New algae and problematica of algal affinity from the permian of the Aseelah Unit of the batain plain (East Oman)

N E W / H JGAE A N D P R O B L E M A T I C A OF AI,G/HJ AFFINITY F R O M THE P E R M I A N OF THE A S E E L A H U N I T OF THE BATAIN P L A I N (EAST ©MAN) DANIELVACHARD, MARC HAUSER, ROSSANAMARTINI, LOUISETTEZANINETTI, ALBERTMATTER, TJERKPETERS Vachard, D., Hauser, M., Martini, R., Zaninetti, L., Matter, A., Peters, T., 2001. New algae and problematica of algal affinity from the Permian of the Aseelah Unit of the Batain Plain (East ©man). [Nouvelles algues et formes affines probl~matiques du Permien de l'Unit~ d'Aseelah (Plaine de Batain, Est ©man)]. Geobios 34 (4), 375-404. Villeurbanne, le 30.09.2001. Manuscrit dSpos~ le 27.07.2000; accept~ d~finitivement le 07.03.2001. ABSTRACT - Permian microfloras occur in limestone boulders of the Aseelah Unit at the lower part of the Batain Group (NE ©man). The boulders are composed of bioclastic limestones deposited during the Permian on a shallow marine warm shelf environment. They yield several rich associations of Dasycladales, tubiphytids and Archaeolithophyllaceae indicating a Yakhtashian to Dzhulfian age (dated by fusulinids). The algal-bearing Aseelah Unit overlies conformably the Permian Qarari Unit deposited on the distal slope of the Arabian carbonate platform. The conglomerate and sandstone sheddings of the Aseelah Unit indicate the end of a major transgressive-regressive cycle at the Permian/Triassic boundary. The oldest age of the depositional event is given by the Dzhulfian age of the youngest reworked boulders. The age of the sandstones and sandy matrix is not well established; it is postDzhulfian, probably Triassic on the basis of some palynological indications. Several new taxa are described: Aphralysiaceae nov. fam., Sparaphralysia orientalis nov. sp., Mellporellopsis corpulenta nov. gen. nov. sp., Aseelahella granieri nov. gen. nov. sp., Claracrustinae nov. subfam., Claracrusta ungdarelloidea nov. sp. and Sphairionia galinae nov. sp. © 2001 Editions scientifiques et m~dicales Elsevier SAS KEYWORDS: ©MAN, BATAIN PLAIN, ASEELAH UNIT, PERMIAN, ALGAE, MICROPROBLEMATICA. R]~SUMt~ - Des microflores permiennes sont pr~sentes dans les galets des conglom~rats de l'Unit~ d'Aseelah, qui constitue la partie inf~rieure du Groupe de Batain (NE ©man). Les galets, contenus dans une matrice sableuse, sont compos~s de calcaires bioclastiques d~pos~s au Permien sur une plateforme marine peu profonde, en climat chaud. Ils contiennent de nombreuses associations de Dasycladales, Tubiphytidae et Archaeolithophyllacea, associ~es ~ des fusulines, indiquant un age yakhtashien ~ djoulfien. L'Unit~ d'Aseelah surmonte stratigraphiquement l'Unit~ permienne de Qarari, d~pos6e darts la partie distale de la plateforme carbonat6e Arabique. L'Unit~ conglom~ratique d'Aseelah est le t~moin de la phase terminale d'un cycle transgressif/r~gressif majeur, ~ la limite Permien-Trias. L'age de l'~v~nement d~positionnel, ~ rorigine de l'Unit~ d'Aseelah, se situe entre le Djoulfien, age le plus jeune identifi6 parmi les galets remani~s, et le Trias inf~rieur, age probable de la matrice sableuse selon quelques donnSes palynologiques. Plusieurs associations permiennes, de rYakhtashien au Djoulfien, sont d~crites. Les taxons c r ~ s sont: Aphralysiaceae nov. faro., Sparaphralysia orientalis nov. sp., MellporeUopsis corpulenta nov. gen. nov. sp., Aseelahella granieri nov. gen. nov. sp., Claracrustinae nov. subfam., Claracrusta ungdarelloidea nov. sp. and Sphairionia galinae nov. sp. © 2001 Editions scientifiques et m6dicales Elsevier SAS MOTS-CLI~S: ©MAN, PLAINE DE BATAIN, UNITI~ D'ASEELAH, PERMIEN, ALGUES, MICROPROBLEMATICA.

INTRODUCTION I n the course of a geological r e s e a r c h project in the B a t a i n P l a i n ( E a s t ©man), r e w o r k e d P e r m i a n limestone boulders were discovered f r o m s a n d y limestone conglomerates belonging to the A s e e l a h U n i t (Peters et al. 2000). T h e s e boulders are m a d e up of s h a l l o w - m a r i n e bioclastic l i m e s t o n e s which yield a n exceptional microflora with new f o r m s of P e r m i a n algae. Algal occurrences are well k n o w n from © m a n , e.g. from the a u t o c h t h o n o u s s e d i m e n t s in the H a j a r S u p e r - G r o u p , or in the so-called ' © m a n Exotics' facies w i t h i n the allochthonous H a w a s i n a s e d i m e n t s (Glennie et al. 1974). F r o m

the B a t a i n Plain, S h a c k l e t o n et al. (1990) mentioned the exposures of p i n k i s h algal a n d reef-limestones associated w i t h g r e y n o d u l a r bedded h e m i p e lagic l i m e s t o n e s . B a s e d on t h e a g e - d i a g n o s t i c micro- a n d m a c r o f a u n a , S h a c k l e t o n et al. (1990) c o m m i t t e d a t i m e r a n g e f r o m E a r l y to L a t e P e r m i a n for this a s s e m b l a g e . This p a p e r a i m s to p r e s e n t the s y s t e m a t i c palaeontological description of the new taxa. F u r t h e r m o r e , a n a t t e m p t is m a d e to c o m p a r e the algae a n d associated p r o b l e m a t i c a from t h e B a t a i n P l a i n w i t h those of other occurrences f r o m © m a n a n d Arabia. The microflora, m i c r o f a u n a a n d microfacies from

376 palaeogeography and geodynamic evolution of this part of the Tethyan Basin. Hadd

al

GEOLOGICAL SETTING The algal-bearing conglomerates of the Aseelah Unit are located in the Batain area, South of the town Sur. The Batain Plain extends about 130 km NE-SW and 40 km in E-W direction (Fig. 1). It is bounded to the North by the Gulf of Oman and to the East by the Arabian Sea; the Wahibah sands separate the Batain Plain in the West from Interior Oman.

Ad D a f f a h

1Khabbah ays h 20kin

I'

Om

FIGURE 1 - Simplified map of the Batain Plain showing the locations of the sampled outcrops of the Aseelah Unit. Carte simpli-

fide de la Plaine de Batain montrant la localisation des affieurements gchantillonnds de l'UniM d'Aseelah.

the Permian of the Batain Plain, compared to occurrences from the Arabian Platform, need a reinterpretation of the Middle and Late Permian assemblages and facies distributions in the Southern Neotethys, as well of the biodiversity, HAWASINA

Maastrichtian Campanlaa Santonian : Coniacian Tumniau

0 "~

O "~

BATAIN NAPPES

NAPPES

Shaekleton et aL (1990)

B~chennec et aL (1992) i:2~o,0oot~o~d ~p

Batain Melange

Al Aridh Gp Hamrat Duru Groul:

....

~

Nayid Fm

Peters et aL (2000) Batain Group

~ ~

Sid'r Fm

Wahrah Fm "~

li

ydM~ t

Callovian

Guwayza Fm

Bathor,ian Aalenian Toareian PliensbaehJala Shlemurian Heltaugian Rbaetian Noriaa Camian Ladini~ Anisian

The Permian boulders were found in the Aseelah Unit, as an assemblage of calcareous sandstones and unsorted reworked polymict limestone conglomerates with a sandy matrix. These conglomerates yield, in contrast to other conglomerates from the Batain Plain, exclusively Permian shallow marine macro- and microfossils. The Aseelah Unit is often associated with the Qarari Unit, or found in tectonic contact with the Sal Formation or the Guwayza Formation (Fig. 2).

Fayah Fm

: O i C~nomanian I Aptian BatrenCan Haaterivian Valan~iman Berriasian Tahorfian Kimmeridgian Oxfordian

The Batain Plain comprises the Batain nappes, which are composed of Permian to Maastrichtian marine sediments, as well as volcanic rocks and the eastern Ophiolite nappe (Peters et al. 2000). Tectonically, the Batain Plain is characterised by intense folding and thrusting, indicating a WNW directed nappe transport and obduction of the Batain nappes onto the Arabian continent (Schreurs & Immenhauser 1999). The Batain nappes are unconformably overlain by autochthonous Late Palaeocene to Miocene continental siliciclastics and shallow marine limestones.

Gwl

Gws

o~

Matbat Fm

Unit

N

"Scymia¢ Aseelah Unit AI Jil F m

Qarari Unit

FIGURE 2 - Overview of the stratigraphy of Oman, showing the different stratigraphic subdivisions in the Batain Plain, and the stratigraphic position of the Aseelah Unit. Ta-

bleau synoptique de la stratigraphie d'Oman, montrant les diffdrentes subdivisions pour la Plaine de Batain , ainsi que la position stratigraphique de l'Unitd d'Aseelah.

377 PREVIOUS WORK

According to Glennie et al. (1974), algal-bearing conglomerates of reworked Permian and Triassic limestones in the Batain Plain displaying an 'Oman Exotics' facies have been assigned to the Ibra Formation; they were included as limestone breccias in the Batain Mrlange by Shackleton et al. (1990) (Fig. 2). Roger et al. (1991), Brchennec et al. (1992a) and Wyns et al. (1992) attribute these conglomerates of the Batain Plain to the A1 Jil Formation, which was defined by Brchennec (1987) in the Oman Mountains and later redefined by Brchennec et al. (1992b). In fact, according to the litho- and biostratigraphic data obtained from this study, the name A1 Jil Formation appears to be inadequate for the Batain Plain. For this reason, Peters and others (2000) introduced, to replace the A1 Jil Formation, a new lithostratigraphic subdivision for the Permian and a part of the Triassic in the Batain Plain, with respectively the Qarari Unit, the Aseelah Unit and the Ad Daffah Unit (Fig. 2); they also include the Batain series into the Batain Group of Immenhauser et al. (1998).

Pillevuit et al. (1997) were the first to distinguish the 'Oman Exotics' according to the sequence of the depositional events; they present a revised classification of the 'Oman Exotics' into different palaeogeographic units, and link the exotic blocks from the Batain Plain to the 'Reworked Permian platform limestones in the Hawasina basin' (Pillevuit et al. 1997). The lithological and palaeontological description of the Aseelah Unit was published in a preliminary paper by Hauser et al. (2000).

STRATIGRAPHY

The algal-bearing boulders were found in Permian unsorted clasts or matrix supported sandy conglomerates and coarse grained sandstones, attributed to the Aseelah Unit by Peters et al. (2000). The Aseelah Unit is well exposed 3 km SW of Aseelah (UTM coord. 769662/2428915), and 6 km NW of the village Bu Fashiqah (UTM coord. 775769/2468896), in the surrounding area of the village Musawa. Most of the algal-bearing boulders were found in the Wadi Hasad (UTM coord. 759068/2444367) 10 km WSW from the village Sal (Fig. 1).

Gastropods

Limestone Nodular bedded limestone

m

(~

Crinoids

&

Fnsulinids

Dolomite

Bivalves

& &

Marly shales

@

Ammonoids

Sandstone

(~

Nautiloids

"~7

Brachiopods

"~

Bryozoa

Conglomerate Breccia beds Conglomerate beds

Ripple L~

Slump

6~

N

%_ FIGURE 3 - Sketch of the Bu Fashiqah outcrop (UTM coord. 775769/2468896), and detail of the lithological section (modified after Wyns et al. 1992).

immmlm£:=::::::::~

Panorama de l'affieurement de Bu Fashiqah (Coordonnges UTM 775769/ 2468896), et ddtail de la colonne lithostratigraphique (modifide d'apr~s Wyns et al. 1992).

378

Jabel Jalaan

NNE

........

~

..............~

. . . . .

FIGURE4 - Overviewofthe WadiHasad outcrop showingthe tectoniccontactbetweenthe limestoneconglomeratefromthe AseelahUnit (As) and the stronglyfoldedlimestonesand shales fromthe Sal Formation(S). Panorama de l'affleurement de Wadi Hasad, montrant le contact tectonique entre les conglomgrats calcaires de l'Unitd d'Aseelah (As) et les calcaires et argilites fortement plissds de la Formation de Sal (S).

Subsystems LOPINGIAN (= LatePermian)

Stages

I

DORASHAMIN',I DZHULFIAN

MIDIAN GUADALUPIAN

MURGABIAN

(= MiddlePermian) KUBERGANDIAN

CISURALIAN (=EarlyPermian)

Samples

III111111111111t1111111 H245, H253,H256 H250 H132; H133;H489;H541

The conglomerate is made up of centimetre to metre-sized exclusively Permian shallow marine limestone blocks and boulders, and rare reworked volcanics. The limestone components are made up of bioclastic wackestones, packstones and rudstones containing a Yakhtashian to Dzhulfian microfauna of fusulinids (Hauser et al. 2000; Vachard et al. in press); Wentzelellid Rugosa corals and Richtofeniid brachiopods also occur.

H249 H180;H236;H240;H241

BOLORtAN

H19

YAKHTASHIAN

H233; H253

SAKMARIAN~s~u~N

the village Sal, is exposed as a thrust slice within the Sal Formation (Fig. 4).

PERMIAN SUBDIVISIONS As the here presented biostratigraphy based on Tethyan fusulinids, the subdivisions proposed by Leven, and summarised for example in Leven (1997), are accepted for the chronostratigraphic names of stages (Fig. 5). The official scale of Jin et al. (1997) slightly differs, because essentially based on North-American endemic fusulinids.

II]11111111111111111111

FIGURE 5 Adopted Permian chronostratigraphy, according to Leven(1997), and age ofthe fossiliferoussample(see alsoHauser and others, 2000). Classification chronostratigraphique du Permien, adoptde d'apr~s Leven (1997), et ¢tge des gchantillons fossilif&es (voir aussi Hauser et al. 2000). -

The mudstones and dolomites from the Qarari Unit are overlain by the moderate to very poorly sorted but well rounded polymict clast-supported limestone conglomerate of the Aseelah Unit. The conglomerate is composed of centimetre up to metre-sized fossiliferous boulders and recrystallised limestone blocks. The sandy limestone conglomerate passes over in a succession of cross-bedded white sandstone, the same as the conglomerate matrix; this sandstone, of fluviatile origin, is made up of very coarse grains, well sorted and sub-angular rounded. SW of Aseelah, the sandstones show two horizons of a dark brown hematite rich duricrust, indicating old weathering surfaces. The total thickness of the Aseelah Unit is difficult to estimate; at the locality NW of Bu Fashiqah section (Fig. 3), it measures approximately 15 m, and SW of Aseelah circa 35 m. The outcrop in the Wadi Hasad, 10 km WSW from

The successive Permian stages, and/or biozones, of the Batain Plain are characterised by Leeina kraftti (Yakhtashian); 'Parafusulina' japonica (Bolorian); Toriyamaia, Skinnerella spp. and smaller foraminifers Endoteba, A g a t h a m m i n a , Neoendothyra and Abadehella (Kubergandian); Neoschwagerina simplex (Early Murgabian); Colania douvillei, Neoschwagerina spp., Kahlerina, Pseudokahlerina, Reichelina and Codonofusiella (Early Midian); an undetermined assemblage of Chusenellenellids and Palaeofusulinids (supposed intermediary between the 'keriotheca stage' and 'postkeriotheca stage' of Sheng 1992) (Late Midian); and finally an association of Codonofusiella schubertellinoides and smaller foraminifers such as Robuloides, Robustopachyphloia, Midiella, etc. (Dzhulfian).

PREVIOUS WORK ON ALGAE The carbonate skeletal Permian algae (sensu lato) belong to five groups: (1) green dasycladale algae;

379 Yakhtashlan Bolodan H233 H253

H19

1 4rchaanlithoporeHa hidensis Shvane#a sp.

Early Late Early Early Murgablan Midian Midian DzhulEan Ht80 H236 H240 H249 H132 H489 H250 H245 H256 H241 Ht33 H641 • • (240) Kubergandian

2

• (240)

FIGURE 6 - Recapitulative table of the Permian algae and pseudo-algae of the Batain Plain. Tableau r~capitulatif des algues et pseudo-algues perrniennes de la Plaine de Batain.

• (541)

"-3-" Sparaphralysia oriantaHs 4

gacine~ sp, Vostonites veaicu/osa Spor~diostrom~s

7 8 _9 10 11"

12 13 14 15

o(132)

i


?

• • (240)

M~ziavelebitana Epimastoporasp. Diplopora sp.

• •

•

•

•

•

Kochanskye#atut~oa

• (~I)

•

•

16 LP,anet/a sp. 17 Aseelehel/agrargefl 18 Parachaetetes sp, 19 Permoca/culussp. Gymnocodium exile 21 Atchano~thophy/lum missoudense 22 Eugonophyllum sp, 23 Ungdare//a ura~ca 24 Eflugeliajohnsanl 25 Ev/antopsis montana 26 Ctaracruata ungda~/toidea 27 Claracrusta hirEpea 28 C/atactusta spp. ~ " Stacheo~s sp. 30 Sphairianle gar~ae 31 Spha'~bn/a s/kmo/des

• ,

•

?

•

?

•(241) •

•

•

•

i • (240) • (~!~

o .......

•

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• (240)

" • (133) e(489)

•

•

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•

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•

•

•

(240)

• (N1} • (54t) ,(!32)

• •

..

•

•

"A~!) • (132) e (489)

3 3 Tub~hytes obscurus

o

•

34 D~o/asphantaceae

35 Pseudoverm~ore/la spp.

•

•

4

•

• •

•

•

I

(2) other green algae; (3) red gymnocodiacean and solenoporacean; (4) other red algae; (5) various pseudo-algae generally attributed to green- (moravamminids) or red algae (aoujgaliids), foraminifers (tuberitinids), or incertae sedis (Sphairionia, Tubi-

phytes, Pseudovermiporella ). Pionneer works on Triassic and Permian dasycladales were conducted by Pia (1912, 1920, 1937), and they are still used. More recent literature on dasyclad algae is composed of two groups of publications: (1) the theoretical approaches based on compilations (Endo 1961b; Johnson 1963; Emberger 1976, 1979; Fl(igel 1977, 1979; Bassoullet et al. 1979; Gtivenc 1979; Roux 1985; Riding & Guo 1991; Chuvashov et al. 1987, 1993; Deloffre 1987, 1988; Deloffre & Granier 1991; Granier & Deloffre 1994; De Castro 1997; Granier & Grgasovic 2000); (2) the practical works on systematics (Accordi 1956; Praturlon 1963; Kord4 1965; Kochansky-Devid4 & Herak 1960; Endo 1961a, 1969: in this work, 33 genera and 64 species were described in the famous Khao Phlong Phrab in Thailand; Praturlon 1963; Elliott 1968; Gtivenc 1979; Kochansky-Devid4 1970a, b; Homann 1972; Chuvashov 1974; Kulik 1978; Vachard 1980, 1985; Vachard & Montenat 1981; Fltigel 1966, 1980, 1990; Flfigel & FltigelKahler 1980; Mu 1982; Mamet et al. 1987; Chuvashov & Anfimov 1988). Other green algae are poorly known, especially the Caulerpales/Codiaceae (see Pia 1937; Bassoullet et al. 1983; Roux 1985).

