Encrusting foraminifera from the miocene reefs of Sinai, Egypt: A significant paleobiogeographic affiliation

GeoResJ 13 (2017) 134–158

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Encrusting foraminifera from the miocene reefs of Sinai, Egypt: A significant paleobiogeographic affiliation Wafaa I. Shahat Geology Department, Zagazig University, 44519 Zagazig, Sharkia, Egypt

a r t i c l e

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Article history: Received 13 November 2016 Revised 24 January 2017 Accepted 27 March 2017 Available online 13 April 2017 Keywords: Encrusting foraminifera Water connection Miocene Early burdigalian, Sinai West pacific

a b s t r a c t First attention is offered to encrusting foraminifera existing in the Miocene reefal deposits of Wadi Gharandal (Sinai, Egypt). The detected encrusting foraminifera are confined to typical reefal limestone development. They belong mainly to the acervulinid, planorbulinid and homotrematid groups; dominated entirely by Tayamaia and Gypsina. Moreover, other forms include Neoplanorbulinella, Planolinderina, Borodinia, Discogypsina, Ladoronia, Sphaerogypsina and Sporadotrema. Besides, they inhabit different paleoenvironments; reef-flat, fore-reef and back-reef lagoonal conditions. This encrusting assemblage shows close paleobiogeographic affinity to the West Pacific region that locates at the same paleolatitudinal position. Consequently, such strong affiliation and copious faunal exchange certainly require direct and short distance connection and water inroad of an assemblage likely indicative for warm temperate to tropical settings. Therefore, the prior marine connection that dubiously proposed by Rögl (1999) in the Early Oligocene, extending north of India, west-east direction from eastern Mediterranean passing through east Iran and expanded directly across Asia to West Pacific is proposed to be the best direct and shortest water connection to the W. Pacific realizing this rigorous faunal similarity. On consequence, this connecting sea is thought to continue open even during Aquitanian and its closure had started in Early Burdigalian time. This closure is synchronous with the analogous restriction of the central basins of Iran which is considered the entry passage to W. Pacific across the proposed connecting sea. The results significantly provide an evidence for interruption also during the Early Burdigalian. © 2017 Elsevier Ltd. All rights reserved.

1. Introduction In Egypt, most efforts for studying the encrusting fossil reefal associations were greatly interested in the algal coralline assemblages ’the nullipores’. The term nullipores was first reported in Sinai (Wadi Gharandal) by Moon and Sadek [78]. Afterwards, the nullipores were dealt broadly for their prevalence and striking association with corals in the Middle Miocene reef accumulations by Souaya [107], Dullo et al. [37], Piller and Rasser [87], Rasser and Piller [93], Imam and Refaat [59] and Hamad [46]. Furthermore, during a detailed planktonic and benthonic foraminiferal biostratigraphic delineation of some Miocene exposures in Wadi Gharandal (Sinai, Egypt), it is observed, strictly plentiful encrusting and attached types of foraminifera are distributed among the reefal type assemblages. Encrusting foraminifera constitute a group of organisms growing and adapting for attachment or cementation on various

Abbreviations: C.N., Crossed Nicols; Gh, Gharandal; PPL, Plane Polarized Light; W. Pacific, West Pacific. E-mail address: [email protected] http://dx.doi.org/10.1016/j.grj.2017.03.001 2214-2428/© 2017 Elsevier Ltd. All rights reserved.

substrates. Within intricate structures of modern and ancient reefs individuals of encrusting foraminifera live either exposed or cryptic [71]. The cryptic types are getting hide either in (i) the lower surfaces of corals [14], (ii) cavity dwellers in and under coral rubble [29,41,42,77,92], and (iii) inside tubular chambers of another foraminifera in the deep sea [44]. The substrates of encrusting foraminifera are commonly hard skeletons of invertebrates (corals, bivalvia, echinoids and barnacles), decayed matter, sea grass, manganese nodules, debris material and others [7,66,68,73,79,80,95]. On such substrates, the encrusting foraminifera are found dwelling diverse bathymetries ranging from shallow to deep seas. Stratigraphically, encrusting foraminifera are first known in the Late Ordovician. Their individuals have limited stratigraphic ranges, but some may extend and/or crowd the modern reef habitats. On modern or ancient reefs, the encrusting foraminiferal assemblages demarcate thoroughly different depth distributional patterns and actual variations along bathymetric gradients [14,72,79,86,93,112]. In modern environments, most experimental case studies have provided valuable information about this perception. This reefal type of foraminifera is considered as a secondary supplier in the reef building process and acts as frame-binder [14,71].

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Fig. 1. Geographical location of the studied section in the western extremes of Wadi Gharandal, Sinai.

In comparison with the ancient records, the modern reefal systems show a global distribution, e.g. Sinai and the W. Pacific island’s occurrences. Accordingly, the reefal encrusting formainefera signify paleoecological and paleogeographical important implications. In Sinai, encrusting foraminifera, the subject of the present study, are a principal component of Miocene reefal constitutions. The encrusting foraminifera reveal accompanying intergrowths with other encrusting organisms including coralline red algae and bryozoans, with bivalves, echinoids, and barnacles. These taxonomically variant taxa are known to live together forming a symbiotic ’consortium’. Scholle and Ulmer–Scholle [102] reported the consortium for the Late Paleozoic and some Mesozoic reefal deposits. On the other hand, the intended Miocene reefal interval in Sinai shows another younger record of such livelihood. As no attention offered before to the Miocene encrusting foraminifera of Sinai, this paper focuses on studying this type of reefal taxa including paleoecological and/or paleogeographical clarifications in the fossil reef reconstruction. Moreover, their identification, taxonomic classification, description and stratigraphic and geographic distributions are studied. Such reefal taxa are abundant and, accordingly, their role in reef building process ought not to be undervalued.

2. Study area and geologic setting Wadi Gharandal is one of the famous valleys in Sinai. It starts up at El Tih escarpment near the center of Sinai and extends widely across ridges of Eocene–Cretaceous rocks [101]; westward direction, it runs narrower and ends its course facing Ras Lagia at the Gulf of Suez (Fig. 1). The ground surface of Wadi Gharandal area is covered with adjacent, well exposed, moderate and low relief successions. In this location, facies generally exhibit a monotonous character, mostly formed of soft clastics, and few successions are seemed different. One of the dissimilar successions exhibits reefal facies criteria and has been selected for the present study.

The selected succession is located in the southern flank of Wadi Gharandal, near the west extremes of the valley (Fig. 1) at longitude 32° 56΄ 10˝ E and latitude 29° 15΄ 56˝ N. It attains a total thickness of 41 m. In the field, this succession is demarcated easily due to the characteristic configuration of its embedded outcrop. It consists of closely spaced successive embankments of limestone and much fewer sandstones (Fig. 2A and B); these banks are continuous even or undulating and have small thicknesses. The bedding planes and their geometry are hardly defined in most parts. This distinctive succession (Fig. 2A–C) has been found particularly suitable for the present study; its crystallized, massive, limestone constitution is dominated with growths of in-situ attached or encrusting organisms. Encrusting foraminifera are the most prominent in successive intervals of this reefal succession. The section parts are composed of a talus slope of reef debris as fore-reef deposits, defined by their original dip and another flat to embedded limestone geometry corresponding to backreef deposits. It is an actual example of typical reefal limestone development without a true solid framework. 3. Material and methods This study is based on thin sections derived from a stratigraphic succession of special facies characters in the southern margin of Wadi Gharandal course trend. In the field, this infrequent section of the reefal structure was hardly offered the opportunity for studying its bedding geometry and limitations. The lithologic features of some markers such as conglomerate and evaporitic beds facilitate its division. A large number of rock samples is therefore collected in order to trace the barely expected change in lithology. A number of 63 rock samples are used for megascopic description and petrographical analysis and finally succeeded in constructing a stratigraphic column consisting of three successive different intervals (interval I, interval II and interval III). The hard rock matrix in the recognized succession have succeeded in preparing good thin sections but failed to allow specimens of encrusting foraminifera to be freed from their encasing sediment. Therefore,

