RODRIGUEZ 1984 a,

b and RODRIGUEZet al. 1986). The paleobiogeography of North Africa and thewestern Mediterranean Carboniferous paleotethys was recently discussed byLEGRAND-BLAINet al. (1987) and FLUGEL& HERBIG (1988).

The distribution of rugose coral genera in North America, northern Spain,Donetz Basin, Gulf of Suez and China during the Moscovian and Kasimovian(Westphalian-Stephanian) Epochs is given in Table 1, compiled from differentsources including Guo (1983), RODRIGUEZ(1984b), RODRIGUEZet al. (1986),FEDOROWSKI(1981,1987) and RODRIGUEZ& KULLMANN(1990). Only the validgeneric names listed by SANDO & SANGREE(1990) are used. Also, identifica-tions from the Gulf of Suez based solely on external morphological features arenot considered here. From this table, it is observed that the similarity of coralfaunas in the Gulf of Suez region and the other basins is very striking. Thus,among the fifteen Late Carboniferous rugose coral genera hitherto knownfrom the western side of the Gulf of Suez, ten (67%) were recorded from theDonetz Basin in the U.S.S.R. and from the Cantabrian Mountains in northernSpain, and eleven (73%) were distributed also in the different parts of NorthAmerica, indicating a good communication between these basins.

It is also evident that northern Spain was one of the richest regions of thesouthern Mediterranean subprovince. The fauna closely resembled that of theNorth American basins and that of the Donetz Basin of the U.S.S.R.; the latterwas considered by FEDOROWSKI(1981) as one of the main faunistic centres,although calcareous and shaly facies as well as terrestrial deposits are interbed-ded there.

According to LEGRAND-BLAINet al. (1987), a few North American benthicelements had entered North Africa in the early Late Carboniferous by a poorlyknown passage; either by the NW Tethyan route or by a subsiding channel atthe junction between the West African Craton and the central Sahara. Themaps of FEDOROWSKI(1981) indicate that the northwest seaway which wasopened for corals from the Mediterranean through the Ural Mountains and theArctic into North America in the Moscovian had been closed by Kasimovian-Gzelian times. Since the coral fauna of the Gulf of Suez region, which is partlyKasimovian in age, exhibits apparent similarity to those of Spain and NorthAmerica, the present study suggests that connection between North Africa,Spain and the Donetz Basin with North America might have continued till theend of the Carboniferous (Fig. 6).


Table 1. Distribution of rugose coral genera in North America, Spain, Donetz Basin, Gulfof Suez and China during the Moscovian and Kasimovian Epochs.

Genera North North Donetz GulfChinaAmerica Spain Basin of Suez

Actinophrentis x x xAkagophyllum xAllotropiochisma x xAllotr1ri°'hhyllum x x x xAman op yllum x x xAmplexizaphrentis x x x x xAmplexocarinia x xAmplexus x x x xAmygdallophylloides x x xAntheria xArachnolasma xAsserculinia xAssimulia x x xAxolithophyllum x x xBarytichisma x xBothroclisia x xBothrophyllum x x x xBradyphyllum x x x xCalophyllum xCaninia xCarinthiaphyllum xCorwenia x xCyathaxonia x x xCyathocarinia xCystolonsdaleia x x xDuplophyllum x x xEmpodesma xFomichevella xIvanovia x x xKionophyllum x x xKoninckocarinia x xLithostrotionella xLom'dahyllum xLons aleoides xLophamplexus xLophocarinophyllum xLopholhyllidium x x x xLytvo asma x x xM eniscophyllum x x xN eokoninck°r.hyllum x x x xOrygmophyl um xParacarruthersella xParaduplophyllum x x x xPetalaxis x xPseudowannerophyllum x xPseudoziuhrentoides x x x xRotiphyl um x x x xSestrophyllum x xSochkine°fuhyllum xSpirophyl um xStylostrotion x xTachylasma xVerbeekiella x x x


I/

/I

(.'.~::~

/\-/..d~,/

,,"

I ~-\'.~J

N. AMERICA

500 km--...Fig. 6. Paleobiogeographic relationships of marine benthic organisms in North Africa

during the Late Carboniferous (modified from LEGRAND-BLAINet al. 1987).

Moreover, the first fossiliferous marine horizon recorded in the WadiAraba area (zone A of EL ADINDANI 1968) contains definite Upper Carbonifer-ous fossils, the majority of which have a west European and North Americanaffinity. Similarly, the uppermost Carboniferous flora described by LEJAL-NICOL (in BANDEL & Kuss 1987) from Wadi Qiseib in the study area isessentially an association of mostly Euramerican forms. On the other hand, thepresence of some corals as Paraduplophyllum, Assimulia, Rotiphyllum andAmplexizaphrentis, which are identified in the present study, as well as aprevious record of Amygdallophylloides ivanovi (DOBROLYUBOVA) by HERBIG& Kuss (1988) form the study area, suggests some faunistic connections withChina.