__,_o~132) /

•

•

•

Among the red algae, the gymnocodiacean are abundantly described (Pia 1937; Elliott 1955; Roux 1985, 1991; Roux & Deloffre 1990). Species of solenoporacean are also numerous (see compilations in Emberger 1976), but recently nothing new was published about these enigmatic algae. Other true Permian red algae are poorly known, with the exception of the genus Archaeolithophyllum (Vachard et al. 1989; Moshier & Kirkland 1993). The Permian moravamminids and aoujgaliids pseudo-algae were described and discussed by Chuvashov (1974), Vachard (1980), Vachard & Montenat (1981), etc. We add to the algal incertae sedis the problematical family of the Tuberitinaceae (nomen translat.), which is more frequently attributed to the foraminifers. The genus Tub# phytes is doubtful, and has been successively interpreted as alga, foraminifer, consortium alga-foraminifer, sponge. Pseudovermiporella, a true porcelaneous foraminifer (Henbest 1963; Loeblich & Tappan 1964), is traditionally considered as an alga or pseudo-alga; this genus is basically derived from the genus Hedraites or other calcivertellid ancestors. Recent works however still admit an algal affinity (i.e. Granier & Deloffre 1994). Few monographs are concerned with Omanese Permian algae (Elliott 1962, 1968; Montenat et al. 1976; Glennie et al. 1974); Elliott (1968) examined a collection in Fahud, composed of Anthracoporella spectabilis P~A, 1920; A. mercurii ELLIOTT, 1968; Diplopora (?) sp.; Pseudoepimastopora cf. likana

380 KOCHANSKY-DEVIDI~ & HERAK, 1960; Pseudovermiporella sodalica ELLIOTT,1958, described from the Jebel Qamar in Oman. Aeolisaccus dunningtoni ELLIOTT,1958, and Anchicodium sindbadi ELLIOTT, 1970, were also defined from Omanese material. Glennie et al. (1974) illustrated Gymnocodium,

Permocalculus, Pseudoepimastopora, Tubiphytes, Mizzia and Ivanovia, and Vachard (in Montenat et al. 1976) identified the first Velebitella and pseudoalgae moravamminids and aoujgaliids from Oman (Jebel Akhdar). The here studied Permian microfloras are relatively different from those of the Khuff Formation in Saudi Arabia (Racz 1959; Okla 1992), which exhibits similarities with the Bellerophonkalk of the Carnic Alps (abundance of Mizzia and Gymnocodiaceae); both formations probably belong to the Upper Dzhulfian or to the Dorashamian.

COMPOSITION AND AGE OF ALGAL MICROFACIES (Fig. 6)

H233 (Yakhtashian): Bioclastic wackestone, poorly sorted, strongly dolosparitized with: algae: Mellporellopsis cf. corpulenta nov. gen. nov. sp. (Fig. 10.4); Epimastopora sp. incertae sedis: Eflugelia johnsoni(FL~GEL, 1966) VACHARDin Massa & Vachard, 1979; Ungdarella uralica MASLOV, 1956; Diplosphaerina inaequalis (DERVILLE, 1931); Mendipsia conili (TIEN, 1979); Tubiphytes obscurus MASLOV,1956. ti242 (Yakhtashian): Bioclastic dolosparitized floatstone with: incertae sedis: Diplosphaerina inaequalis (DERVILLE, 1931); Tubiphytes obscurus MASLOV,1956. H253 (Yakhtashian mixed with Dzhulfian, according to the fusulinids): Bioclastic and intraclastic floatstone, with: algae: Mellporellopsis corpulenta nov. gen. nov. sp. (Fig. 10.5-9); Diplopora sp. (Fig. 14.7). incertae sedis: Eflugelia johnsoni (FL~GEL, 1966) VACHARDin Massa & Vachard, 1979; Diplosphaerina inaequalis (DERVILLE,1931); Tubiphytes obscurus MASLOV,1956. ti19 (Bolorian): Bioclastic rudstone with: algae: Epimastopora sp.; Diplopora sp.

incertae sedis: Diplosphaerina inaequalis (DERVILLE, 1931); Taberitina collosa REITLINGER,1950; and Tubiphytes sp. H180 (Kubergandian): Bioclastic rudstone with: algae: ArchaeolithoporeUa hidensis ENDO, 1961a; Anthracoporella spectabilis PIA, 1920 (Fig. 12.10); Epimastopora sp.; Diplopora sp.; Aseelahella granieri nov. gen. nov. sp. (Fig. 17.9). incertae sedis: Claracrusta hirtipes VACHARD,1980; Claracrusta sp.; Pseudovermiporella nipponica (ENDo in Endo & Kanuma, 1954); Tubiphytes obscuru8 MASLOV,1956 (Fig. 24.5). H236 (Kubergandian): Bioclastic and intraclastic grainstone with: algae: Diploporaceae gen. indet.; Permocalculus sp.; Aseelahella granieri nov. gen. nov. sp.

incertae sedis: Claracrusta hirtipes VACHARD,1980; Claracrusta sp.; Eflugelia johnsoni (FLOGEL,1966) VACHARDin Massa & Vachard, 1979; Tubiphytes obscurus MASLOV, 1956 (numerous morphotypes, Fig. 24.6-7). H240 (Kubergandian): Bioclastic rudstone with: algae: Archaeolithoporella hidensis ENDO, 1961a; Girvanella sp.; Thaiporella aff. uralica CHUVASHOV, 1974 (Fig. 10.10); Anthracoporella cf. spectabilis PIA, 1920; Epimastopora sp.; Diplopora sp. (Fig. 14.6); Permocalculus sp.; Kochanskyella tulipa MILANOVIC,1974; Aseelahella granieri nov. gen. nov. sp. (Fig. 17.11). incertae sedis: Claracrusta hirtipes VACHARD,1980; Claracrusta ungdarelloidea nov. sp. (Fig. 18.3-4); Eflugeliajohnsoni (FLUGEL,1966) VACHARDin Massa & Vachard, 1979; Ungdarella uralica MASLOV,1956; Diplosphaerina inaequalis (DERVILLE, 1931); Tttbiphytes obscurus MASLOV,1956 (numerous morphotypes). H241 (Kubergandian): Bioclastic rudstone with: algae: AnthracoporeUa spectabilis PIA, 1920; Epimastopora sp.; KochanskyeUa tulipa MILANOVlC, 1974 (Fig. 14.2-4); Likanella sp. (Fig, 14.5); Diplopora sp; Aseelahella granieri nov. gen. nov. sp. (Fig. 17.1, 4-8, 10, 12-14). incertae sedis: Claracrusta ungdarelloidea nov. sp. (Fig. 18.2, 5-6); Eflugelia johnsoni (FL~GEL, 1966) VACHARD in Massa & Vachard, 1979 (Fig. 18.7); Diplosphaerina inaequalis (DERVILLE, 1931); Mendipsia conili (TIEN, 1979); Tubiphytes obscurus MASLOV, 1956. H249 (Early Murgabian): Bioclastic dolosparitized rudstone with : algae: Eugonophyllum sp.; Anthracoporella spectabilis PIA, 1920; Mizzia velebitana SCHUBERT,1908. incertae sedis: Ungdarella uralica MASLOV, 1956; Diplosphaerina inaequalis (DERVILLE,1931). H132 (Early Midian): Bioclastic and intraclastic floatstone with: algae: Spongiostromids (into intraclasts); Archaeolithoporella hidensis ENDO, 1961a; Kochanskyella tulipa MILANOVIC, 1974 (Fig. 14.1); Parachaetetes sp. (silicified). incertae sedis: Sphairionia galinae nov. sp. (Fig. 21.2-9); Diplosphaerina inaequalis (DERVILLE, 1931); Pseudovermiporella sodalica ELLIOTT, 1958 (Fig. 21.15); Tubiphytes obscurus MASLOV, 1956 (numerous morphotypes; Fig. 24.9). H133 (Early Midian): Bioclastic and intraclastic rudstone partially silicified with: algae: Archaeolithoporella hidensis ENDO, 1961a; Koivaella sp.; Mizzia sp.; Kochanskyella tulipa MILANOVIC,1974; Parachaetetes sp. incertae sedis: Pseudovermiporella longipora (PRATURLON, 1963) nov. comb. (Fig. 21.16); Tubiphytes obscurus MASLOV,1956 (Fig. 24.8). tt489 (Early Midian): Bioclastic rudstone with: algae: Mizzia yabei (KARPINSKY,1908). incertae sedis: Eflugelia johnsoni (FL~GEL, 1966) VACHARDin Massa & Vachard, 1979; Chuvashovia densifolia VACHARDin Vachard & Montenat, 1981 (Fig. 18.9); Sphairionia galinae nov. sp. (Fig. 21.6, 8); Diplosphaerina inaequalis (DERVILLE, 1931);

381

Pseudovermiporella sodalica ELLIOTT, 1958; Tubiphytes obscurus MASLOV, 1956 (numerous morphotypes). H541 (Early Midian): Bioclastic, intraclastic and partially silicified rudstone with: algae: Girvanella sp.; Anthracoporella spectabilis PIA, 1920; Diplopora sp.; Gymnocodium exile Mu, 1981 (Fig. 10.2-3). incertae sedis: Evlaniopsis montana VACHARD in Montenat & Vachard, 1981; Claracrusta catenoides (HoMANN, 1972) VACHARD,1980; C. hirtipes VACHARD, 1980; Claracrusta sp.; Sphairionia galinae nov. sp. (Fig. 21.3-5, 7); S. sikuoides TIEN, 1989 non 1988 (Fig. 21.1); Diplosphaerina inaequalis (DERVILLE, 1931) (Fig. 21.10); Pseudovermiporella sodalica ELLIOTT, 1958; Tubiphytes obscurus MASLOV,1956. H250 (Late Midian): Algal bindstone with: algae: Archaeolithoporella hidensis ENDO, 1961a; Sparaphralysia orientalis nov. sp. (Fig. 10.1; Fig. 24.2); Bacinella sp.; Thaiporella aff. uralica CHUVASHOV,1974 (Fig. 10.11); Anthracoporella spectabilis PIA, 1920 (Fig. 12.1-9); Kochanskyella (?) sp.; Archaeolithophyllum missouriense JOHNSON, 1956 (Fig. 10.1; Fig. 12.6; Fig. 14.8-10; Fig. 24.2; in process of sparitization); Aseelahella granieri nov. gen. nov. sp. (Fig. 17.2-3). incertae sedis: Claracrusta catenoides (HoMANN, 1972) VACHARD,1980; C. hirtipes VACHARD, 1980; Evlaniopsis montana VACHARDin Vachard & Montenat, 1981; Nostocites vesiculosa MASLOV, 1929 emend. (Fig. 18.1); Tuberitina collosa REITLINGER, 1950; Pseudovermiporella elliotti ERK & BILG~TAY, 1960; Tubiphytes obscurus MASLOV, 1956 (Fig. 24.1, 3-4).

incertae sedis: Stacheoides sp.; Eflugelia johnsoni (FLOGEL, 1966) VACHARDin Massa & Vachard, 1979; Evlaniopsis montana VACHARDin Vachard & Montenat, 1981 (Fig. 18.8); Claracrusta sp.; Diplosphaerina inaequalis (DERVILLE,1931); Tubiphytes obscurus MASLOV,1956.

PALEOECOLOGY Three algal communities are present (Fig. 7): 1) The Kubergandian community of Aseelahella nov. gen., Dasycladales and Tabiphytes, probably accumulated as proximal tempestites. 2) The communities with dominant fusulinids and subordinate algae (Yakhtashian, Bolorian, Early Murgabian, Early Midian, Dzhulfian). 3) The community with Anthracoporella and Archaeolithophyllum, which constitutes a bindstone/baffiestone in the toe of slope, or boundary of the platform, in the Late Midian, because of its association with red wackestones with abundant polyaxone sponge spicules and radiolaria (compare with the basin-facies described off Chinese Permian reefs by Fan et al. 1990; Liu & Rigby 1992). Similar 'micromounds', built by the same algae, were accurately studied in the Early Permian of the Carnic Alps (Krainer 1995). According to Samankassou (1998), the build-ups with Anthracoporella in the Carnic Alps 'occurred below storm wave base, in a quiet environment', contrary to the lagoonal interpretation of Flfigel (1979).

H245 (Dzhulfian): Bioclastic neosparitized packstone without alga.

H256 (Dzhulfian): Bioclastic rudstone/floatstone with: algae: Mellporellopsis corpulenta nov. gen. nov. sp.; Diploporaceae undeterminated.

•

®

®

2

3

4

5

Division CYANOBACTERIA Family APHRALYSIACEAEnov. fam. S y n o n y m s - Wetheredellaceae sensu BERCHENKOin Chuvashov et al. 1987 non VACHARDin Dil et al., 1977; Wetheredella Group sensu Riding, 1991, pp. 72-73 (pars); Lithocodium/ Bacinella Group sensu Riding, 1991, pp. 73-74 (pars).

Diagnosis - Encrusting colonies, nodular, compo-

©

1

SYSTEMATICS (D. Vachard)

6

7

8

sed of dark microgranular elements ('walls'): vesicles, tubules, laminae, networks, etc., filled with clear microsparitic elements. Some 'walls' can be also microsparitic. 9

10

FIGURE 7 - Schematic diagram of environments with algae in the Permian of the Batain Plain. A-C: algae-bearing sediments; A: proximal 'tempestites'; B: wackestones/packstones with algae and foraminifers; C: relatively deep mud-mounds adjacent to basin deposits. 1: hypothetical paleotopography and basement; 2: rudstones; 3: fusulinids; 4: Anthracoporella; 5: resedimented Aseelahella; 6: Archaeolithophyllum; 7: Aseelahella in situ; 8: radiolaria; 9: polyaxone spicules; 10: ammonoids. Reconstitution des environnements ~talgues du Permien de la Batain Plain. A-C: sddiments & algues; A: "tempestites' proximales; B: wackestones/packstones ~t algues et foraminif~res; C: mud-mounds relativement profonds passant ~t des ddp6ts de bassin. 1: paldotopographie suppos~e et substrat; 2: rudstones; 3: fusulines; 4: Anthracoporella; 5: Aseelahella resddimentdes; 6: Archaeolithophyllum; 7: Aseelahella in situ; 8: radiolaires; 9: spicules polyaxones; 10: ammono~des.

Composition -Aphralysia GARWOOD,1914; Sparaphralysia VACHARDin Vachard & Beckary, 1991; Wetheredella auctoris (pars) non WOOD, 1948; P y c n o p o r i d i u m DERVILLE, 1931; Stylocodium DERVILLE, 1931; Archaeolithoporella ENDO, 1961a; Bacinella RADOtCIC,1959; ? Intextulella PETRYKin Petryk & Mamet, 1972 (Fig. 8).

Comparison - The Wetheredellidae or Wetheredellaceae (incertae sedis) differ from the Aphralysiaceae by a hyaline perforated wall, which is a true exoskeletal element. Wetheredella, probably of foraminiferal affinity (see discussion in Belka 1981, replacing the erroneous name Aphralysia by Wetheredella !), possesses a true calcitic wall, but

382 - Height of colonies up to 1 mm; width up to several mm. Height of arches 0.01 ram; width of arches 0.01-0.02 mm. Dimensions

- ? Early Midian of Tebaga (Tunisia). Late Midian of the Batain Plain (Oman). Range

mH,

Genus A r c h a e o l i t h o p o r e l l a ENDO, 1961a

Archaeolithoporella

Aphralysia

Bacinella FIGURE8 - Some Cyanobacteria Aphralysiaceae nov. fam., showing the calcified parts in black and the uncalcified (empty?) parts in white. A possible phylogeny is suggested by the arrows.

Schgmas de quelques cyanobactgries Aphralysiaceae nov. faro., montrant les parties calcifiges en noir, et les parties non calcifides en blanc. Les fl~ches indiquent une possible phyloggnie.

the i n t e r p r e t a t i o n of Wetheredella by K a z m i e r c z a k & Kempe (1992) can be applied to the Aphralysiaceae (i.e. 'vesicular/tubuloid microbial structures'). - Early Carboniferous (Tournaisian) with Aphralysia to Late Cretaceous with Bacinella; cosmopolite. Ranges

Genus S p a r a p h r a l y s i a VACHARDin Vachard & Beckary, 1991

Type species - Archaeolithoporella hidensis

- We agree with the definition and interpretation of the genus by Razgallah & Vachard (1991). Archaeolithoporella is a crust of Cyanobacteria more or less identical to a stromato]ire (Fliigel 1966, p. 52-53, pl. 9, figs 2-3; Kochansky-Devid6 1970a, p. 210, 238, pl. 20, figs 1-2; Fliigel 1977, pl. 2, fig. 3 and p. 325; Vachard & Montenat 1981; Fontaine et al. 1994, pl. 30, fig. 4; and discussion in Kirkland et al. 1998). Archaeolithoporella and other Aphralysiaceae can be interpreted as endostromatolites (i.e. cavity- or fissure-dwelling stromatolites defined by Monty 1982, 1984, 1986). Wood et al. (1994) have already emphasized the cryptobiont behaviour of Archaeolithoporella in the Capitan Reef, USA. The association of Archaeolithoporella and Tubiphytes is very common and reef-building during the whole Permian (Fliigel 1980, 1981, 1994; Fliigel et al. 1984; Babcock 1986; Fliigel & Reinhardt 1989; Fan et al. 1990; Razgallah & Vachard 1991; Riding & Guo 1991; Dawson & Racey 1993; Wang et al. 1994). - Probably all the Permian (until the latest Changxingian/Dorashamian; see Liu & Rigby 1992).

Range

P h y l u m CHLOROPHYCOPHYTA Class CHLOROPHYCEAE Order CAULERPALES F a m i l y UDOTEACEAEF e l d m a n n , 1946

Composition - Sparaphralysia tacania VACHARDin Vachard & Beckary, 1991; Wetheredella starobeshevense BERCHENKO in A i s e n v e r g et al., 1983; Sparaphralysia orientalis nov. sp.

- N o d u l a r thallus, hemispherical or elongate, composed of skeletal elements in arches. White, finely microsparitic wall. Middle Carboniferous (Vachard & Beckary 1991) to Midian (this study).

Range

-

Sparaphralysia orientalis nov. sp. Fig. 10.1; Fig. 24.2 ? 1964 Osagia ? - Glintzboeckel & Rabat~, pl. 72, figs 1-2 ; pl. 73, fig. 1, pl. 77, fig. 1. ? 1991 'Osagia' - Vachard & Beckary, p. 322 (not illustrated). ? 1993 Grains ~ crofite d'Aphralysia - Vachard & Razgallah, pl. 1, figs 1-3, 5-6. Derivatio

nominis

- From the Middle East.

Holotype - Fig. 10.1 (detail Fig. 24.2). Material

-

Ten colonies.

Type locality - Wadi Hasad, 10 km WSW of the village Sal ~(UTMcoord. 759068/2444367), Batain Plain, Oman. Type level - Late Midian (Middle Permian; probable equivalent of the Late Capitanian). Diagnosis - Colonies with n u m e r o u s arches layers, incrusting various supports, generally Anthracopo-

rella. - C o l u m n a r g r o w t h as in the case of some stromatolites; each skeletal series parabolic in section, and composed of t w e n t y to fifty arches of individually mineralised algae; wall very thin and microsparitiC. O m a n e s e colonies are poorly developed, those of the Tebaga (see synonymy) are more i m p o r t a n t in size and n u m b e r of arches, but possibly belong to the same species. Description

1961a.

Remarks

Type species - Sparaphralysia tacania VACHARDin Vachard & Beckary, 1991.

Diagnosis

ENDO,

Genus M e l l p o r e l l o p s i s nov. gen. Type species - Mellporellopsis corpulenta nov. gen. nov. sp. Derivatio

nominis

-

Similar to Mellporella RACZ,1965.