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Fig. 2. A. The reefal limestone succession in Wadi Gharandal consists of embedded massive limestone (at center), fore-reef limestones (to the right) are talus slope of reef debris having original dip and embedded to flat back-reef limestone (to the left); B. Close up of central part showing apparent embedded, continuous, undulating, closely spaced successive thin embankments of limestone; C. Polymictic conglomerate marking the top of interval I.

this study of encrusting foraminifera is established mainly on the reliance of thin sections through two-dimensional observation and random orientation of the specimens in the thin sections. In order to obtain coordinate data, the thin sections ready in each rock sample should be close in number. A total of 80 thin sections have been prepared by the average of five thin sections of varying sizes (25 × 75 mm) and (50 × 70 mm) in each rock sample; this enabled vision of each taxon of encrusting foraminifera in different orientations for a detailed description and accurate definition. About 523 specimens of encrusting foraminifera were investigated in all thin sections; they are wholly considered either complete specimens or only parts of specimens. The vertical distribution and occurrences of the defined taxa are inserted in their enclosing intervals in the stratigraphic column of the studied succession.

This study is of two-fold, one is descriptive and the other is quantitative. For the descriptive part, the following recognitions are demanded in each thin section: (1) taxonomic identification of the encrusting foraminifera, (2) detailed description of all available morphologic features and arrangement of chambers of each defined taxon either in embryonic or adult stage, (3) each taxon is put ready to discuss, synonymous, and/or compare to ensure its definition, (4) information about the enclosing sediments through microscopic examination of the carbonate litho- and biofacies and siliciclastic grains, textures and other common inclusions, (5) the state of the individuals, either free in the matrix, attached or encrusting a substrate and the types of substrates, (6) data about associated fauna are given. For the quantitative part, the division of the studied succession into three stratigraphic intervals helped

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to make a quantitative analysis of the taxa in each interval separately and hold actual comparison between them. Data matrix is therefore arranged, containing the samples studied and examined as thin sections in each interval and displaying the number of individuals of each identified taxon in thin sections. From the total number of individuals in the succession, the percentage of the recorded individuals in each interval (relative abundance) is calculated. Also for each interval, the following quantitative data are provided: (1) the number of individuals and the percentage of abundance of each defined taxon, (2) the percentage of taxa relating to each recorded family of encrusting foraminifera and the percentage of abundance of the individuals of the recorded families in the total succession and in each interval separately. Results are made noticeable by scheming in tables, pie charts, and columns. All the defined taxa of encrusting foraminifera are illustrated in Figs. 9–21. The used morphological terms and the generic and suprageneric classification adopted follow that of Loeblich and Tappan [69] with some modifications and observations according to recent literature. The age diagnostic species of the encrusting foraminifera are used with the aid of another associated index species of larger and planktonic foraminifera to make accurate dating for the studied intervals. A detailed worldwide geographic tracking for the defined species across all available literature is gained and arranged in a table (Table 3) then exemplified in Stacked Venn smart Art Graphic to emphasize gradational relationship. The taxonomic classifications, descriptions, and remarks of the defined encrusting foraminifera from Sinai, Egypt are joined in electronic attachment file. All specimens of rock samples and thin sections have been deposited in the collections of the Department of Geology, Faculty of Science, Zagazig University, Egypt. 4. Results 4.1. Stratigraphic succession and age determination The field data accompanied with a description of the rock samples, petrographical examination and microbio- and lithofacies investigations of multiple thin sections enabled the possible division of the studied succession. Mainly on the lithologic basis, the succession is divided into three different stratigraphic intervals (interval I, interval II and interval III); each interval is entirely homogeneous in its lithologic characters. The boundaries between the three intervals are sharp and eventful; they are picked up at the formation of conglomerate between interval I and interval II and at the start deposition of evaporites marking the change to interval III. From base to top, interval I is 8.5 m thick (Fig. 3). It is composed of highly compact, hard, gritty limestone and the intermittent pellet-rich bands. The limestone is massive with no stratification; generally, dark gray and yellow. Many angular sands increased upwards and caused a roughness touch. The top of interval I is marked by well consolidated, polymictic conglomerate; it is an essential constituent of this interval. The conglomerate appeared immature and poorly sorted with components ranging from coarse sand size to boulders of calcareous lithology (limestone) and other lithoclasts. The clasts are subangular to subrounded, darker in tone than the surrounding material and embedded in a very fine-grained sandy matrix. Encrusting foraminifera are concentrated only in the upper level of interval I (bed no.1, sample no.4). They are represented by Ladoronia vermicularis (Hanzawa) and Planolinderina escornebovensis Freudenthal association with the dasycladacean Cymopolia saipania Johnson. Abundant Amphistegina spp., A. bohdanowiczi Bieda and Miolepidocyclina panamensis (Cushman) are other associated larger foraminifera. Few plank-

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tonic foraminifera comprise Globigerina praebulloides Blow and Globigerinoides primordius Blow and Banner are important accompanying taxa. Other much fossil debris and fragments of echinoids, bivalvia, gastropoda and bryozoans are also present. The Aquitaine Basin (France) showed the first description of P. escornebovensis in Oligocene (Chattian) sediments [39] and in the Oligocene of Turkey [81]. Amphistegina bohdanowiczi Bieda ranges from Late Oligocene to Early Miocene in Zagros Basin, southwest Iran [8], and in the Venetian area, north-east Italy [9]. Miolepidocyclina panamensis (Cushman) was described from the Oligocene [31] to Early Miocene [69] in East Caribbean Sea and California, USA, respectively. Johnson [63] described the holotype C. saipania from the Early Miocene (Aquitanian) of Saipan (Mariana Islands, W. Pacific). The planktonic foraminifera Globigerina praebulloides Blow is a long-ranging species from Late Oligocene to Pleistocene characterizes planktonic foraminifera Zone P22/ N22 [64]. Globigerinoides primordius Blow and Banner is an index Late Oligocene (Chattian) species marking Zone N3; attributed to 23.50Ma in the standard tropical and subtropical biozone, biochron [45]. From above, most of the recorded fauna in interval I are index Late Oligocene species, others extend their ranges to the Early Miocene (Aquitanian). Therefore, Late Oligocene-Early Miocene (Aquitanian) age is indicated for interval I in the studied area. Interval II is 12 m thick, started with a reefal development in the form of small bioherms; locally, grown up above rocky substrate of the former conglomeratic heap and enclose crowded intergrowths of predominant encrusting foraminifera mixed with algae, bryozoans, mollusca and echinoids; all lived in a consistent assemblage. These deposits give rise to embedded, successive undulating embankments, made of massive, crystallized, occasionally dolomitic reefal limestone and identify interval II. In addition, gypsum thin bands and intermittent highly sandy and/or pellet-rich embankments interrupt this interval at its upper levels. A sand rich bank (bed no. 7) is deposited in the upper level of interval II. It contains abundant skeletal material cemented with calcium carbonates. Rich pellets are common in sediments that typify the upper level of interval II. The encrusting foraminifera are concentrated in two levels inside interval II; the basal level is bed no. 3 and the upper level consists of beds no. 6, 7 and 8; while completely absent in the other levels of this interval. In the present study, encrusting foraminifera species reach their maximum diversity in interval II; more than 90% of the detected species are concentrated in this interval, occupying its basal and upper levels. In addition, the relative abundance of the recorded individuals of encrusting foraminifera within intervals II is the greatest (83.2%). Tayamaia marianensis (Hanzawa) is the most prominent encrusting foraminifera species in interval II, its individuals attained their maximum abundance (31.0%) in this interval. In interval II, the basal level (bed no. 3) contains the encrusting foraminifera Tayamaia marianensis (Hanzawa), Borodinia septentrionalis Hanzawa, Planolinderina escornebovensis Freudenthal, Gypsina plana (Carter), Sphaerogypsina globulus (Reuss) and Sporadotrema cylindricum (Carter). The morphologic features of tests are consisting of helmet-shaped tests of T. marianensis, hemispherical tests of S. globulus and high accreted tests of G. plana. These morphologies are concentrated in this level of interval II. In the basal level of interval II, the faunal association accompanied to encrusting foraminifera consists of larger foraminifera as Miogypsinoides dehartii (Van der Vlerk), M. complanatus (Schlumberger), Miogypsinoides sp., and Miogypsinita mexicana (Nuttall). The red algae in this level are Lithothamnium, Archaeolithothamnium taiwanensis Ishijima, A. myriosporum Johnson, Mesophyllum sancti-dionysiien Lemoine, M. pacificum Johnson and the green algae are Halimeda, Chara and Microcodium sp. Planktonic foraminifera as Globigerinoides trilobus (Reuss), G. altiaperturus

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Fig. 3. Stratigraphic column and the encrusting foraminifera assemblage at the different levels of the studied section in Wadi Gharandal.