Similarly, RODRIGUEZ et al. (1986) concluded that the zoogeographiccomparisons of northern Spain with North America and the Donetz Basin bymeans of similarity index suggest that a seaway along the juncture of theAfrican, South American and North American plates provided a communica-tion route for coral gene flow between Spain and North America during theentire Late Carboniferous time. However, the maps recently constructed forthe paleobiogeographic relations in North Africa (LEGRAND-BLAIN et al.1987), although satisfactorily indicating that the Tethyan marine migrationscame from Eastern Europe and Middle Asia with exchanges occurring betweenNorth Africa and Southern Europe (Fig. 6), yet claim that connections withAmerican seas had ceased in the late Upper Carboniferous.

40 N. Jb. Geo!. Palaont. Mh. 1991


References

ABDALLAH, A. M. & EL ADINDANI, A. (1965): Stratigraphy of Upper Paleozoic rocks,

western side of the Gulf of Suez. - Geol. Surv. Egypt, paper no. 25: 1-18; Cairo.

ABD ELSHAFY, E. & ABD ELAzEAM, S. (1990): Geology of the Upper Paleozoic sediments onthe western side of the Gulf of Suez. - 7th symposium on Phanerozoic anddevelopment in Egypt, May 1990, Abstracts, p. 19-20; Cairo.

AWAD, G. H. & SAID, R. (1966): Egypt. - Lexique Stratigr. Internat. 4, Afrique, fasc. 4b:4-73; Paris.

BALL, J. (1916): The geography and geology of west-central Sinai. - Egypt. Surv. Dept.,p. 151-163; Cairo.

BANDEL, K. & Kuss, J. (1987): Depositional environment of the pre-rift sediments, GalalaHeights (Gulf of Suez, Egypt). - Berliner geowiss. Abh. (A), 78: 1-48; Berlin.

BOLL, F. C. (1983): Der Wandel der Rugosen Korallenfaunen der Flachwasser-Fazies im

Karbon des Kantabrischen Gebirges (Nordspanien). - Ph. D. Thesis, Eberhard-Karls-Universitat Tiibingen, 275 p.; Tiibingen.

DE GROOT, G. D. (1963): Rugose corals from the Carboniferous of northern Polenica,Spain. - Leid. Geol. Meded., 29: 1-37; Leiden.

DUBATOLOV, V. N. & VASSILJUK, N. P. (1980): Coral paleozoogeography in the Devonian

and Carboniferous of Eurasia. - Acta Palaeont. Polonica, 25 (3-4): 519-529;Warsaw.

EL ADINDANI, A. (1968): Stratigraphical and paleontological studies on some Permo-

Carboniferous rocks of Egypt. - M. Sc. Thesis, Alexandria Univ., 98 p.; Alexandria.FEDOROWSKI, J. (1981): Carboniferous corals: distribution and sequence. - Acta Palaeont.

Polonica, 26: 87-160; Warsaw.

(1986): The rugose coral faunas of the Carboniferous/Permian boundary interval.-

Acta Palaeont. Polonica, 31 (3-4): 253-275; Warsaw.

(1987): Upper Paleozoic rugose corals from southwestern Texas and adjacent areas:

Gaptank Formation and Wolfcampian corals, Part I. - Palaeontologia Polonica, 48:

1-271; Warsaw.

FLUGEL, H. W. (1975): Zwei neue Korallen der Sardar-Formation (Karbon) Ost-Irans. -Mitt. Abt. Geol. Palaont. Bergb. Landesmus., 35: 45-53; Graz.

FLUGEL, E. & HERBIG, H.-G. (1988): Microfacies of Carboniferous Limestone clasts from

the Rif Paleozoic (Morocco): a contribution to the paleogeography of the westernMediterranean Carboniferous Paleothethys. - Facies, 19: 271-300; Erlangen.

FOMICHEV, V. D. (1953): Rugose corals and stratigraphy of the Middle and Upper

Carboniferous and Permian deposits of the Donetz Basin (Russian). - Trudy vses-

nauchno-issled-geol. Inst. Vsegei Miner. Geol., 622 p.; Moscow.

GRABAU, A. W. (1928): Paleozoic corals of China, Part I, Tetraseptata II. - Palaeontol.Sinica, B, 2: 1-175; Beijing.

Guo, SHENG-ZHE (1983): Middle and Upper Carboniferous rugose corals from southernDahinganling (Mts. Great Khingan). - Acta Palaeontologica Sinica, 22(2): 220-229;Beijing.