- S e g m e n t e d Udoteaceae, with simple elongate thallus; medullar zone poorly calcified constituted by zigzagging relatively big tubes; cortical zone with n u m e r o u s , m e d i u m to fine t h r e a d s , dichotomous, r a n d o m l y disposed (Fig. 9A); white microsparitic wall. Diagnosis

Comparisons - The cortical zone presents some resemblances with the genus Mellporella, generally i n t e r p r e t e d as a dasycladale alga. According to Vachard & Beckary (1991) the algal f r a g m e n t s of the B e c h a r B a s i n (Algeria), d i s t i n g u i s h e d by various a u t h o r s u n d e r the n a m e of Mellporella anthracoporellaeformis RACZ, 1965 are also very similar to the non-dasyclad species A n c h i c o d i u m m a g n u m ENDO, 1951. Mellporellopsis differs from 'Mellporella' or Anchicodium by the more complex stucture of the cortical area. Affinities are also with Boueina, H a l i m e d a and Arabicodium (see discussions about these three genera in Elliott 1957, 1965; in Bassoullet et al. 1983), b u t the difference is in the irregular cortical threads. According to Fliigel (1988) Halimeda-like types a p p e a r massively in the Late Triassic, but the P e r m i a n Siamporid i u m ENDO, 1969, can be s y n o n y m o u s of Arabicod i u m . F u r t h e r m o r e Halimeda soltanensis PONCET, 1989, is a 'phylloid alga', not a true H a l i m e d a , comparable to A n c h i c o d i u m funile JOHNSON, 1946, illust r a t e d by Torres & B a a r s (1992, fig. 1.4), to A. robustum Mu, 1982 or A. zhougbaensis Mu, 1982. Range Omau.

- Yakhtashian and Dzhulfian of the Batain Plain,

383 FIGURE 9 - Anatomy of Mellporellopsis nov. gen. (9A) and 'Thaiporella' (9B). Skeletal part in white; stippled emplacements of filaments. Anatomie de Mellporellopsis nov. gen. (9A) et ~haiporella' (9B). Les parties squelettiques sont laissdes en blanc et le moulage des filaments est en pointilld. Sans dchelle.

not congeneric with the Endo material; and the original microphotograph of T kobayashi does not correspond to the reconstruction (Endo 1966, text fig. 1, p. 171). Some misinterpreted sections of Permian algae possess other similarities with this taxon: Ortonella morikawai ENDO, 1958; Ortonella latifibrosa ENDO in Endo & Kanuma, 1954; Salopekiella cf. velebitana MILANOVIC, 1965 sensu Homann, 1972 (p. 230-231, pl. 7, figs 56-58); Salopekiella (?) sp. sensu Mu, 1982 (p. 230, pl. 4, fig. 7). - Unknown in Thailand. Asselian (Early Permian) in Urals. Kubergandian and Late Midian in Oman (Batain Plain). Range

T h a i p o r e l l a aff. u r a l i c a CHUVASHOV, 1 9 7 4 Fig. 10.10-11

N

? 1974 Thaiporella uralica - Chuvashov, p. 19-20, pl. 5, figs 1-3. ? 1993 ThaiporeUa uralica - Chuvashov et al., pl. 14, fig. 14 (not described). R e m a r k s - Two thalli, measuring 0.96 × 0.80 mm and 1.90 x 1.80 mm, show a set of laterals diverging from a basal and central point. The microsparite of the thallus and the microgranular filling of the filaments are typical of the Caulerpales or the Dasycladales. Range

-

Kubergandian and Late Midian of Oman (Batain

Plain). O r d e r DASYCLADALES M e l l p o r e l l o p s i s c o r p u l e n t a nov. sp. Fig. 10.47, 5-9 D e r i v a t i o n o m i n i s - From the Latin corpulentus, fat. H o l o t y p e - Fig. 10.5. P a r a t y p e s - Fig. 10.6-9. T y p e m a t e r i a l - Thin section H253. O t h e r m a t e r i a l - Thin sections H233 and H256. M a t e r i a l - 12 specimens. T y p e l o c a l i t y - Wadi Hasad, 10 km WSW of the village Sal (UTM coord. 759068/2444367), Batain Plain, Oman. T y p e l e v e l - Yakhtashian. Diagnosis

- A s for t h e m o n o s p e c i f i c g e n u s .

Description - Thallus cylindrical, unbranched; medullar zone weakly calcified, with badly conspicuous interweaving threads of relatively large diameter; cortical area perforated by numerous thin, acrophore, dichotomous threads, randomly dispos e d (Fig. 9A). D i m e n s i o n s - Length (1.25?)-1.70-1.80 mm; diameter 2.002.20 (2.80?) mm; fragment thickness 0.25-0.50 mm; average thread diameter 0.03 mm. - Yakhtashian (H233)-Dzhulfian (H256); Yakhtashian reworked in the Dzhulfian (H253) of Oman.

R e m a r k s - The nomenclature used here is that of De Castro (1997), with the following precision: bifurcation in the absolute sense corresponds to a dichotomy of the thallus; ramification is only used for the dichotomy of the laterals (= branchlets = ramifications sensu stricto = pores sensu lato). Therefore a Dasycladale shows a thallus composed of a large central cell with a calcified cortex. According to the shape of the central cell, the thallus is cylindrical, bifurcate, clavate, moniliform (in collar of pearls), spherical, etc. From this central cavity, arise laterals, which are acrophore (i.e. cylindrical), phloiophore (i.e. acrophore tapering), vesicular (acrophore becoming spherical), etc. in shape. The calcification of the laterals is closely associated with that of the central cavity, or independent. These laterals are grouped by planes called verticilles. If the verticilles are simple, the arrangement is euspondyle; if the euspondyle basal segments (sometimes called vestibules) give rise to a tuft of laterals, the verticille is metaspondyle. If not any verticille is individualised the pattern is aspondyle (atypical for a Dasycladale). The family Seletonellaceae is aspondyle; Dasycladaceae are euspondy]e and Diploporaceae metaspondyle. The mode of reproduction is (1) endospore (spores within the axial cavity), and in this case all the laterals are sterile; (2) cladospore (spores in specialized laterals, generally vesicular); or (3) choristospore (with specialized fertile verticilles generally disc-shaped, with elements called spicules or rays). The choristospore Dasycladales, at least assumed if fossil, constitute the family Acetabulariaceae.

Range

G e n u s ' T h a i p o r e l l a ' ENDO, 1 9 6 6 s e n s u CHUVASHOV, 1 9 7 4 T y p e s p e c i e s - Thaiporella kobayashi ENDO,1966. Description - Thallus probably segmented, constituted by a bundle of radial divergent phloiophore ramifications, apparently not bifurcated. The ramifications are not in a single plane, and thicken toward the ends, terminating in small funnels (Fig. 9B). R e m a r k s - Our specimens are too rare for emendating the genus, but it is clear that the taxon described by Chuvashov is

F a m i l y SELETONELLACEAE K o r d 6 , 1 9 5 0 e m e n d . B a s s o u l l e t e t al., 1 9 7 9 T r i b e DASYPORELLEAE P i a , 1 9 2 0 (?) Description - S e l e t o n e l l a c e a e (i.e. a s p o n d y l e D a sycladale) cylindrical or bifurcated, with numerous acrophore laterals, generally single, sometimes ramified; wall white microsparitic, laterals filled with a micritic cement. R e m a r k s - Anthracoporella is generally included in the tribe Dasyporelleae, and possibly in the subtribe Dasyporellineae (Pia 1920; Shuysky 1973; Bassoullet et al. 1979; Roux 1985; Mamet et al. 1987; Chuvashov et al. 1987; Deloffre 1988), but for three reasons this classification is questionable: (1)

384

1--

2

4

3

5

.,

6

8

FIGURE 10 - 1. Sparaphralysia orientalis nov. sp. Holotype. A thallus encrusting an Anthracoporella and itself encrusted by Archaeolithophyllum missouriense. Late Midian. Batain Plain. H250(2)/924(3). For detail, see Fig. 16: 2. 2-3. Gymnocodium exile Mu, 1981. Early Midian. Batain Plain. 2: Oblique section. H541(1)/924(21). 3: Tangential section. H 541 (7)/924 (27). 4?, 5-9. Mellporellopsis corpulenta nov. gen. nov. sp, (4 is Yakhtashian in age; 5 to 9 are Yakhtashian reworked in the Dzhulfian, as evidenced by the Leeina kraffti and Codonofusiella schubertellinoides contained in the same levels; see Hauser et al., 2000). Batain Plain. 4?: Longitudinal section partly dolomitized. H 233(2)/912(3). 5: Holotype. Longitudinal section showing the general shape and the different types of canalicules. H 253(3)/911(3). 6: Paratype. A fragment of cortical zone. H253(4)/911(4). 7: Paratype. Other fragment. H 253(2)/911(2). 8: Paratype. Subaxial section showing the mode of bifurcation. H 253(7)/911(7). 9: Paratype. Tangential section. H253(8)/911(8). 10-11. Thaiporella aff. uralica CHUVASHOV,1974. 10: Longitudinal section. H240C(2)/912(16). 11: Oblique section. H250B(1)/909(13).

385 Anthracoporella differs from other Dasyporelleae by the bifurcations of the thalli; (2) the laterals of the members of the Dasyporelleae are never ramified and entirely simple and acrophore, contrary to Anthracoporella; (3) the eponym genus Dasyporella STOLLEY,1893, is doubtful as part of its material belongs to the red alga Californiella PONCET, 1987a. De Castro (1997, p. 142) assumed t h a t Anthracoporella is not a Dasycladale, probably because of its morphological resemblance with the Vermiporellaceae, which may be Ulotrichales (according to Kozlowski & Kazmierczak 1968; Emberger 1976; Bassoullet et al. 1979).

Genus Anthracoporella PIA, 1920 T y p e - s p e c i e s -Anthracoporella spectabilis PIA, 1920.

S y n o n y m - ?Couvinianella MAMET& PREAT,1992.

D i a g n o s i s - Large and cylindrical, often bifurcated thallus; numerous simple laterals, aspondyle, acrophore or slightly phliophore, sometimes peculiarly dichotomous ('dichotomous branching' of De Castro 1997, p. 142), or really ramified one time (Fig. 11); yellowish or white, microsparitic wall. Remarks - This well known genus (see Pia 1920; KochanskyDevid6 & H e r a k 1960; Elliott 1968; Chuvashov 1974; Emberger 1976, etc.) is also interesting as a reef-builder (Fliigel 1987; Krainer 1995; Samankassou 1997, 1998). R a n g e - Silurian (Shuysky 1973)?; Late Emsian (Early Devonian) to Couvinian (Middle Devonian) (Shuysky 1973; Saltovskaya 1984; Shuysky & Patrunov 1991; Mamet & Pr6at 1992); Middle Carboniferous (Late Bashkirian of Spain, Racz 1965), Moscovian (Racz 1966; Kochansky-Devid6 1970b; Emberger 1976; Delvolv6 et al. 1987) to Middle Permian (Midian) in Malaysia (Fontaine et al. 1988a), in Afghanistan (Vachard 1980), in Oman (Montenat et al. 1976; this study), and possibly Dzhulfian in Albania (Bignot et al. 1982). Despite the gap between the Early Devonian and the Early Carboniferous (the species attributed to the genus during this period are misinterpreted; see below), no difference between the Devonian species described by Shuysky (1973) and typical Anthracoporella is noticeable.

C o m p o s i t i o n - The genus is composed by two groups of species: (1) the Devonian species with Anthracoporella atypica SALTOVSKAYA,1984; A. delicata SALTOVSKAYA,1984; A. expansa SHUYSKY in Shuysky & Patrunov, 1991; A. laevis SHUYSKY,1973; A. longiramosa SH~YSKYin Shuysky & Patrunov, 1991; A. setosa SHUYSr~Z,1973; ?Couvinianella sartenarii MAMET& PREAT, 1992 (if generically diffe-

rent, this first group must be called Couvinianella); (2) the Late Carboniferous-Early Permian species A. spectabilis PIA, 1920, A. uralensis CHUVASHOV, 1974 and A. vicina KOCHANSKY-DEVID~& HERAK, 1960. But our observations in the Carnic Alps (Vachard and Argyriadis, in press) reveal that A. spectabilis is extremely polymorph and these socalled species could be synonymous. The following species do not belong to Anthracoporella (see also Vachard 1993): A. kasachiensis MASLOV, 1939 and A. fragilissima MASLOV, 1939 (two Nanopora, according to Wood 1964; Mamet & Roux 1975; Emberger 1976); A. magnipora ENDO, 1951, A. torinosensis ENDO, 1961b and A. menchikoffi CHANTON,1965; they correspond to other genera of Dasycladales. A. mercurii ELLIOTT,1968 is probabty a fragment of Pseudovermiporella (i.e. a foraminifer); A. baschkirica KULIK, 1973 probably belongs to the Vermiporellacae; A. girtyi MAMET& Roux in Mamet et al., 1987 is an Anthracoporellopsid pseudo-alga, as well as Anthracoporella sp. (Mamet & Roux 1975, p. 264, pl. 13, fig. 12); A. insolita IVANOVA,1988 probably belongs to the genus Roquesselsia TERMIERet a1.,1977.

Anthracoporella spectabilis PIA, 1920 Fig. 12.1-10

S y n o n y m y - See authors (Pia 1920, 1937; Kochansky-Devid6 & Herak 1960; Fliigel 1966; Homann 1972; Chuvashov 1974; Emberger 1976; Fliigel & Flfigel-Kahler 1980; Granier & Grgasovic 2000). Some diversely attributed algal fragments certainly belong to Anthracoporella: Epimastopora kanumai ENDO in Endo & Kanuma, 1954 (p. 195, pl. 13, figs 8-10); Chuvashov 1974 (p. 16, pl. 1, figs 1-4); Mu 1982 (p. 225, pl. 5, fig. 6); etc.; Epimastopora urtazymensis CHUVASHOV& ANFIMOV, 1988 (p. 60-61, pl. 23, figs 3-7); Anchicodium sindbadi ELLIOTT, 1970 (p. 327-328, pl. 61, figs 3-4); Parapimastopora sp. in Chun & Mamet 1993 (Fig. 4E). Remarks - Our material exhibits typical and atypical forms (Figs 11, 12), with sometimes a thin external finely perforated calcareous layer adjoining the surfaces of the laterals, which was first observed by Pia (1920) in the Carnic Alps; these atypical forms look like Diplopora americana KORDE, 1965. The synonymy of this species with A. spectabilis is possible. D i m e n s i o n s - Length up to 10 ram, outer diameter 0.60-2.50 mm; inner diameter 0~25-1.40 mm, thallus thickness 0.20-0.50 ram, pore diameter 0.01-0.02 mm.

B i o s e d i m e n t o l o g y - Samankassou (1997) indicates that Anthracoporella can build massive reef mounds during the late period of the transgression, prior to the maximum flooding surface, or in an early highstand system tract (Krainer 1995).

atypical Anthracoporetla FIGURE 11 - Comparison of the typical and atypical Anthracoporella of the Late Permian from the Batain Plain. Schdmas comparatifs des Anthracoporella typi-

ques (A) et atypiques (B) du Permien supdrieur de la Batain Plain.

D i s t r i b u t i o n - Common in: Carnic Alps of Austria and Italy (Pia 1920; Homann 1972; Fltigel & FlfigelKahler 1980; Krainer 1991, 1995; etc.); Karawanken (Slovenia); Lika and Velebit (Croatia); Montenegro (Kochansky-Devid6 & Herak 1960); Serbia (Filipovic 1995); Greece: Chios Island (Pia 1920; Renz & Reichel 1945, p. 230) and Attica (Vachard et al. 1993); Iran? (Douglas 1950); Turkey (Teke dere, Lycian Nappes: Argyriadis et al. 1976); Irak; Bukk Mountains (Hungary); Madagascar ? (discussion in Pia 1937); Shakhtau Massif (Urals, Russia: Kulik 1978); Darvaz (Saltovskaya 1984); Japan (Kitakami, Kwanto: compilation by Emberger 1976; Kyushu: Konishi 1954, as Macroporella

386

m

o9

co

7

FIGURE 1 2 - 1-10. Anthracoporella spectabilis PLY, 1920. Late Midian. Batain Plain. 1: Bifurcated typical thallus. H250(2)/909(23). 2: Opposite size of the former specimen encrusted by Sparaphralysia orientalis nov. sp. H250(2)/909(22). 3: Various transverse sections showing the typical aspect (in center) and atypical ones (right and left), more 'diploporid', all encrusted by Archaeolithoporella. H250(9)/909(33). 4: Bifurcated thallus. H250B(2)/909(14). 5: Transverse section with a Schubertellid trapped in the axial cavity. H250(1)/909(21). 6: Transverse section withArchaeolithophyllum in a cyanobacterial crust H250B(3)/909(15). 7: Detail of 6, showing the ramified lateral and the perforated cortex. H250B(3)/909(16). 8: Tangential section. H250/922(31). 9: Oblique section. H250(2)/924(8). 10: Transverse section. H180(2)/914(16).

387 sp., pl. 2, fig. 18; Honshu: H a n z a w a 1961, as Macroporella sp., pl. 71, fig. 1); China a n d Xizang (Mu 1982); U S A (New-Mexico, Texas, Missouri: Bebout & Coogan 1964); C a n t a b r i c M o u n t a i n s (Spain: Racz 1966); P y r e n e e s (France: Delvolv~ et al. 1987); Thail a n d (Endo 1969; F o n t a i n e & S u t e e t h o r n 1988); M a l a y s i a (Fontaine et al. 1988); Viet N a m and Laos (Tu 1970); C a m b o d i a (Tien 1979); S u m a t r a (Font a i n e & Vachard 1984; F o n t a i n e & Gafoer 1989, not u n d e r this n a m e , but as Permocalculus fragilis, m i s i n t e r p r e t e d , pl. 24, fig. 5; pl. 31, figs 1-2); A f g h a n i s t a n (Vachard 1980); O m a n (Elliott 1968; Vachard in M o n t e n a t et al. 1976; a n d this study). The range is mainly Late Carboniferous-Early Permian in the Western Tethys (Emberger 1976). Midian outcrops are situated in Croatia (Kochansky-Devid~ & Herak 1960), in Slovenia (Fliigel et al. 1984), in Afghanistan (Vachard 1980) and in Oman (Montenat et al. 1976; and this study). Range -

F a m i l y ACETABULARIACEAEs e n s u Deloffre, 1987 Tribe LULIPOREAE S h u y s k y in C h u v a s h o v et al., 1987 emend. Definition - D a s y c l a d a l e s w i t h spaced complex verticilles composed of simple l a t e r a l s (L1), or r a r e ly of first a n d second orders (L1 a n d L2); l a t e r a l s

elongate, cylindrical or w e a k l y clavate; wall white microsparitic, typical of the order. - Differs f r o m Clypeineae Elliott, 1968 emend. B a s s o u l l e t et al., 1979 s e n s u Chuv a s h o v et al., 1987 (pars) by the g r e a t e r complexity of verticilles, w i t h l a t e r a l s c o m m u n i c a t i n g w i t h the exterior; b e c a u s e of this character, Luliporeae are morphologically i n t e r m e d i a t e b e t w e e n t r u e Dasycladaceae a n d t r u e A c e t a b u l a r i a c e a e . Comparison

(Figs 13-14 pars) - Lulipora SHUYSKY, 1986; Lopsiella SHUYSKY, 1987; Coticula SHUYSKY& SCHIRSCHOVAin C h u v a s h o v et al., 1987; Gissarella SALTOVSKAYA, 1979; Connexia KOCHANSKY-DEVIDI~, 1970b; Likanella MILANOVIC, 1966; Salopekiella MILANOVIC, 1965; Kochanskyella MILANOVIC, 1974; Oligoporella s e n s u ENDO, 1969 non PIA, 1912 (pars); Clavaphysoporella s e n s u ENDO, 1969 non Endo, 1958 (pars). Composition

- This tribe possesses the typical morphology of the family but the laterals are not still specialized for the reproduction. The members are euspondyle and not metaspondyle as the Diploporaceae. The true Clypeineae appear probably in the Early Midian with Eoclypeina EMBERGERin VACHARD,1985, p. 273, not EocIypeina PONCET,1987b; not only this genus is preoccupied (Granier & Grgasovic 2000), but its type species, E. crassa PONCET,1987b, is a probably synonymous of Clavaporella reinae RACZ,1966. Remarks

Emsian/Elikhovian to Givetian (Early-Middle Devonian) from Urals (Russia). Moscovian from Croatia (KochanskyDevid6 1970b) and Middle Carboniferous of Tadzhikistan (Saltovskaya 1979). Early Permian from the Carnic Alps, Tunisia, Xizang, Sumatra, Guatemala (Vachard et al. 1997, p. 758). Kubergandian-Midian (Middle Permian) from Croatia, Tunisia, Japan, Afghanistan (Vachard 1980). Discovered in the Middle Permian (Kubergandian-Midian) of the Batain Plain, Oman. Range -

Ciype(na

(Triassic-Oligocene)

Eoclypeina (Midian)

J

G e n u s K o c h a n s k y e l l a MILANOVIC, 1974 Type-species - Kochanskyella tulipa MILANOVIC,1974.