Bolli and Globoquadrina dehiscens (Chapman, Parr and Collins) are abundant associates to the encrusting foraminifera assemblage dwelling this level. The upper level of interval II (beds no. 6, 7 and 8) is highly crowded with encrusting foraminifera T. marianensis, P. escornebovensis, Neoplanorbulinella saipanensis Matsumara, Discogypsina vesicularis Silvestri, Gypsina mastelensis Bursch, Gypsina saipanensis Hanzawa, G. plana, S. globulus and S. cylindricum. They are associated to Miogypsina intermedia Drooger, M. gunteri Cole, and Miolepidocyclina burdigalensis (Cumbel). Abundant Lithophyllum spp. and Halimeda plates are associated to encrusting foraminifera in the upper level of interval II. Bryozoans of cheilostomes and cyclostomes types and serpulids are especially abundant in the sandy beach rock near the top of interval II. This sand represents the passage to reefs attached to coast in the lagoonal condition of the next interval III. In the upper level of interval II, it is observed that stretched and expanded morphologic types of both T. marianensis and G. plana are concentrated at this level. The upper boundary of interval II is picked below the lowest gypsum and the fringing reefs of the overlying interval III. On evolutionary bases, Boudagher–Fadel [16] reported that Neoplanorbulinella and Planolinderina continued to exist during the Oligocene-Miocene boundary, but both disappeared within the Early Miocene of the Tethys. On consequence, the time duration of interval II does not go younger than the Early Miocene time. Borodinia seemed to appear only in the Aquitanian of the Indo-Pacific

[16] and T. marianensis is known in conspicuous abundance in the Aquitanian limestone of W. Pacific [49]. Regarding associated species of larger foraminifera, M. dehartii is detected in Aquitanian deposits in its type locality and placed entirely in the Aquitanian [48,111]; Miogypsina gunteri Cole is a Late Aquitanian species [69]. In Miogypsinoides-Miogypsina evolutionary lineage, Miogypsinoides dehartii shows gradual evolution to Miogypsina indonesiensis/ antillea from Early Miocene (Aquitanian) to Middle Miocene [111]; Miogypsina intermedia demarcates the beginning of Burdigalian in America, Aquitaine, Piedmont, Apennines and Greece [36]. In addition, the defined forms of the coralline red and green algae are known as common Early Miocene species in the Western Pacific, Saipan Islands [63]. Furthermore, Globigerinoides trilobus is an index Aquitanian species characteristic for the standard Zone N4a, attributed to 22.96Ma; while Globoquadrina dehiscens characterizes the Aquitanian Zone N4a/N4b and attributed to 22.44Ma. On the other hand Globigerinoides altiaperturus Bolli demarcates the beginning of Burdigalian time Zone N5/N6 and dated to 20.03Ma [45]. From above-mentioned information, most of the recorded fauna in interval II are index Aquitanian species, others confirm the beginning of Early Burdigalian; therefore, Early Miocene (AquitanianEarly Burdigalian) age is confirmed for interval II in the studied area. The presence of these index and zonal marker species correlate the time spanning of interval II with that of the standard IndoPacific and Atlantic tropical-subtropical biozones, biochron [45]. Interval III is thick, mainly evaporitic succession in the topmost part of the studied section. 16 m thick were measured, mainly

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Fig. 5. The percentage of taxa relating to each recorded family of encrusting foraminifera. Fig. 4. Relative abundance of the recorded individuals of encrusting foraminifera within intervals I, II and III.

made up of thick chalky white, well-bedded gypsum and anhydrite beds. In the basal part of interval III, there are mixed deposits of both lagoonal gypsum and anhydrite with fringing reefs consisting of crystallized and highly dolomitic limestone (Fig. 3). Because of the common lack of fossils in the evaporitic milieux, the exact time for the first deposition of evaporites is hardly determined. Regarding fauna defined in the fringing reefs, it is observed that some species of those defined in the just underneath bed in interval II, extend their ranges and occupy the fringing reefs in the basal part of interval III such as N. saipanensis, G. plana, S. globulus, M. intermedia and M. burdigalensis; they are particularly referring to the Early Burdigalian time. Accordingly, the deposition of interval III is started in the Early Burdigalian time.

4.2. Encrusting foraminifera In the studied succession it is observed that the recognized intervals show a noticeable change in the distribution and content of the encrusting foraminifera, both qualitatively and quantitatively. The comparative change is moderate in taxonomic composition of the recorded taxa but high in the relative abundance of the recorded individuals as 4%, 83.2% and 12.8% within intervals I, II and III respectively (Fig. 4). The abundance and the species diversity of encrusting foraminifera reach maximum in interval II, and much decreased in interval I. This disparity in species abundance and diversity from one interval to another is mainly related to the individual’s adaptation to the available environmental conditions. In the present study, individuals of the acervulinid, planorbulinid and homotrematid groups are the main constitutions of the studied encrusting foraminifera. On taxonomic basis, the taxa which belong to the family acervulinidae constitute the largest percentage where more than 60% of the determined taxa are acervulinids. This percentage is arranged as 63.6% belonged to the family acervulinidae, 27.3% belonged to the family planorbulinidae, and 9.1% are of the family homotrematidae (Fig. 5). Considering the number of individuals belonged to each family, it is found that the planorbulinid group has the largest number of individuals in the three intervals, they make 53.3%, exceeding the acervulinids which constitute 39.4% and the homotrematids which consist only 7.3% of the total investigated number of individuals (Fig. 6). The remarkable abundance of the planorbulinid group is related to the prolific existence of the planorbulinid species T. marianensis which known to occur with considerable abundance in the Aquitanian deposits [49] and in the present study, it adapts well to the prevailing environmental conditions in both levels of interval II.

Fig. 6. Relative abundance of the individuals of the recorded encrusting foraminifera families in the total succession.

4.2.1. The planorbulinid group Individuals of the planorbulinid group are the most prominent in the three intervals; they represent 57.1% in interval I, 52.9% in interval II and 52.2% in interval III (Fig. 7). Three taxa are described belonged to the planorbulinidae. Neoplanorbulinella saipanensis [74]; Fig. 9: it is concentrated mainly in the upper levels of interval II (beds no. 8) and in some fringing reefs at the base of interval III (bed no. 9). The enclosing sediment is gray, massive wholly crystallized dolomitic limestone; builds up a reefal rock with much sand input. The associated fauna are plentiful; they consist of many cheilostome bryozoans and abundant Lithophyllum spp. The larger foraminifera species associated to N. saipanensis are the encrusting G. plana, G. saipanensis, T. marianensis, D. vesicularis, S. globulus and P. escornebovensis with M. intermedia, M. gunteri, and Miolepidocyclina burdigalensis. Other associated organisms are abundant bivalvia and echinoids. Geographic distributions of N. saipanensis report its first description in the Early Miocene, Tagpochau Limestone of Saipan (Mariana Islands, W. Pacific) by Matsumaru [74]. Other recent records are from the western Tethys in Venetian area (north-east Italy; [9]) and in Philippine Archipelago [75]. Planolinderina escornebovensis [39]; Fig. 10A and B: in the studied succession, P. escornebovensis is distributed in the three intervals and absent in few levels only. The percentages of the abundance of its individuals are 57.1%, 16.3% and 19.4% in intervals I, II, and III, respectively (Table 2, Fig. 8). It occupied a stratigraphic range from Upper Oligocene to Early Miocene (Early Burdigalian). It exhibits a relatively small size and the specimens investigated in bed no. 7 show good states of preservation. Its enclosing sediment in bed no. 7 builds up a calcareous sandstone bank near the top of interval II. This sandstone describes beach rock, consisting of yellowish brown, coarse-grained, massive, with strong calcareous cement; it is enriched in coarse skeletal material of organisms living mainly on the sandy coral beaches such as bivalvia and many fragments and spines of echinoids. Well preserved bryozoans

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Fig. 8. The percentage of abundance of each defined taxon in the three intervals I, II and III.