HERBIG, H.-G. & Kuss,J. (1988): The youngest Carboniferous rugose corals from Northern

Africa (NE Egypt) - palaeoenvironment and systematics. - N. Jb. Geol. Palaont.Mh., 1988 (1): 1-22; Stuttgart.


HILL, D. (1973): Lower Carboniferous corals.- [In:] Hallam, A. (ed.): Atlas of paleobioge-

ography: 134-142; Amsterdam (Elsevier).

(1981): Rugosa and Tabulata. - Treat. Invert. Paleont., Part F Coelentrata, Sup pI. 1:FI-F428; Lawrense.

IVANOVSKIY, A. B. (1967): Studien iiber friihkarbone Rugosa.- Nauka: 1-92; Moscow.

Jux, U. (1986): On the Paleozoic framing the Arabian-Nubian Shield. - Symposium ongeological problems of NE Africa, May 1984, Whittier College; California.

KLEBELSBERG, R. (1911): Ein Beitrag zur Kenntnis des Sinai-Karbons. -2. dt. geol. Ges., 63:

594-603; Hannover.

KORA, M. (1989): Lower Carboniferous (Visean) fauna from Wadi Budra, west-centralSinai, Egypt. - N. Jb. Geol. Palaont. Mh., 1989 (9): 523-538; Stuttgart.

KORA, M. & Jux, U. (1986): On the early Carboniferous macrofauna from the Urn Bogma

Formation, Sinai. - N. Jb. Geol. Palaont. Mh., 1986 (2): 85-98; Stuttgart.

KULLMANN, ]. & LIAO, W.-H. (1985): Hornformige Einzelkorallen (Rugosa) aus dem

Unterkarbon von Siid-China. - Palaeontographica Abt. A, 189: 125-157; Stuttgart

LEGRAND-BLAIN, M.; CONRAD,].; COQUEL, R.; LEJAL-NICOL, A.; Lys, M.; PONCET,]. &

SEMENOFF-TIAN-CHANSKY (1987): Carboniferous palaeobiogeography of NorthAfrica. - XI Inter. Congr. Carboniferous, Beijing 1987, preprint, 16 p.; Beijing.

OMARA, S. (1967): Contribution to the stratigraphy of the Egyptian Carboniferous

exposures. - 6th Arab Petrol. Congr., paper, 44, (B-3); Baghdad.

(1971): Early Carboniferous tabulate corals from Urn Bogma area, southwesternSinai, Egypt. - Riv. Ital. Paleont., 77 (2): 141-154; Milano.

POTY, E. (1981): Recherches sur les Tetracoralliaires et les Heterocoralliaires du Viseen de la

Belgique. - Meded Rijks. Geol. Dienst, 35 (1): 1-161; Roermond.RODRIGUEZ, S. (1984 a): Corales rugosos del Carbonifero del este de Asturias. - Univ.

Complutense Madrid, Col. Doc. 109/84, 258 p; Madrid.

(1984 b): Carboniferous corals from eastern Cantabrian Mountains: Paleogeo-graphic implications. - Paleontographica Americana, 54: 433-436.

RODRIGUEZ, S. & KULLMANN,]. (1990): Hornformige Einzelkorallen (Rugosa) aus spatober-

karbon is chen Flachwasser-Ablagerungen des Kantabrischen Gebirges (Nordspa-nien). - Palaeontographica, A, 210 (1-3): 19-40; Stuttgart.

RODRIGUEZ, S.; SANDO, W.]. & KULLMANN,]. (1986): Utility of corals for biostratigraphic

and zoogeographic analyses of the Carboniferous in the Cantabrian Mountains,Northern Spain. - Trabajos de Geologia, Univ. de Oviedo, 16: 37-60.

SANDO, W.]. (1980): The paleoecology of Mississippian corals in the western counterminous

United States. - Acta Palaeont. Polonica, 25 (3-4): 619-631; Warsaw.

SANDO, W.]. & BAMBER, E. W. (1985): Coral zonation of the Mississippian System in the

Western Interior Province of North America. - U.S.G.S. Prof. paper 1334: 1-61;Washington.

SANDO, W.]. & SANGREE, A. C. (1990): List of names proposed for genera and subgenera of

Late Paleozoic corals, 1828-1989. - Fossil Cnidaria, 19 (2.2): 1-21; Miinster.

SCHOUPPE, A. & STACUL, P. (1955): Die genera Verbeekiella PENECKE, TimorphyllumGERTH, Wannerophyllum n. gen., und Lophophyllidium GRABAU aus dem Perm von

Timor. - Palaeontographica, Suppl. Bd., 4, V Abt. (3): 95-196; Stuttgart.

WALTHER,]. (1980): Uber eine Kohlenkalk-Fauna aus der agyptisch-arabischen Wiiste.- 2.

dt. geol. Ges., 42: 419-449; Berlin.