Kochanskyella

(Kubergandian-Midian)

~

u

Likanolla

bergandian-Murgabian)

Diagnosis - Thallus composed of loosely connected bowl-shaped segments; s e g m e n t s composed of loosely connected petal-like parts; each p a r t h a s a syst e m of pores, and corresponds to the calcification of an euspondyle (metaspondyle?) verticille with laterals incurved a n d strongly inclined; wall microsparitic. R e m a r k s - This beautiful genus is little known but apparently important. The resemblance between one of the first Atractyliopsis published by Pia (1937), and the transverse sections of petal of Kochanskyella, is evident. The genus 'Atractyliopsis' sensu Mamet & Roux (1975) seems to be different of the taxon of Pia, but paradoxically corresponds more to its definition (i.e. a calcified central stem of an endospore Dasycladale, whose laterals are not mineralized).

The tribal assignment of Kochanskyella is very discussed: questionable Salpingoporelleae for Bassoullet et al. (1979); true Salpingoporelleae for Chuvashov et al. (1987); Diploporaceae Albertaporelleae for Giivenc (1979, p. 634) and Deloffre (1988, p. 173); Columbiaporelleae for Granier & Deloffre (1994).

(Moscovian-earlyPermian)

- Middle Permian (Kubergandian to Murgabian) Croatia, Afghanistan. Discovered in the Middle Permian of the Batain Plain, Oman. Range

Mas/oviporella

(earlyCarboniferous) FmURE 13 - Comparative scheme of Acetabulariacean Dasyclad algae. Schgmas comparatifs et possible phylogdnie de certaines

dasycladales acdtabulaires.

Kochanskyella tulipa MILANOVIC, 1974 Fig. 14.1-4 1937 Atractyliopsis - Pia, pl. 10, fig. 6 (only).

388

¢)

8 ¢D

0

O3

3

5

8--

6

I4

7

9--

10

FIGURE 1 4 - 1-4. Kochanskyella tulipa MILANOVIC,1974. Kubergandian. Batain Plain. 1: Transversal section of a petal. H132(3)/907(26). 2: Transversal section of a petal (center) with Diplopora sp. (above). H241(2A)/923(15). 3: Longitudinal section of a petal. H241(2)/910(20). 4: Oblique section of a petal. H241(1A)/923(5). 5. Likanella (?) sp. A fragment of verticille in longitudinal section. Kubergandian. B a t a i n Plain. H241(3)/910(21). 6-7. Diplopora spp. 6: Transverse section. Kubergandian. B a t a i n Plain. H240(B4)/912(12). 7: Oblique section. Dzhulfian. Batain Plain. H253(6)/911(6). 8-10. Archaeolithophyllum missouriense JOHNSON, 1956. Late Midian. Batain Plain. 8: Longitudinal section of several thalli with shelter porosity. H 250 (3)/924(5). 9: Constituting a bindstone with Archaeolithoporella. H 250(3)/924(4). 10: A diagenetically transformed thallus tending to Kansaphyllum or 'Halimeda' of the authors. H250(7)/909(31).

389 1974 Kochanskyella tulipa - Milanovic, p. 128-130, pl. 1, figs 1-4, pl. 2, figs 1-5, pl. 3, figs 1-9, pl. 4, figs 1-6, pl. 5, figs 1-5. 1980 Kochanskyella tulipa - Vachard, pp. 356-357, pl. 23, fig. 11. 1981 Kochanskyella tulipa -oVachard & Montenat, pp. 39, pl. 5, figs 10, 15. 1994 Kochanskyella tulipa - Granier & Deloffre, p. 53, 68 (not illustrated). 1997 Kochanskyella tulipa - De Castro, pp. 67, 161 (not illustrated). 2000 Kochanskyella tulipa - Granier & Grgasovic, p. 90 (not illustrated, with synonymy). r,'~\ Description

-

A few fragments, all illustrated here.

- Length of petals 1.16-2.50 mm; width of petals 0.40-0.60 mm, diameter of laterals 0.08-0.15 mm, interpores 0.02-0.04 mm.

Dimensions

- This alga is associated with Likanella sp. (Fig. 14.5), as in Croatia.

Remarks

R a n g e - Middle Permian of Croatia (Milanovic 1974). Kuber-

gandian-Murgabian of Afghanistan (Vachard 1980). Middle to Late Permian, Bellerophon Limestone from the Dolomites (Pia 1937). Discovered in the Middle Permian of the Batain Plain, Oman.

Tribe DIPLOPOREAEPia, 1920

Diplopora spp. Fig. 14.6-7 R e m a r k s - These tribe and this genus are characterized by metaspondyle verticilles. Various incomplete sections have been observed from the Kubergandian to the Dzhulfian of the Batain Plain, Oman. Division

RHODOPHYCOPHYTA

Order CRYPTONEMIALES Family ARCHAEOLITHOPHYLLACEAEChuvashov in Chuvashov et al., 1987 emend. - Thallus phylloid or erected, segmented, sometimes ramose and attached; millimetric dimensions; internal tissue differentiated into an important central hypothallus with great cells, and a thin lateral perithallus with small cells; reproductive organs subconical to ovoid, conceptacles with a single atypical aperture; wall calcitic, yellowish or white, probably originally aragonitic. Diagnosis

R e m a r k s - Many 'phylloid algae' appear as the result of the complete sparitization of thalli of Archaeoliphy]laceae, and subsequent microborings by endolithic cyanobacteria or algae, with consecutive micritic filling (Vachard et al. 1989; this study, Fig. 15); this observation has been confirmed in cathodoluminescence by Dawson (1992) and Moshier & Kirkland (1993). The aragonitic composition of the original calcification was established by Kirkland et al. (1991, 1993); despite of that, the classical views (Roux 1985; Mamet et al. 1987; Baars & Torres 1991, etc.) about phylloid algae are still accepted. ArchaeolithophyUum represents, with the Gymnocodiaceae, the only bona fide red algae of the Palaeozoic. Other 'red algae' must be assigned to various incertae sedis groups: Solenoporaceae (see discussion on the affinities of this group in Vachard et al. 1978, with Solenopora, Pseudosolenopora and Parachaetetes), Aoujgaliid pseudo-algae (Ungdarella, Stacheia, Fourstonella, etc.); Stromatoporoidea (Keega); vascular plants ? (Katavella and Lysvaella); etc. Furthermore, the relationship between Archaeolithophyllum and the Peyssonnelliaceae is admitted by various authors (James et al. 1988, Vachard et al. 1989; Dawson 1992; Moshier & Kirkland 1993). In this case the family Archaeolithophyllaceae could be invalid.

FIGURE 15 - Schematic diagram showing of an Archaeolithophyllum passing to a phylloid algae. Schema th~orique montrant le passage des Archaeolithophyllum vrais aux algues dites phyllo~des.

Composition

-

Archaeolithophyllum

JOHNSON,

1956; Principia BRENCKLEin Brenckle et al., 1982; Aseelahella nov. gen.; Eugonophyllum KONISHI WRAY, 1961; Ivanovia KHVOROVA,1946; Kansaphyllure BAARS,1992; ? Anchicodium JOHNSON,1946.

Kansaphyllum, Ivanovia and Eugonophyllum exhibit the phylloid pattern of Archaeolithophyllum, but Anchicodium can be cylindrical (Johnson 1946; Roux 1985; Torres & Baars 1992). Neoanchicodium ENDO in Endo & Kanuma, 1954, sometimes included in the phylloid algae, is clearly different in shape. The genus Amorfia RAcz, 1965, assigned to the family Archaeolithophyllaceae by Chuvashov et al. (1987, p. 131) is an aoujgaliid pseudo-alga, possibly synonymous of FourstoneUa, without relationship with the Archaeolithophyllaceae emend, herein. - Early Carboniferous to Middle Permian (Late Midian). Principia appears in the latest Visean of Europe and Northern Africa (Vachard & Berkli 1992), maybe earlier in USA (i.e. Keokuk Limestone; Brenckle et al. 1982). Archaeolithophyllum appears in the earliest Middle Carboniferous (Early Bashkirian or Feninsky of Ukraine; Vachard & Maslo 1996, p. 361); it is cosmopolite in the Late Carboniferous-Early Permian, and known until the Early Midian in Tunisia (Vachard et al. 1989); and the Late Midian in Oman (this study). Aseelahella is Kubergandian and Late Midian in Oman. Range

Genus Archaeolithophyllum JOHNSON,1956 Type s p e c i e s - Archaeolithophyllum missouriense JOHNSON, 1956.

Archaeolithophyllum missouriense JOHNSON,1956 Fig. 10.1; Fig.12.6; Fig. 14.8-10 1956 Archaeolithophyllum missouriensum - Johnson, p. 54, p. 14, figs 1, 3, 5-6. 1972 Archaeolithophyllum missouriensum - Homann, pp. 163164, pl. 1, fig. 6 (with synonymy). 1976 Archaeolithophyllum missouriensum - Emberger, p. 99 (not illustrated; with synonymy). 1987 Archaeolithophyllum missouriense - Mamet, Roux & Nassichuk, pp. 54-55, pl. 28, figs 3-4, pl. 29, figs 1-2 (with synonymy). 1991 Archaeolithophyllum missouriense - Vachard & Beckary, p. 324, pl. 1, fig. 11 (with synonymy). 1991 Archaeolithophyllum missouriense - Krainer, pp. 161, 168 (not illustrated). 1995 Archaeolithophyllum missouriense - Harris et al., figs 9: 11.

390 1995 Archaeolithophyllum missouriense - Krainer, pl. 38, fig. 2, pl. 40, figs 1-2, 4.

- Thallus ribbon-like with central hypothallus composed of large cells, and lateral perithallus with small cells; conceptacles ovoid, laterally emplaced and slightly protruding in the periphery, their single aperture is cylindrical; various stages of diagenesis are conspicuous among these thalli, and are probably at the origin of misinterpreted genera as Anchicodium, EugonophyUum and Kansaphyllum (compare our Fig. 12.10 with the fig. 14 of Torres & Baars 1992, and the fig. 2 of Baars 1992). Description

D i m e n s i o n s - The ribbons attain a centimetric length for a width of 0.30-0.60 ram. The polygonal hypothallus cells measure 0.06-0.08 x 0.03-0.05 mm. R a n g e - Appearance and acme in the Pennsylvanian of USA (Missouri, Illinois, Kansas: compilation in Emberger 1976, p. 99; Alaska: Harris et al. 1995), Spain, Italy, southern Urals, Russia (Rauser-Chernoussova & Koroliuk 1981) and Ukraine (Aisenverg et al. 1983; Vachard & Maslo 1996). Early Permian of USA, New Mexico; Carnic Alps (Krainer 1995). Rare in the Early Midian of Tunisia (Vachard et al. 1989), and in the Late Midian of the Batain Plain, Oman.

Genus Aseelahella nov. gen. T y p e s p e c i e s - AseelaheUa granieri nov. gen. nov. sp.

Derivatio nominis Batain Group.

- After the Aseelah Unit, part of the

- Thallus cylindrical and bifurcated; well developed hypothallus consisting of some polygonal cells randomly arranged; perithallus reduced, difficult to observe, exhibiting small cells which show little or no orientation; perithallus generally completely microsparitized; cell walls white and microsparitic; reproductive structures unknown (Fig. 16). Diagnosis

Composition

- So far monospecific.

- Differs from Archaeolithophyllum and Principia by the bifurcated character of the thallus and the shape of the hypothallus cells; and from Principia also by the white and not dark wall (if not early diagenetic, this criterion can be used for the differentiation of the transverse sections of both genera). Comparison

R a n g e - Discovered in the Kubergandian and Late Midian of the Batain Plain, Oman.

Aseelahella granieri nov. gen. nov. sp. Fig. 17.1-14

D e r i v a t i o n o m i n i s - Dedicated to Dr. Bruno Granier from Total Company, eminent specialist of algae. H o l o t y p e - Fig. 17.1,5.

P a r a t y p e s - Fig. 17.2-4, 6-14. M a t e r i a l - Circa 200 sections. T y p e l o c a l i t y - Wadi Hasad, 10 km WSW of the village Sal (UTM coord. 759068/2444367), Batain Plain, Oman. T y p e l e v e l - Kubergandian.

Diagnosis

- As for the monospecific genus.

- Thallus generally short (segmented?); only one specimen shows the typical bifurcation with a maximal angle of 70 degrees for the branches. Description

D i m e n s i o n s - Thallus: length 0.50-4.50 mm; diameter 0.601.00 mm; cells of hypothallus 0.10-0.30 x 0.30 -0.50 m m with cell walls of 0.01-0.03 mm; cells of the perithallus 0.03-0.04 x 0.04-0.05 mm.

- Two assemblages: one with very numerous fragments in the proximal 'tempestites' of the Kubergandian; the other one in the Late Midian, with rare remains, possibly transported, in the mudmounds with Anthracoporella and ArchaeolithophyUum in the outer platform. Distribution

R a n g e - Only known from the Batain Plain, in the Kubergandian and Late Midian. Some specimens of the Asselian of Austria (Vachard & Krainer, work in progress) may belong to the new genus.

Incertae sedis algae Genus Nostoeites

MASLOV,

1929

T y p e s p e c i e s - Nostocites vesiculosa MASLOV,1929.

- As already summarized by Groves (1983, p. 31), this genus is characterized by a thallus (?) which is a sheet of loosely packed rounded or dolioliform cells (i.e. barrel-shaped); one cell thick, and with dark, subcylindrical inclusions in the center. Wall calcareous, yellowish, hyaline. Diagnosis

Comparison - Differs from Globochaete by the greater number of cells, the depositional environment and the stratigraphic range: Globochaete is Silurian to Cretaceous (see Perret et al. 1994; Skompski 1996, p. 217). R a n g e - Early Visean (Roux 1985) to Late Midian in the Batain Plain, Oman. FIGURE 16 - Reconstruction ofAseelahella granieri nov. gen. nov. sp. Reconstitution d'Aseelahella granieri nov. gen. nov. sp.

Nostocites

v e s i c u l o s a MASLOV, 1 9 2 9

Fig. 18.1

391

4

i Ltt, tL,~,t,,

Scale bar represents 300 I~

8

5

6 ...........

7

9

3 ,,,,,,,,,,

10

11

,,,

12

14

FIGURE 17 - 1-14. Aseelahelta granieri nov. gen. nov. sp. (1, 4-14: Kubergandian; 2-3: Late Midian). Batain Plain. 1: Holotype. Axial section showing the bifurcated thallus. H241C(8)/914(9). 2: Axial section. H250/922(23). 3: Subtransverse section with well preserved hypothallus and conspicuous perithallus. H250/922(24). 4: Paratype. Transverse section showing a perithallus preserved in the cyanobacterial crust. H241B(23)/913(24). 5: Holotype. Detail of the left branch of the holotype (above). H241C(8)/914(8). 6: Paratype. Longitudinal section with partly dissolved cells of hypothallus. H241B(24)/913(25). 7: Paratype. Several transverse sections with polygonal cells. H241B/913(10). 8: Paratype. Transverse section; the perithallus is apparently absent. H241C(9)/914(10). 9: Oblique section with hexagonal-elongate cells in the hypothallus (right) with Claracrusta ungdarelloidea nov. sp. (left). H180(11)/914(25). 10: Paratype. Longitudinal section with a badly visible ramification. H241C(5)/914(5). 11: Pattern of a specimen with altered hypothallus cells H240D(2)/912(25). 12: Paratype. Transverse section with rounded cells in the hypothallus. H241(1)/910(19). 13: Paratype. Oblique section with thin walls. H241C(1)/914(1). 14: Paratype showing polygonal cells. H241B(13)/913(13).

392

3

i 4

T !

7

8

9

FIGUI~E18 - 1. Nostocites vesiculosa MASLOV,1929. Longitudinal section of a colony. Late Midian. Batain Plain. H250/922(28). 2-6. Claracrusta ungdarelloidea nov. sp. Kubergandian. Batain Plain. 2: Longitudinal section very similar to Ungdarella. H241B(14)/913(14). 3: Paratype. Longitudinal section more irregular in shape. H240C(1)/912(15). 4: Holotype. Large longitudinal section perforated by numerous endobionts. H240C(4)/912(21 & 22). 5: Longitudinal section with more regular perforations which can be confused with algal conceptacles. H241C(11)/914(12). 6: Detail of these false-conceptacles. H241C(11)/914(13). 7. Eflugeliajohnsoni (FLOGEL, 1966) VACHARD in Vachard & Massa, 1979. Longitudinal section showing clearly the growth and the skeletal elements. Kubergandian. Batain Plain (Oman). H241-2B/923 (29). 8. Evlaniopsis montana VACHARDin Vachard & Montenat, 1981. A longitudinal section. Dzhulfian. Batain Plain. H256(12)/911(25). 9. Chuvashovia densifolia VACHARDin Vachard & Montenat, 1981. Longitudinal section (under the three black arrows) encrusting a brachiopod shell and Pseudovermiporella tubes. Early Midian. Batain Plain. H489(14)/907(18).

1929 Nostocites vesiculosa - Maslov, p. 1522, 1538, pl. 70, figs 2-10. 1937 Nostocites vesiculosa - Pia, pp. 807-808 (not illustrated). 1982 Globochaete alpina LOMBARD- Skompski, pp. 48-53, pl. 1-3. 1991 Nostocites ex gr. vesiculosa - Vachard & Beckary, pp. 322323, pl. 2, fig. 10 (with synonymy). 1992 Nostocites vesiculosa - Mamet & Pr~at, p. 60, pl. 1, fig. 6. 1996 Globochaete alpina - Skompski, pl. 16, fig. 4 (not described).

1996 Nostocites vesiculosa - Sebbar & Mamet, p. 164, pl. 3, fig.3. Description - Only one section, with several groups of'cells' apparently linked to a convex micritized substrate; 'cells' composed of yellow, hyaline calcite; dark centers well marked.

D i m e n s i o n s - Length of 'colony' 1.24 ram; height of 'colony' 0.30 mm; maximal dimensions of one cell 0.03 x 0.06 ram.

393

PERMIAN

BACHKIRIAN

~

Berestovia~ ~ Praedonezella

~~ ~ ~

Ungdaretta late VISEAN

early VISEAN

late TOURNAISIAN

~~ ~ I

Stacheoides

....;~

"~"'°

~ ~ ~ ~i

~

~

~r~

/~ / I~amaenefl . a

l ~

,.~,

| ~Stacneoidella~ ! \ ,, I Stach~

t .

early TOURNAISIAN

Pokorninella

FAMENNtAN

graeca

/

/

FRASNIAN

GIVETIAN

Pokorninella

EMSIAN

Praecominella

FIGURE 19 - Hypothetical phylogeny of some Aoujgaliida with location of the Claracrustinae subfam, nov. and possible explanation of the convergences between Claracrusta ungdarelloidea nov. sp. and Ungdarella. 1: evolutionary trends; 2: Claracrustinae subfam, nov. Phylogdnie probable de divers Aoujgaliides, avec la position des Claracrustinae nov. subfam, et une possible explication des convergences existant entre Claracrusta ungdarelloidea nov. sp. et Ungdarella. 1: tendances ~volutives; 2: enveloppe des Claracrustinae nov. subfam.

R a n g e - Same as the genus.