Fig. 7. Relative abundance of encrusting foraminifera families in intervals I, II and III.

of cheilostomes and cyclostomes types, algae and serpulids are associated organisms to P. escornebovensis in the sandstone rock. Escornebéou (Aquitaine Basin, France) showed the first description of P. escornebovensis in the Oligocene (Chattian) sediments [39]. Adding to the present occurrence in Sinai, another record is by Özcan et al. [81] from the Oligocene of Mus¸ , Eastern Turkey. Tayamaia marianensis [49]; Figs. 11A–C, 12A–D and 13A–E: in the studied succession, T. marianens is the most prominent encrusting foraminifera species. It recorded its greatest abundance at two levels inside interval II, less abundant in interval III, and completely absent in interval I (Table 2; Fig. 8). The sediments enclosed T. marianens are only limestones and not detected in sandstones. It encrusted bivalvia and generally, it is characterized by the most prolific abundance of its individuals throughout the succession. The first appearance of T. marianensis is in yellowish gray, frequently sandy, crystalline, fractured, massive, embedded limestone at the base of interval II (bed no. 3). It represented the predominant element in a reefal growth in the form of small bioherms, without framework; conglomerate directly underlies the

Fig. 9. Neoplanorbulinella saipanensis Matsumaru, horizontal section showing aperture openings as foramens (arrows) (Gh8-13), PPL.

reefal construction. The general scarcity of significant coral masses is observed, but only rare fragments of the coral Porites is found in association with abundant and diversified encrusting foraminifera such as P. escornebovensis, G. plana, S. globulus, S. cylindricum and B. septentrionalis. In this level of interval II, faunal association with T. marianensis is profuse; it consists of larger foraminifera such as M. dehartii, Miogypsinoides sp., M. complanatus, M. mexicana; abundant coralline red algae as Lithothamnium spp., Mesophyllum spp., M. sanctidionysiien, M. pacificum, A. myriosporum, and A. taiwanensis; green algae Chara sp. oogonia, Microcodium sp. and Halimeda sp.. Thick encrusting cheilostome and cyclostome bryozoans, abundant large bivalvia skeletal parts, ostracod valves and abundant planktonic foraminifera species such as G. trilobus, G. altiaperturus and G. dehiscens are clearly noticeable associated to T. marianensis. All the

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A

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B

Fig. 10. Planolinderina escornebovensis Freudenthal. A. Tangential to oblique section passing with the outer wall which appeared coarsely perforate, with roughly lobulate peripheral outline (Gh1-4), C.N.; B. Horizontal section in microspheric form showing circled later chambers (Gh7-11), C.N.

recognized T. marianensis specimens at the lower level of interval II are typical helmet-shaped tests. Another important record of T. marianensis is at the upper level of interval II, (beds no. 6 and 8) (Table 1). The stretched (umbrellashaped) tests of T. marianensis are concentrated only at this level. In this level of interval II, T. marianensis represents the essential component of a typical reefal rock, built up of gray, coarsely crystallized, highly dolomitic, frequently gypseous, and pelletrich sandy limestone. The limestone is enriched in encrusting organisms: mainly encrusting foraminifera as P. escornebovensis, N. saipanensis, D. vesicularis, G. plana, G. saipanensis, G. mastelensis, and S. globulus; other larger foraminifera species as M. intermedia, M. gunteri, and Miolepidocyclina burdigalensis and abundant bryozoans (cheilostomes and cyclostomes). The red algae Lithophyllum is the most prominent algae occurred in the upper level of interval II. Coarse echinoid and bivalvia fragments are abundant. Geographic distribution of T. marianensis has been asserted W. Pacific concentration in the Lower Miocene (Aquitanian), Tagpochau Limestone of Mariana Islands (the type locality), Laderan Dago, Mulato, Tinian, Sadog Talofofo, kanat Fahang Lichan, Saipan [30,49]. other records were by Binnekamp and Belford [13] in the Lower Miocene of Central Highlands (New Guinea); by Adams and Belford [3] as a frequent species associated to M. dehartii in the Upper Tertiary e letter stage (Aquitanian) of Christmas Island (Indian Ocean); by Adams [2] in the Indo-West Pacific and by Perrin [84] from the reefal limestone of Mururoa Atoll (French Polynesia).

4.2.2. The acervulinid group In the present collection of encrusting foraminifera, the percentage of taxa relating to the acervulinid group is the highest (63.6%); they are the most diverse. At the same time, the relative abundance of its individuals in the total succession and in each interval separately occupies the second grade after the planorbulinid group (Figs. 6 and 7). The following taxa belong to the acervulinidae. Borodinia septentrionalis [48]; Fig. 14A–C: in the present investigation, only one specimen of Borodinia septentrionalis is found at the basal level of interval II (bed no. 3). It is associated with T. marianensis and its accompanying fauna of red and green algae, larger and planktonic foraminifera species at the base of this interval. It adapted to this level to environmental conditions of the reefal biohermal growth, formed directly over conglomerate.

Borodinia septentrionalis is first defined by Hanzawa [48] in the Lower Miocene (Aquitanian), subsurface of Daitô Island, E. of Ryukyu Islands (W. Pacific) and by Hanzawa [49] in the Aquitanian, Tagpochau limestone from Saipan, Tinian, Rota, and Mariana Islands (Micronesia). Other records are from: Southeast Asia (Philippines; [51]); Early Miocene Limestone of Australia and New Zealand [23–25]; Aquitanian of the Shimizu Formation (Japan; [76]); and reefal limestone of Mururoa Atoll (French Polynesia; [84]). Discogypsina vesicularis [106]; Fig. 15A–C: it is concentrated in the upper level of interval II (bed no. 8) which made up of highly dolomitic, coarsely crystallized sandy limestone, also detected in the fringing reefs at the base of interval III. It is a most frequent reefal element, played a considerable role in the reef manufacture. Fauna found associated to D. vesicularis include N. saipanensis, P. escornebovensis, T. marianensis, G. plana, G. saipanensis, S. globulus, Miogypsina intermedia Drooger, M. gunteri, and Miolepidocyclina burdigalensis. Profuse cheilostome and cyclostome bryozoans are essential components; abundant Lithophyllum, green algae, much echinoids and bivalvia are common. D. vesicularis encrusts barnacles and the substrate is indefinite for some specimens. Göes [43] recorded D. vesicularis firstly in Recent reefs of the Caribbean Sea; another Recent record is in Funafuti Atoll (South Pacific) by Chapman [22]. Other records in the geologic past include: the Eocene of Borneo (Klias Peninsula; [114]); the Aquitanian of Philippines [115]; the Upper Oligocene sediments of Great Kei (Indonesia; [18]); Micronesia [49], Saipan and Mariana Islands [30]. Gypsina mastelensis [18]; Fig. 16A and B: three species of the genus Gypsina dominate the examined assemblage of encrusting foraminifera; they are concentrated mainly in interval II (its lower and upper levels) and the base of interval III. Gypsina mastelensis recorded only in interval II (bed no. 6), where 13 specimens are defined in five thin sections; they are moderately preserved. The percentage of abundance of individuals of G. mastelensis is relatively weak (3%) compared with the other taxa in interval II (Fig. 8). In the studied section, the sediment enclosed G. mastelensis (bed no. 6) is yellow, dirty, partly dolomitic, compact, massive, hard, gritty, highly fossiliferous limestone; alternated with brittle, yellow, highly sandy, shelly bands, contain abundant echinoid and bivalvia shell fragments; abundant subangular carbonate grains are mainly peloids. The substrate is unclear. The fauna associated to G. mastelensis, arranged in order of abundance, are encrusting

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Table 1 Data matrix includes list of thin sections and quantitative data represent the distribution of the recorded taxa and their individuals in the three intervals of the studied succession.