614 M. Kora and Y. Mansour, Late Carboniferous solitary rugose corals

WEYER, D. (1972): Rugosa (anthozoa) mit biformen Tabularium. - Jahrb. Geol., 4:439-463.(1975): Zur Taxonomie der Antiphyllinae lUNA. - Z. geol. Wiss., 3: 755-775; Berlin.

WILSON, E. C. (1990): Permian corals of Bolivia. - J. Paleont., 64 (1): 60-78; Menasha.

Bei der Tiibinger Schriftleitung eingegangen am 4. Marz 1991.

Anschrift der Verfasser:

Dr. MAHMOUD KORA, YASSERMANSOUR,Department of Geology, Faculty of Science,Mansoura University, Mansoura 35516, Agypten.

Biostratigraphy and paleoecology of some Cenomanian successions 617

HECKEL,P. H. (1972): Recognition of ancient shallow marine environments. [In:]RIGBY,J. K. & HAMBLIN,W. K. (eds.): Recognition of ancient sedimentaryenvironments. - Soc. Econ. Paleont. Miner., Spec. Publ., 16: 226-286; Tulsa.

KLITZSCH, E. & HERMINA, M. (1989): The Mesozoic. [In:] HERMINA, M.; KLITZSCH,E. & LIST,F. K. (eds.): Stratigraphic lexicon and explanatory notes to thegeological map of Egypt 1 :500000: 77-140; Cairo (Conoco Inc.).

KORA,M. & EL BEIALY,S. (1989): Early Cretaceous palynomorphs from GabalMusaba Salama area, southwestern Sinai, Egypt. - Rev. Palaeobot. Palynol.,58: 129-138; Amsterdam.

KORA,M.; SHAHIN,A. & SEMIET,A. (1993): Stratigraphy and macrofauna of theCenomanian exposures in west-central Sinai, Egypt. - Mans. Sci. Bull. 20(2).: 227-260; Mansoura.

Kuss, J. (1986): Facies development of Upper Cretaceous-Lower Tertiarysediments from the Monastery of St. Anthony, Eastern Desert, Egypt. -Facies, 15: 177-194; Erlangen.

LIPSON-BENITAH,S. (1980): Albian to Coniacian zonation of the western coastalplain of Israel. - Cretaceous Research (1980), 1: 3-12,4 figs.; London.

MANCINI,E. A. (1977): Depositional environment of the Grayson Formation(Upper Cretaceous) of Texas. - Gulf Coast Assoc. Geol. Soc. Trans., 27:334-351; Washington.

McRAE, S. G. (1972): Glauconites. - Earth Sci. Rev., 8: 397-440; Amsterdam.MURRAy,J.W. (1973): Distribution and ecology of living benthic foraminiferids. -

Crane: 1-274; New York (Russak & Co.).OMARA,S. (1956): New foraminifera from the Cenomanian of Sinai. - J. Paleont.,

30 (4): 883-890; Menasha.SAID,R. & BARAKAT,M. G. (1957): Cenomanian foraminifera from Gebel Asagil,

northern Sinai. - Egypt. J. Geol., 1 (1): 65-83; Cairo.SCHLUMBERGER,C. (1893): Foraminifera. [In:] THOMAS,P. (ed.): Exploration

Scientifique de la Tunisia: 5-7, 1 pI.; Paris (Imprimerie Nationale).SHAHIN,A. (1988): Fossil fauna and stable isotopic composition within the Late

Cretaceous-Early Tertiary at Gebel Nezzazat, Sinai, Egypt. - Ph. D. thesis,Mansoura Univ., 212 p.; Mansoura.

SHAHIN,A. &KORA,M. (1991): Biostratigraphy of some Upper Cretaceous succes-sions in the eastern Central Sinai, Egypt. - N.Jb. Geol. PaHiont. Mh., 1991(11): 671-692; Stuttgart.

SHATA,A. (1960): The geology of Sinai Peninsula. [In:] Encyclopedia of Sinai (inArabic), Essay no. 5: 124-167; Cairo (Government Press).

WASPI,S. & HATABA,H. (1983): Observations on the genus Nezzazata OMARAandits significance to the Cenomanian-Turonian boundary in the Gulf of Suez.- Benthos, 83: 2nd Intern. Symp. Benthonic foraminifera; Pau.

Bei der Tiibinger Schriftleitung eingegangen am 17. Marz 1993.

Anschrift der Verfasser:

Dr. MAHMOUD KORA, Dr. ABDALLA SHAHIN and AHMED SEMIET, Department ofGeology, Faculty of Science, Mansoura University, Mansoura, Agypten.

,"",,","1-

late carboniferous solitary rugose corals from the western side of the gulf of suez, egypt

Documents