Incertae sedis (or 'Pseudo-algae' auctoris) Class ISCHYROSPONGIATermier & Termier, 1973 emend. Termier, Termier & Vachard, 1977 R e m a r k s - The Class Ischyrospongia (including the 'algal' Palaeosiphonocladales of the authors), principally composed of two orders: Aoujgaliida and Moravamminida, is relatively rare in our material, with Eflugelia johnsoni (FLOGEL, 1966) VACHARDin M a s s a & Vachard, 1979 (Fig. 18.7), Chuvashovia densifolia VACHARD, 1980 (Fig. 18.9), Evlaniopsis m o n t a n a VACHARD, 1980 (Fig. 18.8) and Claracrusta spp. (Fig. 17.9, Fig. 18.2-6) with especially a new species Claracrusta ungdarelloidea nov. sp. Aoujgaliida differ from the red algae by the functional character of the opening between the skeletal units ('cells' of the authors), a n d the perforated wall. Moravamminida differ from the blue-green algae or cyanobacteria by the type of calcitic wall, the presence of a septation, and the multiple convergences with the Aoujgaliida. The Wetheredellaceae lack septation, but can be hypothetically included in this group. R a n g e - From Silurian to Late Permian, particularly diversified during the Carboniferous. Little studied in the Permian, b u t largely present in Afghanistan (Vachard 1980), S u m a t r a (Fontaine & Vachard 1984), Italy (Vachard & Miconnet 1989), etc.; rare in the Permian of the B a t a i n Plain, Oman.

Order MORAVAMMINIDAPokorny, 1951 nomen translat, and emend. Termier, Termier & Vachard, 1975 ex family Moravamminidae Family DONEZELLIDAETermier, Termier & Vachard, 1975 (or Donezellaceae nomen translat. herein) Subfamily CLARACRUSTINAEnov. subfam. C o m p o s i t i o n - Claracrusta VACHARD,1980; Berestovia BERCHENKO, 1982; Ungdarella auctoris pars non MASLOV, 1956; Stacheoidella MAMET& Roux in Mamet et al., 1987, Iberiaella RACZ, 1984, ? Kleinbergella MAMET & BOULVA~N, 1992; ? Pseudostacheoides PETRYK& MAMET, 1972. D i a g n o s i s - Encrusting colonies; skeletal elements composed of large horizontal laminae and sometimes vertical pillars; erected parts can exist in some species of Claracrusta; wall yellowish, calcitic, hyaline, sometimes weakly differentiated as in

Donezella. C o m p a r i s o n s - From the typical Donezellidae, considered herein as the subfamily Donezellinae s.s., the Claracrustinae differ by the great development of the encrusting part. They seem to represent the phylogenetic link between the Donezellidae and other pseudo-algae like Fasciella or Calcifolium. From the Ungdarellidae, they differ by the badly organised and rarely and shortly ramified development. In the material from Forni Avoltri, Italy (Vachard & Argyriadis, in press), various sections are identical to the illustrated representatives of 'Ungdarella'peratrovichensis MAMET& RUDLOFF, 1972, but look more like Claracrusta than Ungdarella. For us, 'Ungdarella'peratrovichensis is a typical Claracrustinae, although generically different from Claracrusta. Atypical Donezellinae, like Kleinbergella and Pseudostacheoides, are atypical Claracrustinae by the stronger development of vertical elements. R a n g e (Fig. 19) - Doubtful in the Late Devonian (Frasnian) of Belgium with Kleinbergella. V i s e a n with Stacheoidella. Abundant from the Bashkirian to the Early Permian with Claracrusta-Berestovia-Iberiaella, constituting some complex biopisolites, generally called Ottonosia. Present in the Middle Permian. Rare in the Late Permian.

Genus

Claracrusta

VACHARD, 1980

T y p e s p e c i e s - Girvanella catenoides HOMANN, 1972.

Claracrusta ungdarelloidea nov. sp. Fig. 18.2-6 D e r i v a t i o n o m i n i s - Similar to Ungdarella. H o l o t y p e - Fig. 18.4. P a r a t y p e s - Fig. 18.3,6. T y p e l o c a I i t y - Wadi Hasad, 10 k m WSW of the village Sal (UTM coord. 759068/2444367), B a t a i n Plain, Oman. T y p e l e v e l - Kubergandian.

D i a g n o s i s - A Claracrusta characterized by the great number of superimposed elements and the convergence with Ungdarella.

394 - Irregular shape, rather elongate and ramose, with bluntly small protuberances and smooth extremities; central network relatively regular with repetitively spaced constrictions, pierced in the center by a small aperture. Our specimens often appear perforated by endobiont microorganisms. Description

D i m e n s i o n s - Colonies: 0.60 x 1.50 mm to 3.00 x 11.00 mm;

thickness of skeletal elements 0.01-0.02 ram; intervals between laminae 0.05 mm; intervals between constrictions 0.080.10 mm. Comparisons - Differs from the congeneric species C. c a t e n o i d e s , C. c l a r a c r u s t a a n d C. h i r t i p e s b y t h e

accumulation

of horizontal skeletal units.

R a n g e - Discovered in the Kubergandian of the Batain Plain,

Oman. Incertae sedis (may be umbellids) Genus Sphairionia

TIEN, 1 9 8 9 n o n 1 9 8 8

T y p e s p e c i e s - Sphairionia sikuoides S y n o n y m - Pseudosphairionia

TIEN,

PRGNINA,

1989 non 1988.

1996.

- Test composed of a large internal cavity with 2 or 3 protruding apertural necks; wall calcitic, y e l l o w i s h , f i b r o u s r a d i a t e , w i t h w e l l i n d i v i dualized fibres, bordered by an internal layer, dark, m o r e o r l e s s m i c r o g r a n u l a r ( F i g . 20). Diagnosis

R e m a r k s - The first valid description of the genus is for us the

publication of Tien (1989, Beijing Carboniferous Congress). Other names exist in the literature (see below the synonymy of the type-species). The 'subgenus' Pseudosphairionia of Pronina (1996) is synonymous (Fontaine et al. 1998). R a n g e - Midian of Peninsular Thailand (Tien 1988; Fontaine

& Suteethorn 1988; Fontaine et al. 1998), Peninsular Malaysia (Fontaine et al. 1988), Cambodia (Tien 1979), Afghanistan (Vachard 1980), Turkey (KSylfioglu & Altiner 1989), Iran (Baghbani 1993), Italy (Vachard & Miconnet 1989), Greece: Peloponnesus (Trifonova 1985). Early Midian of Transcaucasia (Pronina 1996), Crimea (Kotlyar et al. 1999). Midian? of Sumatra (Pia 1937). Discovered in the Early Midian of the Batain Plain, Oman.

Sphairionia

s i k u o i d e s TIEN, 1 9 8 9 n o n 1 9 8 8 Fig. 21.1

? 1937 'ein recht sicherer Schnitt durch ein Characeenoogonium' - Pia, p. 835 (not illustrated). 1979 Incertae sedis sp. 2 - Tien, pp. 143-144, pl. 32, figs 3-7. ? 1979 Umbellina (?) sp. 3 - Tien, p. 78, pl. 3, figs 19-20 (ou S. galinae nov. sp.). v. 1980 Permumbella serotina - Vachard, p. 102 (not illustrated). 1985 Umbellina ? sp. - Trifonova, pl. 4, fig. 9 (not described). 1987 Sphairionia sikuoides nov. gen. nov. sp. - Tien, p. 90 (not described, not illustrated: nomen nudum). 1988 Sphairionia sikuoides nov. gen. nov. sp. - Fontaine & Suteethorn, p. 45 (not illustrated: nomen nudum). 1988 Sphairionia sikuoides - Tien in Fontaine & Suteethorn, p. 109-110, pl. 9, figs 7-15 (holotype not designated: nomen nudum). 1988 Sphairionia, a newly described genus, common in Southeast Asia - Fontaine et al., fig. 6:5 (nomen nudum). v. 1989 Sphairionia sikuoides TIEN - Vachard & Miconnet, p. 301, pl. 1, figs 10-11 (with a reference to the Tien's abstract of 1987). * 1989 Sphairionia sikuoides - Tien, pp. 75, 77, pl. 1, figs 1-6, pl. 2, figs 1-9. 1989 Incertae sedis 1 - KSyliioglu & A]tiner, p. 481, pl. 11, fig. 28. 1993 Sphairionia sikuoides - Baird, Dawson & Vachard, p. 250, fig. 3: 8. 1993 Soroorina ghezelbashi - Baghbani, pl. 16, fig. 1 (nomen nudum). 1994 Sphairionia sikuoides - Fontaine et al., pl. 7, figs 1-3, pl. 14, figs 5-6, pl. 15, figs 5-6, pl. 16, figs 6-7, pl. 22, fig. 6. 1995 Sphairionia sikuoides - Leven, p. 11 (not illustrated). 1996 Sphairionia sikuoides - Pronina, p. 114~ pl. 1, fig. 12, pl. 2, fig. 13. 1996 Sphairionia (Pseudosphairionia) tienii subgen, et sp. nov. - Pronina, p. 115, pl. 1, fig. 11, pl. 2, figs 14-16. 1998 Sphairionia sikuoides - Fontaine et al., p. 13 (not illustrated). 1999 Sphairionia (Pseudosphairionia) tienii PRONINA- Kotlyar et al., pp. 309, 312 (not illustrated). R e m a r k s - Only one specimen has been found, whose dimen-

sions are: diameter including the protuberances 0.93 mm, outer diameter 0.75 mm, inner diameter 0.50 mm, wall thickness 0.13 mm, thickness of internal layer 0.03 ram. R a n g e - Murgabian or Midian of Afghanistan: Abtchakan,

Koh-e Arca, Wardak (material of Vachard 1980, revisited). Early Midian of Italy, Turkey, Crimea, Iran, Central Pamir; Arpa Formation of Transcaucasia (for references, see the analysis of the genus). Late Midian with Lepidolina in Thailand, Malaysia and Cambodia (Sphairionia is not only Early Midian in age, as indicated by Pronina 1996). Discovered in the Early Midian of the Batain Plain, Oman. S p h a i r i o n i a g a l i n a e nov. sp. Fig. 21.2-9 D e r i v a t i o n o m i n i s - Dedicated to Dr. Ganelina P. Pronina,

from Saint Petersburg, Russia, for her contributions to the biostratigraphy of the Permian. H o l o t y p e - Fig. 21.6. P a r a t y p e s - Fig. 21.2-5,7-9,

T y p e l o c a l i t y - 8 km from the village Bu Fashiqah (UTM coord. 776242/2472356), Batain Plain, Oman. T y p e l e v e l - Early Midian. - A small species with very weakly developed protuberances. Diagnosis

- This very small species exhibits characteristically the wall of Sphairionia, and some sections show the typical necks. Only small extern a l p r o t u b e r a n c e s a r e v i s i b l e (Fig. 21.5-6,9). A s t h i s

Description

FIGURE 20 - Comparison between Sphairionia sikuoides TIEN, 1989 non 1988, and S. galinae nov. sp. Comparaison entre Sphairionia sikuoides TIEN, 1989 non 1988 et S. galinae nov. sp.

395

3

8

5

7

6

11

14

9

10

12

15

16

FIGURE 21 - 1. Sphairionia sikuoides TIEN, 1989 non 1988. Transverse section, rare in our material, to compare with S. galinae nov. sp. Early Midian. Batain Plain. H541/922(7). 2-9. Sphairionia galinae nov. sp. 2: Transverse oblique section with juvenile Yangchienia sp. Early Midian. Batain Plain. H132(42)/908(28). 3: Transverse section with the typical wall. Early Midian. Batain Plain. H541/922(2). 4: Transverse section with some external rugosities, compare with Fig. 21.1. Early Midian. Batain Plain. H541/922(8). 5: Transverse section with a polygonal outline. Early Midian. Batain Plain. H541(42)/907(6). 6: Holotype. Axial section Early Midian. Batain Plain. H489(7)/907(14). 7: Subaxial oblique section. Early Midian. Batain Plain. H541(29)/906(28). 8: Paratype. Transverse section with half stellate outline. Early Midian. Batain Plain. H489(23)/907(22). 9: Transverse typical section. Early Midian. Batain Plain. H132(14)/907(36A). 1O. Diplosphaerina ex gr. inaequalis (DERVILLE, 1931). Transverse section. Early Midian. Bataln Plain. H541(36)/906(36). 11-14. Pseudovermiporella elliotti ERK • BILG~2TAY,1960. Note the particular semi-recrystallization of the tube. Late Midian. Batain Plain. 11: Transverse section. H250/922(27). 12: Subtransverse sections. H250(10)/909(35). 13: Oblique section. H250(10)/909(37). 14: Transverse section H250(8)/909(32). 15-16. Pseudovermiporella longipora (PRATURLON, 1963). Early Midian. Bataln Plain. H133(1)/913(29). 15: Transverse section encrusting a Productacean brachiopod. H132(42)/908(28). 16: Oblique section. H133(2)/913(31).

396 species is common in our material, and seems to be of taxonomic significance, it is described herein.

and/or symbiosis, but its erroneous interpretation as a pure alga resists victoriously to all the arguments.

D i m e n s i o n s - Outer diameter 0.23-0.37 mm, i n n e r diameter 0.16-0.20 mm, wall thickness 0.03-0.06 mm, maximal height of protuberances 0.08 mm.

T y p e s p e c i e s - Pseudovermiporella sodalica ELLIOTT,1958.

Comparisons

-

Differs from the type-species

Sphairionia by the smaller dimensions (outer diameter of 0.50 to 1.00 mm for example) and the inconspicuous apertural necks. Differs from the calcispherids by the wall microstructure. R a n g e - So far only known from the Early Midian of the B a t a i n Plain, Oman.

Incertae sedis (calcispherids) Family TUBERITINACEAEnomen translat, herein o r TUBERITINIDAE Miklukho-Maclay, 1 9 5 8 Remarks - Tuberitinaceae are generally considered as foraminifers or 'calcispheres'. The doubt is possible, due to both conspicuous generations, called diplospheric and t u b e r i t i n a t e by Conil & Lys (in Conil et al. 1973, fig. 1, p. 211). Appearing very early in the Silurian, the Tuberitinaceae go on without modification until the end of the Permian. The gener a are probably only based on diagenetic modifications of the wall (see discussion in Vachard 1994). Permian Tuberitinaceae were accurately studied by Tien (1979). The family is poorly represented in Oman.

Incertae Sedis (or true foraminifers) Remarks - Convergences (or symbiosis ?) between foraminifers and algae are n u m e r o u s in t h e literature (see review in Vennin et al. 1997). For example, we t h i n k that: (a) Ellesmerella MAMET • ROUX in M a m e t et al. 1987 is p a r t of a porcelaneous foraminifer similar to some Nubeculariids and not a n alga; (b) Tubiphytes is a consortium alga/foraminifer (as well as Ramovsia, Koivaella, Aeolisaccus, Rigidicaulis, Galaenella, Miliolipora); (c) the example of Lithocodium seems now well elucidated (Schmid & Leinfelder 1995); (d) Pseudovermiporella constitutes at least another case of convergence

Genus Pseudovermiporella ELLIOTT, 1958 Remarks - Although this genus was correctly i n t e r p r e t e d as a porcelaneous foraminifer, m a n y algologists a t t r i b u t e Pseudovermiporella to enigmatic algae, probably because, as indicated by Henbest (1963, p. 33): 'it presents, however, a n e x t e r n a l pore s t r u c t u r e t h a t suggests k i n s h i p w i t h t h e Dasycladacean algae and t h a t is u n u s u a l for the foraminifers'. PseudovermiporeUa sodalica ELLIOTT, 1958, Vermiporella (?) nipponica ENDO in Endo & Kanuma, 1954 and Vermiporella sumatrana PIA, 1937, are well known and generally considered as t h r e e synonymous (Homann 1972; E m b e r g e r 1976). In our opinion, P. sodalica is slightly different by its thicker wall. P. elliotti ERK & BILG?JTAY,1960 is rare in the Omanese microfacies, and has been poorly described; however, the species is perfectly valid and characterized by rounded pores, and interpore wall segments, equal to the pore diameter (Fig. 21.11-14). P. longipora (PRATURLON, 1963) nov. comb. (Fig. 21.15-16) possesses long incurved pores.

Pseudovermiporella elliotti ERK & BIT,GOTAY,1960 Fig. 21.15-16. 1960 Pseudovermiporella elliotti - Erk & Bilgtitay, pp. 110111, pl. 2, figs 1-3. 1968 Pseudovermiporella elliotti - Elliott, p. 72 (not illustrated). 1976 Vermiporella elliotti - Emberger, p. 14 (not illustrated). 1990 Pseudovermiporella elliotti - Fliigel, pl. 2, figs 5, 7. non 1992 Pseudovermiporella elliotti - Sha et al., p. 181, pl. 1, fig. 1 (= Pseudovermiporella nipponica).

R e m a r k s - Our specimens are r e m a r k a b l y similar to the Turkish material, in shape and dimensions, especially with identical pore and interpore diameters of circa 0.04 ram. R a n g e - Bolorian of Turkey (Erk & Bilgfitay 1960; Flfigel 1990). Its presence in the Changxingian of South C h i n a (Sha

Tubiphytes sensu stricto

transverse section

rubiphytes / cavities stage

® fncr~ of~ cavil

growthof lateralcavities

© ®

transverse section

phytes I foraminifer stage singof wall':

fo~raminifer ~ iliolid odophtalmiid gitudinal section

~al section

FIGURE 22 - A-D. Reconstruction of Tubiphytes. A: Various aspects of the central and lateral vacuoles. B: Modifications of the aspect of a cavity in the case of a Tubiphytes related with a Nubeculariid foraminifer. C-D: Sections at different stages of the central cavity wall dissolution; C: the cavity remains multiple; D: the cavity appears simple. Reconstitution de Tubiphytes en temps que micro-dcosyst~me. A: diffdrents aspects des vacuoles centrales et latdrales. B: Modifications de l'aspect des cavitds dans le cas d'un Tubiphytes lid ~t un foraminif~re Nubdculariidd. C-D: Sections 8t diffdrents stades de dissolution de la cavitd centrale; C: la cavitd reste multiple; D: la cavitd appara~t unique.

397 SCHaeFER• SENOWBARI-DARYAN1983 , (Fig. 24.6) and forms more similar to foraminifers (H132, Fig. 24.8-9). Various authors consider Shamovella as a prioritary synonym of Tubiphytes (Riding 1993; Le Mone 1995; and probably Kirkland et al. 1998, which speak of'Shamovella/Tubiphytes', text-fig. 14, p. 965), for others (Chuvashov et al. 1993) the two forms exist, represented at the level of two subspecies 'Tubiphytes obscurus shamovella RAUSER-CHERNOUSSOVA'(pl. 14, figs 2-4) and 'Tubiphytes obscurus obscurus MASLOV'(pl. 14, fig. 5).

1®

,

Tubiphytes from Oman were already mentioned by Elliott (1962) and by Glennie et al. (1974, fig. 4.4.6 to 4.4.9). Tubiphytes is cosmopolitan and ecologically widespread along the ramps or carbonate platforms (Wahlman 1988; Chun & Mamet 1993; Asquith & Drake 1985; Beauchamp 1989; Sano et al. 1990), therefore the 'Tubiphytes community' of Flfigel (1977, 1979) is an imprecise concept. - Cosmopolite from the Moscovian to the Early Cretaceous (Razgallah & Vachard 1991; Riding & Guo 1992; Chun & Mamet 1993). T obscurus is cited in the Early Moscovian, but according to our observations, the transformation from Miliolid encrusting foraminifer (more or less Palaeonubecularia REITLINGER,1950) to Tubiphytes occurs in the latest Moscovian/earliest Kasimovian (Middle/Late Carboniferous boundary) in the Donetz Basin (Vachard, unpublished). Range

Fmt~E 23 - Hypothetical life-cycle of Tubiphytes as a case of coevolution. 1: algal mucilage; 2: foraminiferal or algal host-skeletons; 3: organic or lithic substrate. Stippled parts correspond to an alga or a dominantly algal organism. A: free generation; B: aspects of the host tubes of Koivaella (right) or nubeculariid foraminifer (left); C: attachment of the free alga; D: interaction between alga and host, dissolution of the host wall; E: the host wall is replaced by a hollow, in the center of the alga; F: transfer of substrate and true attachment of the Tubiphytes with its internal cavity; G: multiplication of this type of Tubiphytes. A: a part of this colony can become free again. Cycle de vie hypoth~tiquede Tubiphytes considdr~ comme un cas de co-dvolution entre une algue et son substrat. 1: mucilage algaire; 2: squelette externe des hStes, algues ou foramini~res; 3: substrat organique ou lithique. Les zones en pointilld correspondent aux algues proprement dites ou aux stades & dominante algaire. A: gdndration libre; B: aspects des h6tes: tubes d'algue Koivaella (& droite) ou foramini~re Nubgculariidd (~ gauche); C: fixation de l'algue; D: interaction de l'algue et de son support avec notamment dissolution de la paroi de celui-ci; E: remplacement de la paroi de l'h6te par une cavitd centrale; F: changement de substrat et premiere vdritable fixation de Tubiphytes porteur de sa cavitd interne; G: bourgeonnement de ce type de Tubiphytes; A: retour d'une partie de ces colonies ~t la vie libre.

et al. 1992) seems to be misinterpreted. Discovered in the Late Midian of the Batain Plain, Oman.