(continued on next page)

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Table 1 (continued)

Table 2 Numbers of individuals and percentage of abundance of each encrusting foraminifera taxon in the three intervals.

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A

B

C Fig. 11. Tayamaia marianensis (Hanzawa). A. Transverse section in a typical helmet-shaped test showing the three zones of layers: ventral zone with chambers filling hollow on ventral side, median zone with proloculus (arrow) and dorsal zone with quadrangular chambers on dorsal side (Gh6-10D), C.N.; B. Transverse sections in umbrella-shaped test stretched on substratum (Gh8-12C), C.N.; C. Vertical section in a test encrusting bivalvia (Gh3-6A), PPL.

foraminifera T. marianensis, P. escornebovensis; echinoids; Halimeda plates; bryozoans; bivalvia; coralline algae Lithophyllum and sponge spicules. Gypsina mastelensis is first described by Bursch [18] in the Tertiary of Moluccas; then, in Micronesia (W. Pacific; [49]); in the Middle Miocene of Australia [23,24]; recently, from the reefal limestone of Mururoa Atoll (French Polynesia; [84]). Gypsina plana [19]; Fig. 17A–C: it is considered a vital segment of the present reef structure. G. plana is much frequent, distributed

at many levels in both interval II and interval III, while totally absent in interval I. In interval II, it occurs in both the lower (bed no. 3) and the upper (beds no. 7 and 8) levels and in the reefs attached to coast at the base of interval III. In the lower level of interval II, it is abundant in bioherms; mainly associated to encrusting foraminifera T. marianensis, B. septentrionalis, S. globulus, S. cylindricum and P. escornebovensis; with M. dehartii, Miogypsinoides sp., M. complanatus, M. mexicana; coralline red algae as Lithothamnium spp., Mesophyllum spp., M.

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A B

C

D

Fig. 12. Tayamaia marianensis (Hanzawa). A. Vertical section showing the median zone with spiral, annular chambers and proloculus (arrow) occupies the apex (Gh8-13), C.N.; B. Horizontal section showing different types of chambers in median zone (Gh3-6B), PPL; C. Close up of B showing proloculus (white arrow) at center and spiral chambers around (black arrows), PPL; D. Close up of B showing annular chambers (arrows), PPL.

sancti-dionysiien, M. pacificum, A. myriosporum, and A. taiwanensis; green algae Chara sp., Microcodium sp. and Halimeda sp.; cheilostome and cyclostome bryozoans; bivalvia; ostracoda and planktonic foraminifera G. trilobus, G. altiaperturus and G. dehiscens. In this level of interval II, G. plana tests form nodules and accreted clusters over their surfaces. In the upper level of interval II, and base of interval III, expanded and stretched tests of G. plana are common. The percentage of abundance of its individuals reached a maximum in interval III (16.4%). It occurred in grey, dolomitic, crystallized, peloidal sandy limestone and found associated to T. marianensis, P. escornebovensis, N. saipanensis, D. vesicularis, S. cylindricum, G. saipanensis, G. mastelensis, S. globules, M. intermedia, M. gunteri,

Miolepidocyclina burdigalensis, Lithophyllum spp., Halimeda sp., bryozoans, echinoid and bivalvia. G. plana usually occurred in subtropical and tropical regions of the Indo-Pacific and Caribbean areas [88] and in the Holocene of Mauritius, Amirante and Providence Islands (Indian Ocean; [68]). Hanzawa [49] recorded plentiful Acervulina inhaerens plana Carter in the Miocene of Micronesia (W. Pacific). The wide Recent occurrence is in the eastern Caribbean [91]; in the Atlantic Ocean, Bahamas [112]. Gypsina saipanensis [49]; Fig. 18A–C: it is restricted at the upper level of interval II (bed no. 8) (Table 1); few inhabit fringing reefs at the base of interval III. Thin-bedded, highly crystallized, dolomitic and gypseous shelf limestone constitutes the enclosing

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Fig. 13. Tayamaia marianensis (Hanzawa). A. Horizontal section showing irregular chambers of ventral zone (Gh8-13), C.N.; B. Tangential section (Gh8-13), C.N.; C., Transverse section showing arcuate chambers of ventral zone (Gh3-6A), PPL; D. Vertical section showing subquadrangular chambers of ventral zone developed to fill hollow on ventral side (Gh8-12C), C.N.; E. Horizontal section showing more or less irregular chambers of dorsal zone (Gh3-6B), PPL.

sediment of G. saipanensis. The associated fauna comprising D. vesicularis, N. saipanensis, P. escornebovensis, T. marianensis, G. plana, S. globulus, M. intermedia, M. gunteri, Miolepidocyclina burdigalensis, cheilostome and cyclostome bryozoans, lithophylloid algae, green algae, echinoids, and bivalvia are common. The present species is described by Hanzawa [49] from the type locality in Ogso Talofofo and east of Sadog Doga (Saipan Island, W.

Pacific) and recently, from the reefal limestone of Mururoa Atoll (French Polynesia; [84]). Ladoronia vermicularis [49]; Fig. 19A and B: it is restricted only to interval I, representing the only encrusting foraminifer found with P. escornebovensis in this interval. The percentage of abundance of individuals of P. escornebovensis is 57.1%, exceeds that of L. vermicularis (42.9%) in interval I. In the present study,

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Fig. 14. Borodinia septentrionalis Hanzawa. A. Inverted ’V’ shaped transverse to subhorizontal section (Gh3-6), PPL; B. Spatuliform chambers in the median angular part (Gh3-6), C.N.; C. Close up of B, C.N.

the planorbulinid individuals are more abundant than those of the other detected families in the three intervals (Fig. 7). The sediment enclosing L. vermicularis is determined at the top part of interval I (bed no. 1, sample no. 4), directly below the conglomerates. It consists of gritty fossiliferous, yellow, massive limestone with the much clastic influx, of coarse to fine, angular and subangular sand, increased greatly upwards. Intermittent hard, gray, yellow, pelletiferous bands (2–20 cm) are distinguished in-between. Associated fossils are mainly abundant Amphistegina spp., especially A. bohdanowiczi, many barnacles, and bivalvia, together with dasycladacean C. saipania and Lithophyllum Geographic distribution of L. vermicularis reported the first description from the Tertiary coral reefs of Saipan (W. Pacific); it is

Fig. 15. Discogypsina vesicularis Silvestri. A. Subvertical section (not axial) showing rounded thick periphery (Gh8-13A), PPL; B. Median section passing with the equatorial disc, encrusting barnacles (Gh8-13A), C.N.; C. Close up of B showing large rounded proloculus (arrow), surrounded by typically arched chambers, PPL.

also recorded in the Plio-Quaternary reefal limestone of Mururoa Atoll (French Polynesia; [84]). Sphaerogypsina globulus [96]; Fig. 20A–C: the present specimens of S. globulus encrusted either coralline algae in the lower level of interval II (bed no. 3) or reef rubble in the upper level of interval II (bed no. 8). It also occurs in the reef fringing in the base of interval III. It is associated with T. marianensis and its accompanying fauna in the two levels (bed no. 3 at the base of interval II), where