Genus Tubiphytes MASLOV,1956 emend, herein T y p e s p e c i e s - Tubiphytes obscurus MASLOV,1956. S y n o n y m s - Shamovella, Nigriporella. R e m a r k s - The enigma of Tubiphytes is absolute (Vachard & Montenat 1981; Roux 1985; Mamet et al. 1987; Razgallah & Vachard 1991; Riding & Guo 1992; Senowbari-Daryan & Flfigel 1993; Wang et al. 1994). According to Vennin et al. (1997) Tubiphytes is a non-Linnean microecosystem of unknown affinity, possibly a very complex symbiosis of foraminifers and algae, and not only an incrustation as suggested by Senowbari-Daryan & Flfigel (1993), since the association of 'small miliolid forms' with that forming Tubiphytes is found in the Palaeozoic as well as the Late Jurassic (in this case, see for example Barattolo & Pugliese 1987). The ecotypes, observed in Oman are very numerous, with typical T obscurus (H250, Fig. 24.1) but also with T gracilis

New hypothesis (Figs 22; 23; 24.1,3-9) - We interpret Tubiphytes basically as a free cyanobacteria or alga. During the period of its life, to continue to grow, this alga developed symbiosis with other organisms, e.g. paleaonubeculariid, simple or ramified tubes which probably belong also to the miliolid, etc. The symbiosis between algae and foraminifers (the algal cells occupying the intercrystalline spaces into the wall of the foraminifer) is complete and balanced. The result of the increasing wall is, at the beginning, some doubtful 'nubeculariids' foraminifers, for example 'Cordiformis' of Gfivenc (1965, nomen nudum) or Nodophthalmidium (Bernier 1984; Baratollo & Pugliese 1987), or 'incertae sedis 5' of Tien (1979), and at the end, the aspect illustrated by Tellez-Giron & Nestell (1983, pl. 3, fig. 2; pl. 9, fig. 7) or Vachard & Ferri~re (1991, pl. 1, fig. 4). With the tubular foraminifers, the first growth gives an Aeolisaccus kotori stage (i.e. a cylinder with an anormally thick 'wall'). Progressively, the 'wall' becomes predominant and proportionally enormous, growing probably by indistinct concentric layers, whose existence can be deduced from the accumulation of spicules (Fig. 24.4), foraminifers, and other organic silt-sized remains included in the wall of some Tubiphytes. Later, the equilibrium is broken and the alga begins to feed on the wall of the foraminifer, rapidly replaced by a larger, more irregular and asymmetrical cavity. This cavity is discontinuous (see pl. 3, fig. 1 ofVachard et al. 1993). The shape of Tubiphytes becomes typical with only one cavity within an algal mucilage; 'metamorphosis' is anyway necessary for giving a true Tubiphytes. This occurs when the link with the symbiont is broken, and when Tubiphytes leaves the symbiont for another definitive support, i.e. corals, calcisponges or more generally another Tubiphytes. These successive encrusting stages of Tubiphytes give the classical aspect of this microfossil. Finally, Tubiphytes can be encrusted by Archaeolithoporella, but this relation seems to be entirely passive; Archaeolithoporella layers have been included in the diagnosis of Tubiphytes by Maslov (1956). Various patterns and stages of Tubiphytes can narrowly coexist in the microfacies (see Vachard & Ferri~re 1991, pl. 1, fig. 1; and this study Fig. 24.1,3 or 6-7). The presently described material and the new taxa are part of Marc Hauser thesis collection, Mineralogisch-petrographisches Institut und geologisches Institut, University of Bern, Switzerland. - The present paper is a result of a geological mapping and research program (sheets Sur and A1 Ashkahrah 1:100'000) of the Universities of Bern and Geneva supported by the Swiss National Foundation (project n ° 2043'056.95 T.P. and n ° 20-50577.97 L.Z.); and the Director General of Minerals, Mohammed Bin Hussein Bin Kassim, Ministry of Acknowledgements

398

5

6

7

i FIGURE24 - 1, 3-9. Tubiphytes obscurus MASLOV,1956. Very diversified morphotypes. 1: Typical piriform longitudinal section. Late Midian. Batain Plain. H250(4)/914(27). 3: Atypical morphotype similar to a Nubeculariid foraminifer. Late Midian. Batain Plain. H250(4)/909(19). 4: Typical individual showing some spicule clearly trapped in an algal mucilage, and not a sponge skeleton. Late Midian. Batain Plain. H250(4)/924(6). 5: Atypical foraminiferal aspect (right) passing to a typical aspect (links). Kubergandian. Batain Plain. H180(5)/914(19). 6: Morphotype especially slender. Kubergandian. Batain Plain. H236(3)/910(10). 7: Typical transverse section. Kubergandian. Batain Plain. H236(4)/910(9). 8: Atypical aspect very similar to a true Nubeculariid foraminifer. Early Midian. Batain Plain. H133(5)/913(34). 9: Completely atypical pattern never described, apparently cribrate. Early Midian. Batain Plain. H132(10)/907(34). 2. Sparaphralysia orientalis sp. nov. Detail of the holotype (see Fig. 10.1). Late Midian. Batain Plain. H250/909(27).

399 Commerce and Industry, Sultanate of Oman, and Dr. Hilal A1 Azry, Director of the Geological Survey. Drs M.F. Perret-Mirouse et B. Mamet are thanked for their helpful reviews, and J. Metzger for the graphical assistance.

REFERENCES Accordi, B., 1956. Calcareous Algae from the Upper Permian of the Dolomites (Italy), with stratigraphy of the Bellerophonzone. Journal of. Paleontology of the Society India, inaug. number 1, 75-84. Alsenverg, D.E., Astakhova, T.V., Berchenko, O.I, Brazhnikova, N.E., Vdovenko, M.V., Dubaeva, N.N., Zernetskaya, N.V., Poletaev, V.I., Sergeeva, M.T., 1983. Verkhneserpukhovskii podyarus Donetskogo basseina (Late Serpukhovian substage in the Donets Basin). Akademiya Nauk Ukrainskoi SSR, Institut Geologicheskikh Nauk, 164 pp. [in Russian]. Argyriadis, I., de Graciansky, P.C., Lys, M., 1976. Datation de niveaux rouges dans le Permien marin p6ri-4g6en. Bulletin de la Soci6t6 g6ologique de France 18 (2), 513-519. Asquith, G.B., Drake, J.F., 1985. Depositional history and reservoir development of a Permian Fistulipora-Tubiphytes Bank Complex, Blalock Lake East Field, West Texas. In: Roehl, RE., Choquette, P.W. (Eds), Carbonate Petroleum Reservoirs. Springer Verlag, Berlin, 311-316. Baars, D.L., 1992. Kansaphyllum, a new Late Pennsylvanian phylloid algal genus. Journal of Paleontology 66, 697-701. Baars, D.L., Torres, A.M., 1991. Late Paleozoic phylloid algae, a pragmatic review. Palaios 6, 513-515. Babcock, J.A, 1986. The puzzle of alga-like problematica, or rummaging around in the algal wastebasket. In: Hoffman, A., Nitecki, M.H. (Eds), Problematic fossil taxa. Oxford Monographs on Geology and Geophysics 5, 15-26. Baghbani, D., 1993. The Permian sequence in the Adadeh region, Central Iran. In: Nairn, A.E.M., Koroteev, V. (Eds), Contributions to Eurasian geology, Part II. Occasional publ. ESRI, Columbia, new series 9 (2), 7-22. Baird, A., Dawson, O., Vachard, D., 1993. New data on biostratigraphy of the Permian Ratburi limestone from North Peninsular Thailand. Proceedings International Symposium on Biostratigraphy of Mainland Southeast Asia, Facies and Paleontology, Chiang Mai, 243-259. Barattolo, F., Pugliese, A., 1987. I1 Mesozoico dell'Isola di Capri. Quaderni dell'Academia Pontiana 8, 1-37. Bassoullet, J.P., Bernier, P., Deloffre, R., Genot, P., Jaffrezo, M., Vachard, D., 1979. Essai de classification des Dasycladales en tribus. Bulletin des Centres de Recherche ExplorationProduction Elf-Aquitaine 3 (2): 429-442. Bassoullet, J.P., Bernier, P., Deloffre, R., Genot, P., Poncet, J., Ronx, A., 1983. Les algues Udot4ac6es du Pal6ozoique au C6nozo~que. Bulletin des Centres de Recherche ExplorationProduction Elf-Aquitaine 7 (2): 449-621. Beauchamp, B., 198. Lower Permian (Sakmarian) Tubiphytesbryozoan buildup, Southwestern Ellesmere Island, Canadian Arctic Archipelago. In: Geldsetzer, H.H.T., James, N.P., Tebbutt, G.E. (Eds), Reefs, Canada and adjacent areas. Canadian Society of Petroleum Geologists Mem. 13, 585-589. Bebout, D.C., Coogan, A.H., 1964. Algal-genus Anthracoporella PIA. Journal of Paleontology 38, 1093-1096. B6chennec, F., 1987. G6ologiedes nappes Hawasina dans les parties orientale et centrale des Montagnes d'Oman. Document du B.R.G.M. 127, 1-417. B6chennec, F., Roger, J., Chevrel, S., Le Metour, J., 1992a. Geological map of Al Ashkharah sheet NF 40-12, scale 1:250'000 with explanatory notes. Directorate General of Minerals, Oman Ministry of Petroleum and Minerals, Muscat. B6chennec, F., Roger, J., Le Metour, J., Wyns, R., 1992b. Geological map of Seeb sheet .NF 40-03, scale 1:250'000 with explanatory notes. Directorate General of Minerals, Oman Ministry of Petroleum and Minerals, Muscat. Belka, Z., 1981. The alleged algal genus Aphralysia is a foraminifer. Neues Jahrbuch ftir Geologie und Palfiontologie Monatshefte 1981 (5), 257-266. Berchenko, O.I., 1982. Novye vidy zelenykh vodoroslei iz otlozhenii verkhneserpukhovskogo podyarusa Donbassa (New spe-

cies of green algae from the Late Serpukhovian substage deposits of Donbass). In: Sistematika i evolyutsiya dreniikh rastenii Ukrainy (Systematics and evolution of fossil plants of Ukraine). Kiev Naukova Dumka, 51-55 [in Russian]. Bernier, P., 1984. Les formations carbonat6es du Kimm6ridgien et du Portlandien dans le Jura m4ridional, stratigraphie, micropal6ontologie, s6dimentologie. Documents des Laboratoires de G6ologieLyon 92 (2), 1-803. Bignot, G., Kodra, A., Neumann, M., Pirdeni, A., 1982. Le Permien sup6rieur des Alpes Albanaises; 6tude pr61iminaire. Comptes Rendus de l'Acad6mie des Sciences, Paris 295, 883886. Brenckle, P.L., Marshall, F.C., Waller, S.F., Wilhelm, M.H., 1982. Calcareous microfossils from the Mississippian Keokuk Limestone and adjacent formations, Upper Mississippi River Valley: their meaning for North American and intercontinental correlation. Geologica et Palaeontologica 15, 47-88. Chanton, N., 1965. Nouvelle contribution ~ l'6tude des algues calcaires du Carbonif~re Saharien. Bulletin de la Soci6t6 g6ologique de France 7 (7), 402-409. Chun, L., Mamet, B., 1993. Late Carboniferous and Early Permian algal microflora (Liuzhou, South China). Bulletin of Canadian Petroleum Geology 41 (1), 70-78. Chuvashov, B.I., 1974 - Permskie izvestkovye vodorosli Urala (Permian calcareous algae from Urals). In: Papulov, G.N., Chuvashov, B.I. (Eds), Vodorosli, brakhiopodi i miospory iz permskikh otlozhenii zapadnogo Urala (Algae, brachiopods and miospores from the Permian deposits of western Urals). Akademiya Nauk SSSR, Uralskii Nauchnyi Tsentr, Trudy Instituta Geologii i Geokhimii 109, 1-76 [in Russian]. Chuvashov, B.I., Anfimov,A.L., 1988. Novye izvestkovye vodorosli srednego Karbona-nizhnei Permi Urala i Priuralya (New calcareous algae from Middle Carboniferous to Early Permian of Urals and Preural). In: Zhuravleva, I.T., Puchkov, V.M. (Eds), Izvestkovye vodorosli i stromatolity (Calcareous algae and stromatolites). Akademiya Nauk SSSR, Sibirskoe Otdelenie, Instituta Geologii i Geofiziki, 54-70 [in Russian]. Chuvashov, B.I., Luchinina V.A., Shuysky, V.P., et al. 1987. Iskopaemye izvestkovye vodorosli (Fossil calcareous algae). Akademiya Nauk SSSR, Sibirskoe Otdelenie, Trudy Instituta Geologii i Geofiziki 674, 1-200 [in Russian]. Chuvashov, B.I., Shuysky, V.P., Ivanova, R.M., 1993. Stratigraphical and facies complexes of the Paleozoic calcareous algae of the Urals. In: Barattolo, F. et al. (Eds), Studies on fossil benthic algae. Bolletino Societa Paleontologica Italiana 1, 93119. Conil, R., Groessens, E., Lys, M., 1973. Etude micropal6ontologique de la tranch6e d~/ves-Gomez6e. Bulletin de la Soci6t6 belge de G4ologie 82 (1), 201-239. Dawson, O., Racey, A., 1993. Fusuline-calcareous algal biofacies of the Permian Ratburi Limestone, Saraburi, Central Thailand. Journal of Southeast Asian Earth Sciences 8 (1-4), 4965. Dawson, W.C., 1992. Phylloid algal microstructures enhanced by epifluorescence petrography. Journal of Paleontology 66 (3), 523-525. De Castro, P., 1997. Introduzione allo studio in sezione sottile delle Dasicladali fossili; an approach to thin-section study of fossil Dasycladales. Quaderni dell'Academia Pontaniana 22, 1-261. Deloffre, R., 1987. Nouvelle classification des algues D~isycladales fossiles. Comptes Rendus de l'Acad6mie des Sciences, Paris, (2) 305, 1017-1020. DeIoffre, R., 1988. Nouvelle taxonomie des algues Dasycladales. Bulletin des Centres Recherche Exploration-Production ElfAquitaine, 12 (1), 165-217. Deloffre, R., Granier, B., 1991. Hypoth~se phylog6nique des algues Dasycladales. Comptes Rendus de l'Acad6mie des Sciences, Paris, (2) 312, 1673-1676. Delvolve, J.J., Perret, M.F., Vachard, D., 1987. D6couverte du Kachirien (Moscovieninf6rieur) ~ Fusulines et Algues dans le Massif de Cinco Villas (Pyr6n6es Basques, Espagne). Geobios, 20 (4), 407-414. Derville, H., 1931. Les marbres du calcaire carbonif6re en BasBoulonnais. Boehm Editeur, Strasbourg, 322 pp. Dil, N., Termier, H., Termier, G., Vachard, D., 1977. Contribution l'6tude stratigraphique et pal6ontologique du Vis4en sup6rieur et Namurien inf6rieur du bassin houiller de Zonguldak

400 (N.O. de la Turquie). Annales de la Soci4t6 g6ologique de Belgique 99, 401-449. Douglas, J.A., 1950. The Carboniferous and Permian faunas of South Iran and Iranian Baluchestan. Memoirs Geological Society of India, Palaeontologica Indica, new series, 22 (7), 1-57. Elliott, G.F., 1955. The Permian calcareous alga Gymnocodium. Micropaleontology 1 (1), 83-90. Elliott, G.F., 1957. New calcareous algae from the Arabian Peninsula. Micropaleontology 3 (3), 227-230. Elliott, G.F., 1958. Fossil microproblematica from the Middle East. Micropaleontology, 4 (4), 419-428. Elliott, G.F., 1962. More microproblematica from the Middle East. Micropaleontology 8 (1), 29-44. Elliott, G.F., 1965. The interrelationships of some Cretaceous Codiaceae (calcareous algae). Palaeontology 8 (2), 199-203. Elliott, G.F., 1968. Permian to Paleocene calcareous algae (Dasycladaceae) of the Middle East. Bulletin of the British Museum (Natural History), Geology suppl. 4, 1-111. Elliott, G.F., 1970. New and little-known Permian and Cretaceous Codiaceae (calcareous algae) from the Middle East. Palaeontology 13 (2), 327-33. Emberger, J., 1976. Les algues (Euchlorophyceae, Prasinophyceae, Rhodophyceae) du Carbonif'ere et du Permien; essai d'un inventaire bibliographique, g6ographique, stratigraphique. Bulletin de l'Institut de G6ologie du Bassin d'Aquitaine, num. sp., 168 pp. Emberger, J., 1979. Les algues (Euchlorophyceae, Prasinophyceae, Rhodophyceae) du Trias; essai d'un inventaire bibliographique, g6ographique, stratigraphique. Bulletin de l'Institut de G6ologie du Bassin d'Aquitaine, num. sp., 157 pp. Endo, R., 1951. Stratigraphical and paleontological studies of the later Paleozoic calcareous algae in Japan, I. Several new species from the Sakamotozawa section, Hikoroichi-mura, Kesen-gun in the Kltakami mountainous land. Transactions and Proceedings of the Paleontological Society of Japan 4, 121-129. Endo, R., 1958. Stratigraphical and paleontological studies of the later Paleozoic calcareous algae in Japan, XIII: a restudy of the genus Physoporella. Transactions Proceedings Paleontological Society Japan 31, 265-269. Endo, R., 1961a. Stratigraphical and paleontological studies of the later Paleozoic calcareous algae in Japan, XIV: Fossil algae from the Nyugawa Valley in the Hida Massif. The Science Reports of the Saitama University, series B, commemorative volume dedicated to Professor Riuji Endo, 177-207. Endo, R., 1961b. Calcareous algae from the Jurassic Torinosu Limestone of Japan. The Science Reports of the Saitama University, B, commemorative volume dedicated to Professor Riuji Endo: 53-75 [Endo 1961a, b (sic): mistake in the pagination and composition of the commemorative volume]. Endo, R., 1966. Contributions to the geology and paleontology of South-East Asia, XXIX. Some calcareous algae from Thailand. Japanese Journal of Geology and Geography 37 (3-4), 169173. Endo, R., 1969. Fossil algae from the Khao Phlong Phrab District in Thailand. Geology and Palaeontology of Southeast Asia 7, 33-85. Endo, R., Kanuma, M., 1954. Stratigraphical and paleontological studies of the later Paleozoic calcareous algae in Japan, VII: Geology of the Mino Mountain Land and southern part of Hida Plateau, with descriptions of the algal remains found in those districts. The Science Reports of the Saitama University B, 1 (3), 177-208. Erk, A.S., Bilgiitay, U., 1960. Pseudovermiporella'nin Ttirkiye' deki zuhuru hakkinda (On the occurrence of Pseudovermiporella in Turkey). Tfirkiye.Jeologii Kurumu Biilteni 7 (2), 108-118. Fan Jiasong, Rigby, J.K., Qi Jingwen, 1990. The Permian reefs of South China and comparisons with the Permian reef complex of the Guadalupe Mountains, West Texas and New Mexico. Bulletin of Yunnan University, Geology Studies 36, 15-55. Feldmann, J., 1946. Sur l'h6t6roplastie de certaines Siphonales et leur classification. Comptes Rendus de l'Acad6mie des Sciences, Paris (13) 222, 752-753. Filipovic, I., 1995 (ed.). The Carboniferous of Northwestern Serbia. Rasprave Geoloskog Zavoda 'Gemini' 25, 1-104. Flfigel, E., 1966. Algen aus dem Perm der Karnischen Alpen. Carinthia II, 25, 1-76.

Flfigel, E., 1977. Environmental models for Upper Paleozoic benthic calcareous algal communities. In: Flfigel, E (Ed.), Fossil algae. Springer Verlag, 314-343. Flfigel, E., 1979. Paleoecology and microfacies of Permian, Triassic and Jurassic algal communities of platform and reef carbonates from the Alps. Bulletin des Centres de Recherche Exploration-Production Elf-Aquitaine 3 (2), 569-587. Flfigel, E., 1980. Die Mikrofazies der Kalke in den TrogkofelSchichten der Karnischen Alpen. Carinthia II 36, 51-99. Flfigel, E., 1981. Lower Permian Tubiphytes/Archaeolithoporella buildups in the southern Alps (Austria and Italy). In: Toomey, D.F., (Ed.), European fossil reef models. Society of Economic Paleontologists and Mineralogists, Sp. Publ. 30, 143-160. Flfigel, E., 1987. Reef-Mound Enstehung: Algen-Mounds im Unterperm der Karnischen Alpen. Facies 17, 13-90. Flfigel, E., 1988. Halimeda: paleontological record and palaeoenvironmental significance. Coral Reefs 6, 123-130. Flfigel, E., 1990. Einschnitte in der Entwicklung permischer Kalkalgen. Mitteilungen des naturwissenschaftlichen Vereines Steiermark 120, 99-124. Flfigel, E., 1994. Pangean shelf carbonates: controls and paleoclimatic significance of Permian and Triassic reefs. In: Klein, G.D. (Ed.), Pangea: paleoclimate, tectonics and sedimentation during accretion, zenith and breakup of a supercontinent. Boulder, Colorado. Geological Society of America, Sp. Pap. 288, 247-266. Flfigel, E., Flfigel-Kahler, E., 1980. Algen aus den Kalken der Trogkofel-Schichten der Karnischen Alpen. Carinthia II 36, 113-182. Flfigel, E., Kochansky-Devid6 & Ramovs A. 1984. A Middle Permian calcisponge/algal/cement reef: Straza near Bled, Slovenia. Facies, 10: 179-256. Flfigel, E., & Reinhardt J. 1989. Uppermost Permian reefs in Skyros (Greece) and Sichuan (China): implications for the Late Permian extinction event. Palaios 4, 502-518. Fontaine, H., Gafoer, S., (Eds.), 1989. The Pre-Tertiary fossils of Sumatra and their environments. CCOP Technical Bulletin 19, 113-148. Fontaine, H., Ibrahim, B.A., Khoo, H.P., Tien, N.D., Vachard, D., 1994. The Neoschwagerina and Yabeina-Lepidolina zones in Peninsular Malaysia and Dzhulfian and Dorashamian in Peninsular Malaysia: the transition to the Triassic. Geological Survey of Malaysia 4, 1-74. Fontaine, H., Lys, M., Tien, N.D., 1988. Some Permian corals from East Peninsular Malaysia: associated microfossils, palaeogeographic significance. Journal of Southeast Asian Earth Sciences 2 (2), 65-78. Fontaine, H., Suteethorn, V., 1988. Late Paleozoic and Mesozoic fossils of West Thailand and their environments. CCOP Technical Bulletin 20, 1-217. Fontaine, H., Suteethorn, V., Vachard, D., 1998. Khao Yoi, a Permian limestone hill of the Ratburi area, Peninsular Thailand. CCOP Newsletter 23 (3), 12-14. Fontaine, H., Vachard, D., 1984. New paleontological data for the Upper Paleozoic of Sumatra. M6moires de la Soci6t6 g6ologique de France 147, 49-53. Garwood, E.J., 1914. Some new rock building organisms from the Carboniferous beds of Westmoreland. Geological Magazine 6, 265-271. Glennie, W.K., Boeuf, M.G.A., Hughes-Clarke, M.W., MoodyStuart, M., Pilaar, W., Reinhardt, B.M., 1974. Geology of the Oman Mountains. Verhandelingen van het Koninklijk Nederlands geologisch mijnbouwkunding Genootschap 31 (2), 423 p. Glintzboeckel, C., Rabat6, J., 1964. Microfaunes et microfaci~s du Permo-carbonif'ere du Sud-Tunisien. E. J. Brill, Leiden. Granier, B., Deloffre, R., 1994. Inventaire critique des algues Dasycladales fossiles, IIIe partie: les algues Dasycladales du Permien et du Trias. Revue de Pal6obiologie 14 (1), 49-84. Granier, B., Grgasovic, T. 2000. Les Algues Dasycladales du Permien et du Trias. Geologica Croatica 53 (1), 1-197. Groves, J.R., 1983. Calcareous foraminifers and algae from the type Morrowan (Lower Pennsylvanian) region of northeastern Oklahoma and northwestern Arkansas. Oklahoma Geological Survey, Bulletin 133, 1-65. Gfivenc, T., 1965. Etude stratigraphique et micropal6ontologique du Carbonif'ere et du Permien des Taurus occidentaux dans l'arri6re-pays d'Alanya (Turquie). Th6se de Doctorat Universit6 Paris, 2 volumes, 273 pp.

401 Gtivenc, T., 1979. Dasycladac6es m4taspondyles du Pal6ozoique sup6rieur et du Trias. Bulletin des Centres de Recherche Exploration-Production Elf-Aquitaine 3 (2), 625- 637. Hanzawa, S., 1961. Facies and micro-organisms of the Paleozoic, Mesozoic and Cenozoic sediments of Japan and her adjacent islands. E. J. Brill Publisher Leiden, 421 pp. Harris, A.G., Brenckle, P.L., Baesemann, J.F., Krumhardt, A.P., Gruzlovic, P.D., 1995. Comparison of conodont and calcareous microfossil biostratigraphy and lithostratigraphy of the Lisburne Group (Carboniferous, Sadlerochit Mountains, Northeast Brooks Range, Alaska). In: Dumoulin, J.A., Gray, J.E. (Eds), Geologic studies in Alaska by the U.S. Geological Survey, 1995. U. S. Geological Survey, Professional Paper 1574, 195-219. Hauser, M., Vachard, D., Martini, R., Matter, A., Peters, T., Zaninetti, L., 2000. The Permian sequence reconstructed from reworked carbonate clasts in the Batain Plain (Northeastern Oman). Comptes Rendus de l'Acad4mie des Sciences, Paris 330, 273-279. Henbest, L.C., 1963. Biology,mineralogy and diagenesis of some typical Late Paleozoic sedentary foraminiferal and algal-foraminiferal colonies. Cushman Foundation for Foraminiferal Research, Special Publication 6, 1-44. Homann, W., 1972. Unter- und tief-mittelpermische Kalkalgen aus den Rattendorf Schichten, dem Trogkofel-Kalk und Tressdorfer Kalk der Karnischen Alpen (Osterreich). Senckenbergiana Lethaea 53, 135-313. Immenhauser, A., Schreurs, G., Peters, T., Matter, A., Hauser, M., Dumitrica, P., 1998. Stratigraphy, sedimentology and depositional environment of the Batain Group. Eclogae geologicae Helvetiae 91, 217-235. Ivanova, R.M., 1988. Izvestkovye vodorosli vizeiskogo yarusa Urala (Calcareous algae of Visean stage from Urals). In Biostratigrafiya i litologiya verkhnego Paleozoya i Mezozoya Urala (Biostratigraphy and lithology from Late Paleozoic and Mesozoic of Urals). Sverdlovsk Uralskie Filial Akademiya Nauk SSSR, 4-23 [in Russian]. James, N.P., Wray, J.L., Ginsburg, R.N., 1988. Calcification of encrusting aragonite algae (Peyssonneliaceae); implications for the origin of Late Paleozoic reefs and cements. Journal of Sedimentary Petrology 58, 291-303. Jin Yu-gan, Wardlaw, B., Glenister, B.F., Kotlyar, G.V., 1997. Permian chronostratigraphic subdivisions. Episodes 20 (1), 915. Johnson, J.H., 1946. Lime-secreting algae from the Pennsylvanian and Permian of Kansas. Bulletin of the Geological Society of America 57, 1087-1120. Johnson, J.H., 1956. Archaeolithophyllum, a new genus of Paleozoic coralline algae. Journal of Paleontology, 30 (1): 53-55. Johnson, J.H., 1963. Pennsylvanian and Permian algae. Quarterly of the Colorado School Mines, 58 (3), 211 p. Karpinsky, A., 1908. Einige problematische Fossilien aus Japan. Verhandlungen der russischen mineralogischen Gesellschaft, St. Petersburg 2 (46), 257-272. Kazmierczak, J., Kempe, S., 1992. Recent cyanobacterial counterparts of Paleozoic Wetheredella and rela~ted problematic fossils. Palaios 7, 293-304. Khvorova, I.V., 1946. (On a new algal genus from the Middle Carboniferous deposits of Moscow basin). Doklady Akademiya Nauk SSSR 53 (8), 737-739 [in Russian]. Kirkland, B.L., Dickson, J.A.D., Wood, R.A., Land, L.S., 1998. Microbialite and microstratigraphy: the origin of encrustations in the Middle and Upper Capitanian Formation, Guadalupe Mountains, Texas and New Mexico, USA. Journal of Sedimentary Research 68 (5), 956-969. Kirkland, B.L., Moore, C.H. Jr., Dickson, J.A.D., 1991. Aragonitic Pennsylvanianphylloid algae from New Mexico: the missing link. American Association at Petroleum Geologists Bulletin 75 (D3), p. 610. Kirkland, B.L., Moore, C.H. Jr., Dickson, J.A.D., 1993. Well preserved, aragonitic phylloid algae (EugonophyUum, Udoteaceae) from the PennsylvanianHolder Formation, Sacramento Mountains, New Mexico. Palaios 8, 111-120. Kochansky-Devid4, V., 1970a. Permski mikrofosili zahodnih Karavank. Geologiji, Razprave in Porocila 13, 175-256. Kochansky-Devid6, 1970b. Die Kalkalgen des Karbons vom Velebit-Gebirge (Moskovien und Kassimovien). Palaeontologia Jugoslavica 10, 1-32.

Kochansky-Devid6, V., Herak, M., 1960. On the Carboniferous and Permian Dasycladaceae of Yugoslavia. Geoloski Vjesnik 13, 65-93. Konishi, K., 1954. A new species of Gymnocodium and its algal associates in the Permian Kosaki Formation of southern Kyushu, Japan. Japanese Journal of Geology and Geography 25 (1-2), 1-19. Konishi, K., Wray, J.L., 1961. EugonophyUum, a new Pennsylvanian and Permian algal genus. Journal of Paleontology 35 (4), 659-666. Kord6, K.B., 1950. Ostatki vodoroslei iz Kembriya Kazakhstana (Algal remains from the Cambrian of Kazakhstan). Doklady Akademiya Nauk SSSR, n.s. 73 (4), 809-812 [in Russian]. Kord6, K.B., 1965. Algae. In: Ruzhentsev, V.E., Sarycheva, T.G., (Eds), Razvitie i smena morskikh organizmov na rubezhe Paleozoya i Mezozoya (Evolution and change of marine organisms at the Paleozoic-Mesozoicboundary). Akademiya Nauk SSSR, Trudy Instituta Paleontologicheskikh Nauk 108, 268284 [in Russian]. Kotlyar, G.V., Baud, A., Pronina, G.P., Zakharov, Yu.D., Vuks, V.Ya., Nestell, M.K., Belyaeva, G.V., Marcoux, J. 1999. Permian and Triassic exotic limestone blocks of the Crimea. Geodiversitas 21 (3), 299-323. KSyltioglu, M., Altiner, D., 1989. Micropal6ontologie (foraminiferes) et biostratigraphie du Permien sup6rieur de la r6gion d'Hakkari (SE Turquie). Revue de Pal6obiologie 8 (2), 467503. Kozlowski, R., Kazmierczak, J., 1968. On two Ordovician calcareous algae. Acta Palaeontologica Polonica 13 (3), 325-346. Krainer, K., 1991. The limestone facies of the Auernig and Carnizza Formations (Auernig Group, Pontebba Supergroup; Carnic Alps). Giornale di Geologia 53 (1), 161-169. Krainer, K., 1995. Anthracoporella mounds in the Late Carboniferous Auernig Group Carnic Alps (Austria). Facies 32, 195214. Kulik, E.L., 1973. Cyanophyta. In: Einor, O.L. (Ed.), Stratigrafiya i fauna kamennougolnykh otlozhenii reki Shartym, Yuzhnyi Ural (Stratigraphy and fauna of the Carboniferous deposits of the Shartym river, southern Urals). Uralskoe Geologicheskoe Upravlenie, Kievskii Ordena Lenina GosudarstvennyiUniversitet ira. T.G. Shevchenko, Lvov, 39:48 [in Russian]. Kulik, E.L., 1978. Izvestkovye zelenye (Sifonovye) vodorosli Asselskogo i Samarskogo yarusov biogermogo massiva Shakhtau, Bashkiriya (Green calcareous algae, Siphoneae, from the Asselian and Sakmarian stages of the biohermal massif of Shakhtau, Bashkiria). Voprosy Mikropaleontologii 21, 182-215 [in Russian]. Le Mone, D.V., 1995. The role ofShamovella ('Tabiphytes') in bioherms. In: Garber, R.A., Lindsay, R.F. (Eds.), WolfcampianLeonardian shelf-margin facies of the Sierra Diablo. Seismic scale models for subsurface exploration. Annual Field Guidebook. West Texas Geological Society, Publication 95-97, 135-144. Leven, E.Ya., 1995. Permian and Triassic of the Rushan-Pshart zone (Pamir). Rivista Italiana di Paleontologia e Stratigrafia 101 (1), 3-16. Leven, E.Ya., 1997. Permian stratigraphy and Fusulinida of Afghanistan with their paleogeographic and paleotectonic implications. Geological Society of America, Special Paper 316, 1-134. Liu, H., Rigby, J.K., 1992. Diagenesis of the Upper Permian Jiantianba Reef, West Hubei, China. Journal of Sedimentary Petrology 62 (3), 367-381. Loeblich,A., Tappan, H., 1964. Sarcodina chiefly 'Thecamoebians' and Foraminiferida. In: Moore, R.C. (Ed.), Treatise on Invertebrate Paleontology, part C, Protista 2. Geological Society of America and University of Kansas Press, 2 volumes, 900 pp. Mamet, B., Boulvain, F., 1992. Microflore des monticules micritiques frasniens 'F 2j' de Belgique. Revue de Micropal4ontologie 35 (4), 283-302. Mamet, B., Preat, A., 1992. Algues du D6vonien moyen de Wellin (synclinorium de Dinant, Belgique). Revue de Micropal6ontologie 35 (1), 53-75. Mamet, B., Roux, A., 1975. Dasycladac6es d6voniennes et carbonif6res de la T6thys occidentale. Revista Espafiola de Micropaleontologia 7 (2), 245-295. Mamet, B., Roux, A., Nassichuk, W.W., 1987. Algues carbonif'eres et permiennes de l'Arctique Canadien. Bulletin Geological Survey of Canada 342, 1-83.

402 Mamet, B., Rudloff, B., 1972. Algues carbonif'eres de la partie septentrionale de l'Am6rique du Nord. Revue de Micropa16ontologie 2, 75-114. Maslov, V.P., 1929. Mikroskopicheskie vodorosli kamennougolnykh izvestnyakov Donetskogo basseina (Microscopical algae from Carboniferous limestones of Donetz). Bulletin of Geological Comity 48 (10), 115-138 [in Russian]. Maslov, V.P., 1939. A discovery of green algae in the Paleozoic of Kazakhstan. Problems of Paleontology 5, 290-296 [in Russian with English abstract]. Maslov, V.P., 1956. Iskopaemye izvestkovye vodorosli SSSR (Fossil calcareous algae from USSR). Akademiya Nauk SSSR, Trudy Instituta Geologicheskikh Nauk 160, 301 pp. [in Russian; French translation: B.R.G.M. n ° 3517]. Massa, D., Vachard, D., 1979. Le Carbonif~re de Libye occidentale, biostratigraphie et micropal6ontologie, position dans le domaine t6thysien d'Afrique du Nord. Revue de l'Institut Franfais du P6trole 34 (1), 3-66. Miklukho-Maclay, A.D., 1958. Novoe semeistvo foraminifer, Tuberitinidae M.-Maclay fam. nov. (A new family of foraminifers, Tuberitinidae M.-Maclay fam. nov.). Voprosy Mikropaleontologii, 2:130-135 [in Russian; French translation: Editions Technip Paris 1960, 155-159]. Milanovic, M., 1965. Salopekiella, novi rod familije Dasycladaceae iz permskikh sedimentata Velebita (SalopekieUa, a new genus of the Dasycladaceae family from the Permian sediments of the Velebit range). Acta Geologica, Zagreb 5, 373-382 [in Serbo-Croate with English abstract]. Milanovic, M., 1966. Likanella, a new Permian genus of the family Dasycladaceae. Geoloski Vjesnik 19, 9-14. Milanovic, M., 1974. Kochanskyella (Chlorophyta, Dasycladaceae), a new Permian genus of Mt. Velebit, Croatia. Geoloski Vjesnik 27, 127-132. Montenat, C., Lapparent, A.F de., Lys, M., Termier, H., Termier, G., Vachard, D., 1976. La transgression permienne et son substratum dans le Jebel Akhdar (Montagnes d'Oman, P6ninsule Arabique). Annales de la Soci6t6 g6ologique du Nord 96 (3), 239-258. Monty, C.L., 1982. Cavity or fissure dwelling stromatolites (endostromatolites) from Belgian Devonian mud mounds. Annales de la Soci6t6 g6ologique de Belgique 105, 343-344. Monty, C.L., 1984. Stromatolites in earth history. Terra Cognita 4, 423-430. Monty, C.L., 1986. Interactions 6v6nements g6ologiques-stromatolithes. Bulletin des Centres de Recherche Exploration-Production Elf-Aquitaine 10 (2), 537-553. Moshier, S.O., Kirkland, B.L., 1993. Identification and diagenesis of a phylloid alga: Archaeolithophyllum from the Pennsylvanian Providence Limestone, Western Kentucky. Journal of Sedimentary Petrology 63 (6), 1032-1041. Mu Xinan, 1981. Upper Permian algae from western Guizhou. Acta Palaeontologica Sinica 20 (1), 34-48. Mu Xinan, 1982. Some calcareous algae from Xizang. In Paleontology of Xizang. The Series of the Scientific Expedition to the Qinghai-Xizang Plateau, Science Press, Beijing 5, 205-240. Nguyen Lan Tu, 1970. Some Permian fossil algae from Vietnam, Cambodia and Laos. Archives g6ologiques du Viet-Nam 13 (2), 1-42. Okla, S., 1992. Permian algae and algal microfacies from Unayzah, Quassim District, Saudi Arabia. 27, 217-224. Perret, M.F., Vachard, D., Aguirre, P., Crasquin-Soleau, S., 1994. Micropal6ontologie des calcaires 6pibathyaux h Globochaete (algue probl6matique) du Carbonif~re des Pyr6n6es. Geobios 27 (6), 659-675. Peters, T., Hauser, M., Moser, L., Immenhauser, A., Blasi, H.P., Bataschi, M., Raijh, A.S., 2000. Geological maps of Sur and A1 Ashkahrah sheets NF 40-8F and 40-12C, scale 1:100'000 with explanatory notes. Directorate General of Minerals, Oman Ministry of Industry and Commerce, Muscat. Petryk, A.A., Mamet, B.L., 1972. Lower Carboniferous algal microflora, southwestern Alberta. Canadian Journal of Earth Sciences 9 (7), 767-802. Pia, J. von, 1912. Neue Studien fiber die triadischen Siphoneae Verticillatae. Beitr~ige zur Pal~iontologie und Geologie 0sterreich-Ungarns, 25: 25-85. Pia, J. von, 1920. Die Siphoneae Verticillatae vom Karbon bis zur Kreide. Abhandlungen der zoologisch-botanischen Gesellschaft in Wien 11, 1-263 [French translation by Gubler M.