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Fig. 16. Gypsina mastelensis Bursch. A. Vertical axial section showing flat base and polygonal chambers with highly arched thick roofs and walls (Gh6-10B), C.N.; B. Close up of A showing three juvenile chambers at the center of the flat basal layer (arrows), C.N.

red algae Lithothamnium spp. and Mesophyllum spp. are common and bed no. 8 at top of interval II where Lithophyllum spp. and cheilostome, cyclostome bryozoans are common reefal organisms. In the Tropical Pacific of the ’Albatross’, Todd [110] recorded S. globulus having fairly ovoid shapes of unevenly depressed spheres similar to those detected in Sinai. S. globulus is abundant in the Early Miocene (Aquitanian) Tagpochau Limestone of Micronesia, recorded in Hirippo, Rota, Mariana Islands (W. Pacific; [49]); in the Middle Miocene of Czechoslovakia [89] and of Romania [90]. Other records are in mid-Tertiary of Australia and New Zealand [23–25]; in the Neogene of Queensland Plateau (Australia; [12]); in Plio-Quaternary reefal limestone of Mururoa Atoll (French Polynesia; [84]). Furthermore, in the Middle Miocene of Tongan Platform (Indo-Australian Plate; [26]); in the Upper Eocene of the northern Italy [14]; in the Middle Eocene of Istrian peninsula of the Adriatic Carbonate Platform [33]. It is also distributed in the Late Eocene (Priabonian) to the Holocene of Philippine Archipelago [75]; in the Atlantic Ocean (Bahamas; [112]) and in the Lower-Middle Miocene rocks of southeast Zagros basin (south of Iran; [54]). 4.2.3. The homotrematid group In the present study, taxa belonged to homotrematidae are the least encrusting foraminifera. They represent a percentage of 9.1% of the total collection, and the relative abundance of its individuals in the total succession is 7.3%. Individuals of the homotrematidae correspond to one species Sporadotrema cylindricum. Sporadotrema cylindricum [21]; figures 21A–D: it is concentrated mainly in interval II, occupying entirely two levels (bed no. 3 and bed no. 7). The enclosing sediment in bed no. 3 is crystalline, massive, embedded limestone forming small bioherms at base of interval II. In bed no. 7, S. cylindricum occurred in beach sandstone rock, highly calcareous, coarse-grained, well cemented and stratified. S. cylindricum dominates modern reefs, cosmopolitan [69]. It is recorded in the Holocene of the Indian Ocean and Maldives Islands; in the Timor Sea (north Australia); also in the Philippine Archipelago from the Late Oligocene (Chattian) to the Middle Miocene (Serravalian) by Matsumaru [75]. 5. Discussion 5.1. Paleoecology Data resulted from the distribution of determined encrusting foraminifera taxa revealed restriction to a certain level and/or only a few levels inside each interval and complete absence in others.

In general, the light intensity is the primary factor limiting distribution of the light dependent encrusting algae and the symbiotic foraminifera [56,57,111]. In this study, the extended species to Recent are easy to interpret environmentally by direct correlation with the habitat of their living forms. On the other hand, the reported species as only fossils are difficult to apply a strict uniformitarian approach. The present encrusting foraminifera were interpreted environmentally into three eczones according to (i) the change in taxonomic composition of the detected taxa, (ii) their distribution throughout the studied succession, (iii) enclosing sediments and facies characters, (iv) associated fauna, (v) encrusted substrates and (vi) morphologic variations. Ecozone 1: It occupies the uppermost level of interval I (bed no. 1, sample no. 4). Planolinderina escornebovensis Freudenthal and Ladoronia vermicularis (Hanzawa) are the only encrusting foraminifera inhabited this level. Both P. escornebovensis and L. vermicularis are characterized by a relatively small size. According to Hottinger [58], the smaller size has, in general, a wider ecological range and geographical distribution than the larger. Moreover, Hansen and Buchardt [47] and Haunold et al. [53] stated that the abundance of Amphistegina is greatest between 50 m and 70 m depth in the Red Sea. Ecologically, the occurrence of Amphistegina spp. is greatly influenced by substratum [47]. In the Bay of Safaga, Amphistegina in macroid facies occurred commonly on rocky surfaces with thin sand veneers [52]. Kroeger et al. [65] recorded abundant Amphistegina lessonii in facies with 10–50% siliciclastic sand. This occurrence is found in agreement with the lithologic characters of the level occupied by ecozone 1, which consists of gritty fossiliferous limestone with abundant Amphistegina bohdanowiczi and much clastics. According to Reiss and Hottinger [94], Bassi et al. [9] considered Amphistegina bohdanowiczi as A. lessonii are inhabitant for the upper photic zone proving a negative phototropic movement response in escaping photoinhibition at mid-day. In addition, Amphistegina occurs in regions with minimum winter sea-surface temperature of 14 °C. [65]. Lemoine [67] has shown the abundant presence of dasycladacean Cymopolia reveals shallow water and commonly a muddy or silty bottom down to 10 m depth and are rarely found at more than 25 m depth. The reef-flat facies [62] are greatly found in accordance with the presence of large skeletal debris of bivalvia and barnacles in facies of ecozone 1. As common biocomponents in clean skeletal lime rudstone facies, reef-building metazoans masses typify the present facies in ecozone 1. In addition, the gritty substrate and the intermittent pellet-rich bands

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Fig. 17. Gypsina plana (Carter). A. Vertical section perpendicular to surface (Gh36B), C.N.; B. Vertical section showing apertural pores (arrows) (Gh9-14), C.N.; C. Close up of A showing plane, rectangular, brick-like chambers with few apertures (arrow), PPL.

provide evidence for fluctuations in waves that sweep sands and form barriers creating small protected environments. It was observed, such results are in agreement with the reef-flat deposits in James [62]. Consequently, the bio and litho-components and facies pattern in ecozone 1 can indicate reef-flat conditions in the upper photic zone characterized by moderate wave energy of only a few meters depth at most. The reduced number of species (relative abundance is 4%) of encrusting foraminifera in this

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ecozone signifies that both P. escornebovensis and L. vermicularis have no opportunity to flourish. The preferred occurrence of P. escornebovensis and L. vermicularis with Amphistegina bohdanowiczi and Cymopolia on gritty, rough bottom indicates reef-flat conditions in the upper photic zone that don’t exceed 25 m depth. Ecozone 2: It occupies the lowermost level of interval II (bed no. 3), directly overlying conglomerate. It forms a reefal construction in the form of small bioherms, without framework; made up of yellowish gray, frequently sandy, crystalline, fractured, massive, embedded limestone in which encrusting foraminifera and coralline algae are the predominant elements. The resulted rock describes foraminiferal red algal bindstone facies; its micro-biocomponents are tightly packed with sparite cement in few lime mud matrix. Under conditions typifying ecozone 2, encrusting foraminifera individuals reach a considerable high percentage of abundance consisting of T. marianensis, G. plana, B. septentrionalis, S. globulus, P. escornebovensis and S. cylindricum association. This association is interpreted environmentally with the aid of published data on modern and fossilized encrusting foraminifera. Abundant coralline red and green algae are accompanying for encrusting foraminifera in ecozone 2 as Lithothamnium spp., Mesophyllum spp., M. sanctidionysiien Chara sp., Microcodium sp. and Halimeda sp. Similarly, some planktonic foraminifera species such as Globigerinoides trilobus, and Globoquadrina dehiscens are important associates. In this study, such associations are available as depth indicators. The photosynthetic red algae group use the blue light penetrating the deep waters. So, they exist up to 125 m depth [102]. The melobesioids comprising Lithothamnium and Mesophyllum are the deepest red algal taxa in Recent and Neogene environments, existing down to 110–120 m water depth [6,17]. This is in accordance with the dominance of melobesioid taxa in the basal part of interval II (ecozone 2), where overall deepening is indicated by the presence of abundant planktonic foraminifera. Regarding planktonic foraminifera, Bernoulli et al. [11] recorded the planktonics Globigerinoides trilobus, Globigerinoides sp. and Globoquadrina dehiscens from the Miocene shallow-water limestones of São Nicolau (Cabo Verde). On the other hand, in Great Britain and Norway, Lithothamnion are found mostly in water depth shallower than 20 m [55,60] depending on shading to survive in shallow water [38]. Mesophyllum sancti-dionysii from Crete suggested a preference of shaded settings [65]. Generally, algae forming thin crusts or sheet-like masses live from tide level down to more than 100 m [67]. In addition, Mesophyllum rarely occurs in cold environments [60]. Kroeger et al. [65] concluded that Lithothamnion and Mesophyllum associations from Crete are restricted to the lower photic zone and shaded environments. Lund et al. [70] suggested that recent Lithothamnion and Mesophyllum from eastern Australia are most abundant at water depths below 40 m. Regarding the above-mentioned discussion, the general absence of Amphistegina bohdanowiczi in ecozone 2 led to suggest the water depth is more than 25 m. The general scarcity of significant coral masses is observed at this level. In the Red Sea modern reefs, Porites is reported from water depths of no more than 25 m [98]. Therefore, the Porites fragment detected in only one thin section is a reworked fragment from shallower depths. On considering the calcified green algae, they commonly form biohermal thickets or mounds mainly in warm-temperate to tropical areas [102]. Halimeda thrives best in relatively shallow waters, although some species grow at depths of over 120 m [63]. The deepest zone of growth of green calcareous algae on modern reefs is about 70 m [62]. Furthermore, comparing with Recent occurrences of the encrusting foraminifera; in the northern Red Sea; Rasser and Piller [93] reported acervulinid foraminifera are more dominant in deep water (below 45 m) where light conditions reduce the competi-

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A

B

C

Fig. 18. Gypsina saipanensis Hanzawa. A. Sub-vertical section (Gh8-13A), C.N.; B. Transverse section (axial) showing embryonic spherical proloculus at the center (arrow) with equatorial chambers around (moderate preservation) and intertwined chambers near periphery (Gh8-13A), PPL; C. Horizontal to slightly oblique section (Gh8-13A), PPL.

tion of coralline algae, while shallow occurrences were recorded around 25 m. Sphaerogypsina globulus is black in color and always spherical in shape, recorded in considerable abundance from 15–88 m in the Atlantic Ocean, Lee Stocking Island, Bahamas [112]; in this occurrence, its mean abundance per cm2 ranges from 1.2–1.4 in depth ranging from 30–88 m, while totally absent after 88 m. Gypsina plana is known to occur on exposed substrates in fore-reef environments [85]. In the eastern Caribbean and in association with coralline algae, G. plana may form large (2–15 cm) nodules, usually between 30–60 m water depths [91]. In Atlantic Ocean (Bahamas), the mean abundance per cm2 of Gypsina plana ranges from 0.6–0.8 in depths 30–73 m, while totally absent after 73 m [112]. This is in accordance with results of high abundant S. globulus and G. plana in ecozone 2. Test morphology is greatly controlled by water energy; the globose and hemiglobose morphologic types dominate low energy and more protected habitats, while the stretched, flat-encrusting

tests characterize exposed substrates under high energy conditions [72,94]. Accordingly, in ecozone 2 the concentration of forms consisting of: (i) helmet-shaped tests of T. marianensis with complete absence of the stretched tests, (ii) G. plana tests growing out into projections forming knobs and nodules over their surfaces and (iii) hemispherical to spherical tests of S. globulus well indicate low-energy conditions at the level occupied by ecozone 2. Bosellini and Papazzoni [14] interpreted the globose morphotypes preferred to encrust coral upper surfaces in order to reduce lateral spatial competition with algae as competition is generally high on coral upper surfaces. This is in agreement with the abundance of the globose morphotypes of the encrusting foraminifera T. marianensis, S. globulus and G. plana in association with red algae in ecozone 2. As they constitute vertical expand higher than algal crusts to reduce lateral competition of place, these high accreted forms can play the role of providing shading to the symbiont algae. This confirms that the association inhabits ecozone 2 is living in

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A

151

B

Fig. 19. Ladoronia vermicularis (Hanzawa). A. Horizontal section (Gh1-4A), C.N.; B. Close up of A showing raspberry-like chambers (embryonic stage), acervulinid chambers (second stage) and vermicular chambers (adult stage) (arrows from right to left), PPL.

the photic zone intervals. These results significantly assert shell morphology and shading by other organisms are key factors that influence water depth and water energy. It can be concluded that encrusting foraminifera species that populate ecozone 2 indicate a shallow water depth in the fore-reef environment, of more than 25 m and doesn’t exceed 70 m. This is accompanied with preferred occurrence in the lower photic zone where low energy and protected shaded settings are available. Ecozone 3: It occupies the upper part of interval II (beds no. 6, 7, 8) and the lower part of interval III (the fringing reefs at the base of bed no. 9). It represents the most enriched zone in encrusting foraminifera species. This ecozone shows the maximum diversity of species detected in Sina, it includes more than 80% of the determined species. The biocomponents of encrusting foraminifera in ecozone 3 include T. marianensis, P. escornebovensis, N. saipanensis, D. vesicularis, G. mastelensis, G. plana, G. saipanensis, S. globulus and S. cylindricum in association with the environmentally indicator Halimeda and only Lithophyllum is the most prominent red algae in association to encrusting foraminifera in this ecozone. In rocks containing abundant Halimeda, coralline algae tend to be rare [63]. Lithology in ecozone 3 shows change from pellet-rich carbonates to coarsely crystallized, dolomitic, gypseous banded limestone and a calcareous sandstone transitional bank in-between resulted in pelmicrite, sublitharenite and floatstone facies. The main encrusted substrates are algae, reef rubble, and barnacles. The sandstone bank depositional characters describe beach rock (sandy coral beach) which is known from intertidal zones in tropical or subtropical regions [10]; it represents a transition to reefs attached to coast. From about tide level down to depths of 15–20 m. Regarding associated fauna to encrusting foraminifera in ecozone 3, Halimeda plates are most common in near-backreef areas; they grow mainly in warm temperate to tropical areas with nearnormal salinity water [63,102]. Lithophyllum is widely documented shallow depths in environments of both modern and fossil settings [27,28]; down to 20 m water depth [15]. It also showed maximum abundance in the upper photic zone shallower than nearly 40 m, with annual minimum seawater surface temperature well below 18 °C [65]. Generally, lithophylloids are common in modern tem-

perate shallow-water environments in the Mediterranean [5] and dominate the subtropical and warm-temperate areas [28]. Other important associates to the encrusting foraminifera assemblage in ecozone 3 are the cheilostomes and cyclostomes types of bryozoans; they known to typify the bryozoan-rich sediments in temperate-water conditions [102]. Data about encrusting foraminifera from modern reefs documented the expanded environmentally controlled morphology of Gypsina plana encrusting coral rubble in patch reefs and lagoons [85] and spread over the substratum for 3–4 inches diameter or more [[68]; in Indian Ocean] or make much expansive crusts (up to numerous centimeters in width) [85]. In Bahamas Islands (Atlantic Ocean) the spreading capacities of G. plana are restricted and its encrusting pattern on shell surfaces is the same on both exposed and unexposed surfaces, it documented the greatest value of mean abundance per cm2 (2.75) at 15 m depth [112]. This is in accordance with the concentration of stretched T. marianensis in the level occupied by ecozone 3 (upper levels of interval II) as a vital reefal component and the abundant spreading patches of G. plana that reached maximum percentage of abundance of its individuals (16.4%) in interval III (the upper part of ecozone 3) (Fig. 8). So that, both T. marianensis and G. plana as eco-morph controlled types can indicate high water energy levels in ecozone 3. It is also added, Sporadotrema cylindricum dominates modern reefs and inhabits warmer waters [69]; it reveals a high percentage of abundance of its individuals (8.7%) in the upper level of interval II corresponding to lower part of ecozone 3. Compiling data resulted from analysis of biocomponents, morphologic types, encrusted substrates, lithlogic characters and facies analysis, and unifying conditions surrounding the co-occurring insitu growing organisms provide many evidences either of fauna or facies that indicate the prevalence of shallow water conditions and suggest the interval occupied by ecozone 3 is linked to gradation to the most shoreward reef reaches. It therefore, corresponds to nearshore tidal reefs in the agitated upper photic zone, extending down to depths of 15–20 m and prevailed back-reef warm lagoonal settings indicative to subtropical and warm-temperate to tropical regions.