1961. Les Siphon6es Verticill6es du Carbonif'ere au Cr6tac6. Editions Technip, Paris, 236 pp.]. Pia, J. von, 1937. Die wichtigsten Kalkalgen des Jungpal~iozoikums und ihre geologische Bedeutung. Comptes Rendus du 26me Congr~s de l'Avancement des Etudes Stratigraphiques sur le CarboniFere, Heerlen 1935, 2, 785-856. Pillevuit A., Marcoux J., Stampfli G. & Baud A. 1997. The Oman Exotics: a key to the understanding of the Neotethyan geodynamic evolution. Geodinamica Acta 10 (5), 209-238. Pokorny, V., 1951. The Middle Devonian foraminifera of Celechovice, Czekoslovakia. Vest. Kral. C. Spol. Nauk, Trida Mat.Prirod. 9, 1-29. Poncet, J., 1987a. Cr6ation du genre Californiella nov. gen. (Rhodophycophyta) et r6vision du genre Dasyporella STOLLEY,1893 (Dasycladales). Geobios 20 (6), 837-842. Poncet, J., 1987b. Eoclypeina, un nouveau genre d'algue verte calcaire (Dasycladaceae) dans le Carbonif'ere sup6rieur du bassin de B4char (Sahara alg6rien) avec Eoclypeina crassa nov. sp. Revue de Micropal6ontologie 30 (2), 117-121. Poncet, J., 1989. Pr6sence du genre Halimeda LAMOUROUX,1812 (Algue verte calcaire) dans le Permien sup4rieur du Sud Tunisien. Revue de Micropal6ontologie 32 (1), 40-44. Praturlon, A., 1963. Dasycladaceae from the Upper Permian of the Dolomites (Italy). Geologia Romana 2, 119 -150. Pronina, G.P., 1996. Genus Sphairionia and its stratigraphic significance. Reports of Shallow Tethys 4, International Symposium Albrechtsberg (Austria) 8-11 September 1994, Supplemento agli Musei Civici di Rovereto, Sezione Archeologia, Storia e Scienze Naturali 11, 105-118. Racz, L.G., 1965. Carboniferous calcareous algae and their associations in the San-Emiliano and Lois-Ciguera Formations (Province Leon, NW Spain). Leidse Geologische Mededelingen 31, 1-112. Racz, L.G., 1966. Late Paleozoic calcareous algae in the Pisuerga basin (N Palencia, Spain). Leidse Geologische Mededelingen 31, 241-260. Racz, L.G., 1984. Iberiaella, a new fossil alga from the Middle Carboniferous of NW Spain. Geologie en Mijnbouw 63, 333336. Racz, R., 1959. Permian algae from Saudi Arabia. Journal of Paleontology 33 (4), 531-539. Radoicic, R., 1959. Some problematic microfossils from the Dinarian Cretaceous. Serbia Zavod. Geol. i Geofiz. Istraz., Vesnik 17, 87-92 [in Serbo-Croate with English abstract]. Ranser-Chernoussova, D.M., Koroliuk, I.K., 1981. K morfologii i sistematike pozdnemoskovskikh sifonovykh vodoroslei yuzhnogo Urala ikh ob v rodi b porodoobrazovanii (On the morphology and systematics of Late Moscovian siphonal algae in the south Urals and their significance in rock formation). Voprosy Mikropaleontologii 24, 157-170 [in Russian]. Razgallah, S., Vachard, D., 1991. Syst6matique et bios6dimentologie des algues constructrices permiennes Tubiphytes et Archaeolithoporella suivant l'exemple du Jebel Tebaga (Murghabien de Tunisie). Palaeontographica B, 221 (5-6), 171-205. Reitlinger, E.A., 1950. Foraminifery srednekammenougolnykh otlozhenii tsentralnoi yasto Russkoi platformy (isklyuchaya semeistvo Fusulinidae) [Foraminifers from the deposits of the Middle Carboniferous of the central part of the Russian platform (with exception of the family Fusulinidae)]. Akademiya Nauk SSSR, Trudy Geologischeskii Institut, 126, seriya geologichevskaya 6, 1-126 [in Russian]. Renz, C., Reichel, M., 1945. Beitr~ige zur Stratigraphie und Pal~iontologie des ostmediterranen Jungpal~iozoikums und dessen Einordnung im griechischen Gebirgssystem. Eclogae Geologicae Helvetiae 38 (2), 211-270. Riding, R., 1991. Calcified cyanobacteria. In: Riding, R. (Ed.), Calcareous algae and stromatolites. Springer Verlag, Berlin, 55-87. Riding, R., 1993. Shamovella obscura: the correct name for Tubiphytes obscurus (Fossil). Taxon 42, 71-73. Riding, R., Guo, L. 1991. Permian marine calcareous algae. In: Riding, R. (Ed.), Calcareous algae and stromatolites. Springer Verlag, Berlin, 452 480. Riding, R., & Guo, L. 1992. Affinity of Tubiphytes. Palaeontology, 35 (1), 37-49. Roger, J., B6chennec, F., Janjou, D., Le Metour, J., Wyns, R., Beurrier, M., 1991. Geological map of Ja'alan sheet NF 40-8E, scale 1:100'000 with explanatory notes. Directorate General

403 of Minerals, Oman Ministry of Petroleum and Minerals, Muscat. Roux, A., 1985. Introduction ~ l'6tude des algues fossiles pal6ozoiques (de la bact4rie ~ la tectonique des plaques). Bulletin des Centres de Recherches Exploration-Production Elf-Aquitaine, 465-687. Roux, A., 1991. R6vision des Gymnocodiaceae (Algues rouges, Permien-Cr6tac6), taxonomic, biostratigraphie, pal6obiog6ographic; 2e partie: inventaire taxonomique critique des esp~ces de Gymnocodiac6es du Permien et du Trias. Revue de Micropal6ontologie 34 (2), 136-173. Roux, A., Deloffre R. 1990. R4vision des Gymnocodiaceae (Algues rouges, Permien-Cr6tac6), taxonomic, biostratigraphie, pal6obiog6ographie; 16re partie: g6n6ralit6s sur la famille. Revue de Micropal6ontologie 33 (2), 123-137. Saltovskaya, V.D., 1979. Novyi rod izvestkovykh vodoroslei iz srednego Karbona Tadzhikistana (New genus on calcareous algae from Middle Carboniferous of Tadzhikistan). Izvestiya Akademiya Nauk Tadzhiskoi SSR, Otdel. Biol. Nauk 74 (1), 88-93 [in Russian]. Saltovskaya, V.D., 1984. Nekotorye izvestkovye vodorosli Paleozoya Tadzhikistana (Some calcareous algae of the Paleozoic from Tadzhikistan). In: Novye vidy iskopaemoi flori i fauny Tadzhikistana (New species of fossil flora and fauna from Tadzhikistan). Akademiya Nauk Tadzhiskoi SSR, Institut Geologii, Dushanbe, 141-160. Samankassou, E., 1997. Palaeontological response to sea-level change: distribution of fauna and flora in cyclothems from the lower Pseudoschwagerina limestone (latest Carboniferous, Carnic Alps, Austria). Geobios 30 (6), 785-796. Samankassou, E., 1998. Skeletal framework mounds of Dasycladaceae alga, Anthracoporella, Upper Paleozoic, Carnic Alps, Austria. Palaios 13, 297-300. Sano, H., Horibo, K., Kumamoto, Y., 1990. Tubiphytes-Archaeolithoporella-Girvanella reefal facies in Permian buildup, Mino terrane, central Japan. Sedimentary Geology 68, 293-306. Schfifer, P, Senowbari-Daryan, B., 1983. Die Kalkalgen aus der Obertrias von Hydra, Griechenland. Palaeontographica B, 185 (4-6), 83-142. Schmid, D.U., Leinfelder, R.R., 1995. Lithocodium aggregatum ELLIOTTn'est pas une algue mais un foraminWere encrofitant, commensalis6 par le foraminif~re Troglotella incrustans WERNLI& FOOKES.Comptes Rendus de l'Acad6mie des Sciences, Paris 330, 2a, 531-538. Schreurs, G., Immenhauser, A., 1999. WNW-directed obduction of the Batain Group on the E-Oman continental margin at the Cretaceous-Tertiary boundary. Tectonics 18 (1), 148-160. Schubert, R.J., 1908. Zur Geologic der 5sterreichischen Velebit (nebst pal~iontologischem Anhang). Jahrbuch der geologischen Reichsantalt 58 (2), 347-382. Sebbar, A., Mamet, B., 1996. Algues benthiques calcaires du Carbonif'ere inf6rieur et moyen, Bassin de B6char, Alg6rie. Revue de Micropal6ontologie 39 (2), 153-167 Senowbari-Daryan, B., Flfigel, E., 1993. Tubiphytes MASLOV,an enigmatic fossil: classification, fossil record and significance trough time, part I: Discussion of Late Paleozoic material. In: Barattolo, F., de Castro, P., Parent, E M. (Eds),'Studies on fossil benthic algae. Bollettino della Societ~ Paleontologica Italiana, spec. vol. 1, 353-382. Sha Jin-geng, Zhang Lin-xin, Luo Hui, Xu Shan-hong, Bao Huiruing, 1992. On the closure age of the Late Paleozoic rift in Hohxil, Qinghai. Acta Micropalaeontologica Sinica 99 (2), 177182 [in Chinese with English abstract). Shackleton, R.M., Ries, A.C., Bird, A.C., Filbrandt, J.B., Lee, C.W., Cunningham, G.C., 1990. The Batain Melange of NE Oman. In: Robertson, A.H.F., Searle, M.P., Ries, A.C. (Eds), The geology and tectonics of the Oman regions. Geological Society of London 49, 673-696. Sheng, J.Z., 1992 - Development of fusuline foraminifers in China. Studies in Benthic Foraminifera, Benthos' 90, Sendai 1990, Tokai University Press, 11-22. Shuysky, V.P., 1973 - Izvestkovye rifoobrazuyuchshie vodorosli nizhnego Devona Urala (Reef-building calcareous algae from Early Devonian of Urals). Akademiya Nauk SSSR, Uralskii Nauchnyi Tsentr, Institut Geologii i Geokhimii, 1-155. Shuysky, V.P., 1986. Novyi predstavitel dazikladievykh vodoroslei iz nizhnego Devona Urala (New representatives of Dasycladale algae from the Early Devonian of Urals). Paleontologicheskii Zhurnal 1986, 2, 118-122 [in Russian).

Shuysky, V.P., 1987. Lopsiella mansica, novyi predstavitel atsetabulyarievykh vodoroslei iz givetskikh otlozhenii Urala (Lopsiella mansica, new representative of Acetabulariaceae algae from the Givetian deposits of Urals). Paleontologicheskii Zhurnal 1987, 3:67-75 [in Russian]. Shuysky, V.P., Patrunov, D.K., 1991. Calcareous algae of the lower and Middle Devonian of the southern part of Novaya Zemlya islands. Nauka, Moscow: 1-80 [in Russian). Skompski, S., 1982. The nature and systematic position of the microfossils Globochaete alpina LOMBARD,1945. Acta Geologica Polonica 32 (1-2), 47-66. Skompski, S., 1996. Stratigraphic position and facies significance of the limestone bands in the subsurface Carboniferous succession of the Lublin Upland. Acta Geologica Polonica 48 (3-4), 171- 268. Stolley, E., 1893. Ueber silurische Siphonen. Neues Jahrbuch ffir Mineralogie 2, 135-146. Tellez-Giron, C., Nestell, M., 1983. Microfacies y zonificaci6n del P6rmico de Las Delicias, Coahuila, M6xico. Revista del Instituto Mexicano del Petroleo 15 (3), 7-23. Termier, H., Termier, G., 1973. Stromatopores, Scl6rosponges et Phar6trones, les Ischyrosponges. Annales des Mines et de la G6ologie, Tunis, Livre Jubilaire Marcel Solignac, 26, 285-297. Termier, H., Termier, G., Vachard, D., 1975. Recherches micropal6ontologiques dans le Pal6ozoique sup6rieur du Maroc Central. Cahiers de Micropal6ontologie 4, 1975, 1-99. Termier G., Termier, H., Vachard, D., 1977. Etude comparative de quelques Ischyrosponges. G6ologie M6diterran6enne 4 (2), 139-180. Tien Nguyen Duc, 1979. Etude micropal6ontologique (Foraminif6res) de mat6riaux du Permien du Cambodge. Th~se Universit6 Paris Sud, Facult6 d'Orsay, 167 pp. Tien Nguyen Duc, 1987. Sphairionia sikuoides gen. nov. sp. nov., a Permian incertae sedis organism. 11e Congr6s International de G6ologie du Carbonif'ere, Beijing 1987, Abstracts, 90. Tien Nguyen Duc, 1988. Note on two 'incertae sedis' from the Permian of West Thailand. CCOP Technical Bulletin 20, 109111. Tien Nguyen Duc, 1989. Sphairionia sikuoides nov. gen nov. sp., a Permian incertae sedis organism. 11e Congr6s International de G6ologie du Carbonif~re, Beijing 1987, Compte Rendu 3, 73-78. Torres, A.M., Baars, D.L., 1992Anchicodium JOHNSON:branched or phylloid? Journal of Paleontology 66, 675-677. Trifonova, E., 1985. Upper Permian foraminifers from south-eastern Lakonia (Peloponnessus, Greece). Geologica Balcanica 15 (4), 83-92. Vachard, D., 1980. T6thys et Gondwana au Pal6ozo~que sup6rieur, les donn6es afghanes, biostratigraphie, micropal6ontologic, pal6og6ographie. Documents et Travaux IGAL 2 vol., 463 pp. Vachard, D., 1985. Remarques sur les Dasycladales (Algues vertes) du Permien sup6rieur du Jebel Tebaga. Acres du Premier Congr6s National des Sciences de la Terre, Tunis 1, 271-281. Vachard, D., 1993. Algues, pseudo-algues et microfaci6s carbonat6s du D6vonien du Domaine Lig6rien (MassifArmoricain, France). Palaeontographica B, 229 (1-3), 53-113. Vachard, D., 1994. Foraminif'eres et Moravamminides du Giv6tien et du Frasnien du Domaine Lig6rien (MassifArmoricain, France). Palaeontographica A, 231 (1-3), 1-92. Vachard, D., Argyriadis, I., 2000. Pr6cisions biostratigraphiques sur le Permien m6sog6en, de l'Italie ~ la Turquie. M6moires de l'Association des G6ologues du Permien, Livre comm6moratif Nad6ge Toutin-Morin (in press). Vachard, D., Beckary, S., 1991. Algues et foraminif~res bachkiriens des coal balls de la Mine Rosario (Truebano, Leon, Espagne). Revue de Pal6obiologie 10 (2), 315-357. Vachard, D., Berkhli, M., 1992. Importance des coupes du bassin de J6rada (Maroc) pour la connaissance du Vis6en terminal. Revue de Micropal6ontologie 35 (4), 307-328. Vachard, D., Ferri6re, J., 1991. Une association ~ Yabeina (foraminif~re fusulinoi'de) dans le Midien (Permien sup6rieur) de la r6gion de Whangaroa (Baie d'Orua, Nouvelle-Z41ande). Revue de Micropal6ontologie 34 (3), 201-230. Vachard, D., Fourcade, E., Mendez, J., Cosillo, A., Alonzo, M., Requena, J., Azema, J., Cros, P., 1997. Foraminif~res et algues du Permien du Guatemala. Geobios 30 (6), 745-784.

404 Vachard, D., Gargouri-Razgallah, S., Chaouachi, M.C., 1989. Sur les biohermes ~ algues Sol~noporac6es et phyllo~des du Permien sup4rieur de Tunisie (Murghabien du Djebel Tebaga) et sur les incidences de la diagen~se carbonat6e sur la syst6matique algaire. Revue de Pal4obiologie 8 (1), 121-141. Vachard, D., Krainer, K. Biodiversity of smaller Foraminifera and some other carbonate components from Auernig to Upper Pseudoschwagerinenkalk Formations in Carnic Alps (Late Carboniferous-Early Permian, Austria). Rivista Italiana di Paleontologia i Stratigrafia (in preparation). Vachard, D., Martini, R., Zaninetti, L., Zambetazkkis-Lekkas, 1993. R4vision micropal6ontologique (foraminif~res, algues) du Permien inf4rieur (Sakmarien) et sup~rieur (Dorashamien) du Mont Beletsi (Attique, Gr6ce). Bollettino della Societ~ Paleontologica Italiana 32 (1), 89-112. Vachard, D., Maslo, A., 1996. Pr6cisions biostratigraphiques et micropal6ontologiques sur le Bashkirien d'Ukraine (Carbonif'ere moyen). Revue de Pal6obiologie 15 (2), 357-383. Vachard, D., Miconnet, P., 1989. Une association ~ Fusulinoides du Murghabien sup6rieur au Monte Facito (Appennin m6ridional, Italie). Revue de Micropal6ontologie 32 (4), 297-318. Vachard, D., Montenat, C., 1981. Biostratigraphie, micropal6ontologie et pal4og6ographie du Permien de la r6gion de Tezak (Montagnes Centrales d'Afghanistan). Palaeontographica B, 178 (1-3), 1-88. Vachard, D., Razgallah, S., 1993. Discussion sur l'~ge murgabien ou midien des s4ries permiennes du Jebel Tebaga (Sud Tunisien). Rivista Italiana di Paleontologia i Stratigrafia 99 (3), 327-356. Vachard, D., Termier, G., Termier, H., 1978. Sur l'appartenance des algues Sol6noporac4es. Comptes Rendus de l'Acad6mie des Sciences, Paris 286, 1865-1868. Vermin, E., Vachard, D., Proust, J.N., 1997. Taphonomie et syn6cologie du 'genre' Tubiphytes dans les bioconstrutions de Tratau et de Nizhni-Irginsk (Permien inf4rieur de l'Oural, Russie). Geobios 30 (5), 635-649. Wahlman, G.P., 1988. Subsurface Wolfcampian (Lower Permian) shelf-margin reefs in the Permian Basin of West Texas and southeastern New Mexico. Midcontinent SEPM, sp. publ. 1,

177-204. Wang, S.H., Fan, J.S., Rigby, J.K., 1994. Archaeolithoporella and Tubiphytes: affinities and paleoecology in Permian reefs of South China. Science in China B, 37 (6), 723-743. Wood, A., 1948. Sphaerocodium, a misinterpreted fossil from the Wenlock Limestone. Proceedings Geological Association London 59, 9-22. Wood, A., 1964. A new dasycladacean alga, Nanopora, from the lower Carboniferous of England and Kazakhstan. Palaeontology 7 (2), 181-185. Wood, R.A., Dickson, J.A.D., Kirkland-George, B., 1994. Turning the Capitanian Reef upside down: a new appraisal of the ecology of the Permian Capitan Reef, Guadalupe Mountains, Texas and New Mexico. Palaios 9, 422-427. Wyns, R., B4chennec, F., Le Metour, J., Roger, J. & Chevrel, S. 1992. Geological map of Sur sheet NF-40-08, scale 1:250'000 with explanatory notes. Directorate General of Minerals, Ministry of Petroleum and Minerals, Muscat. D. VACHARD Universit6 des Sciences et Technologies de Lille U.F.R. des Sciences de la Terre, UPRESA 8014 du CNRS Laboratoire de Pal6ontologie du Pal6ozo~que F-59655 Villeneuve d'Ascq Cedex E-mail: Daniel.Vachard@univ-lille 1.fr M. HAUSER, A. MATTER & To PETERS Mineralogisch-petrographisches Institut und geologisches Institut, Universitfit Bern Baltzerstrasse 1 CH-3012 Bern E-mail: [email protected]

R. MARTINI & L. ZANINETTI D6partement de G4ologie et Pal4ontologie 13 rue des Maralchers CH-1211 Gen6ve E-mail: [email protected]