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A

B

C

Fig. 20. Sphaerogypsina globulus (Reuss). A. Median section (Gh3-6B), PPL; B. Horizontal to tangential section near the outer surface showing thick walls between chambers (Gh8-13B), PPL; C. Centered, median section showing megalospheric juvenile (Gh3-6B), PPL.

5.2. Paleobiogeography Wadi Gharandal represents one of the most complete Oligocene-Miocene marine successions in Egypt. Notably, its sediments contain considerably abundant planktonic and benthic foraminifera of high abundance occupying many levels of thick clastic and, frequently, carbonate succession [1]. On the horizontal scale, the Miocene sedimentation in Sinai displayed diversified synchronous depositional realms and facies reflecting different submergence and irregular topography of the basin [105,108,113]. On the vertical scale, the Oligocene and Miocene intervals influenced the paleogeography and the paleobiogeography of Mediterranean region [4,99,100,61,97,109]. Here, the geographic distributions of the defined encrusting foraminifera species (Table 3) show that the W. Pacific region have the greatest concentration of this reefal Miocene fauna (Fig. 22) comprising Japan, Daitô Island, Mariana Islands, Hirippo, Rota,

Saipan Islands, Ogso Talofofo, Sadog Doga, Tinian, Laderan Dago, Mulato, Sadog Talofofo and kanat Fahang Lichan of Micronesia. According to Table 3, it is observed that the rough distributions of encrusting foraminifera are recorded gradationally to India-south Asia, South Pacific, Australia, Mediterranean and Europe, Caribbean Sea and the Atlantic Ocean. Accordingly, the Miocene encrusting foraminifera of Wadi Gharandal show a close affinity to those of equivalent age in the W. Pacific region located on near paleolatidudinal position than those in other places of low distributions. Moreover, this affiliation of encrusting foraminifera is correlated by many species belonging to other groups of organisms (e.g. larger foraminifera and red/green algae). Consequently, similarity and real matching of faunal assemblage are proved in both W. Pacific and Sinai. It is observed that the larger foraminifer’s collection encompasses the genotypes, subgenotypes and the type species of encrusting foraminifera from Sinai. Regarding Cole and Bridge

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A

C

153

B

D

Fig. 21. Sporadotrema cylindricum (Carter). A. Longitudinal section in the end part of a branch (Gh7-11A), PPL; B. Longitudinal section in a meeting point of two branches (7–11B), PPL; C. Longitudinal section in meeting point of two branches (Gh3-6A), PPL; D. ?Longitudinal section showing chambers and their communicating fine passages (Gh3-6B), PPL.

[32] and Cole [30], the collection of larger foraminifera from Saipan (W. Pacific) is related to high Tertiary e (Aquitanian) to possible low Tertiary ƒ (Burdigalian) age (Tagpochau limestone). In comparison with the present study, larger foraminifera from Saipan show perfect synonyms of the studied encrusting foraminifera species in Sinai; especially T. marianensis, together with important Miogypsinoides and Miogypsina. Then, many genera and species of encrusting foraminifera from Wadi Gharandal show the same stratigraphic ranges in the W. Pacific. Moreover, this faunal affinity is easily recognized for both red and green algal collections, e.g. Lithothamnium, A. taiwanensis, A. myriosporum, M. pacificum, M. sancti-dionysiien, Lithophyllum spp., J. vetus, C. saipania, Halimeda sp. and Microcodium sp. Such algal collection shows a good matching with those observed in the Early Miocene Tagpochau Limestone of Saipan and Mariana Islands [63], which enhances the present affiliation. Paleoenvironmentally, the encrusting foraminifera from W. Pacific islands of Micronesia [49] from the Aquitanian-Burdigalian (Tagpochau Limestone) inhabited biohermal, lagoonal and fore-reef detrital facies of coral reefs. On consequence, the assemblage of [49] correlates the constitutions of both ecozone 2 and ecozone 3, corresponding to Interval II and base of interval III in Sinai.

This great coincidence in assemblage characters of similar abundance, diversity, and paleoecological conditions proved in both W. Pacific and Sinai show an exact and entirely Miocene W. Pacific paleobiogeographic affiliation. As the studied region in Sinai is generally considered part of the temperate zone, a pronounced incorporation occurred between the high latitude warm temperate (subtropical) sea and the nearest lower one of tropical settings. These climatic conditions are highly confirmed by the presence of faunal associations comprising lithophylloids, green algae Chara sp., Microcodium sp. and Halimeda sp., G. plana, S. globulus, cheilostome and cyclostome bryozoan-rich rocks and the specific lithology of the beachrock. Regarding paleogeography, Rögl [[100], p. 339; Fig. 2 timeslice] dubiously proposed a marine connection during the Early Oligocene extended north of India to the W. Pacific across Asia. Further, this connection is reduced in Oligocene and Early Miocene (Aquitanian). More diminution and complete closure in such proposed connection are occurred, respectively, during Early and Late Burdigalian [[100], Figs. 3–6 time-slices]. On the contrary with Rögl [100], the rigorous similarity of the Miocene encrusting assemblage in Sinai with W. Pacific region is certainly required quite direct and short distance connection.

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Table 3 Geographic distribution of the recorded encrusting foraminifera species.

Accordingly, the long distance Indo-Pacific influences are disqualified. Furthermore, Early Oligocene water elongation across Asia to W. Pacific [100] is proposed to be the main connection that responsible for the observed profuse faunal exchange and copious water inroad constituting such paleobiogeographic affiliation. On consequence, this connecting sea is thought to still open even during Aquitanian (Fig. 23). The restriction of the Iranian basins in the Early Burdigalian [97] which considered the entry of the passage to the W. Pacific led to the ceasing of the connection with W. Pacific across the proposed west-east connecting sea. The interruption of both Iranian basins and the present studied basin in Sinai during the Early Burdigalian is, therefore, contemporaneous. In interval III of the studied succession, evaporites (lagoonal gypsum and anhydrite) and fringing reefs were deposited reflecting marginal basins and document restricted marine conditions

accompanied by emersions. Notably, the evaporitic formation in interval III is synchronous with the Early Burdigalian analogous restriction of the central basin of Iran (Qom basin). All are actually relating to the ongoing plate collision of Africa and Arabia with Eurasia which ultimately resulted in the closure of the ’ Iranian gate’ between the studied basin in Sinai and the proposed westeast connecting sea to the W. Pacific, and at the same time, the restriction of the Tethyan Sea and the closure of its proficient ’Iranian gateway’ to the Proto–Indopacific [97]. This interruption during the Early Burdigalian is confirmed by the generation of a biogeographic land barrier for marine biota coeval the Early Burdigalian time. In the same region in Sinai, studies by Abd-Elazez [1] have proved a change in the course of basin connection after the Early Burdigalian isolation. It is indicated by the dominance of subsequent Mediterranean and Paratethyan faunal affinities characteristic for younger exposures of Langhian age.

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6. Conclusion

Fig. 22. Stacked Venn Smart Art Graphic shows the degree of affiliation to the different geographical locations gradually increases from the Atlantic Ocean to the W. Pacific (the darker tone represents the place of most affiliation).

In a concordant association of symbiotic encrusting foraminifera, coralline algae and bryozoans, the encrusting foraminifera proved to be the dominant organisms among fossil reef assemblage in Wadi Gharandal, Sinai, Egypt. The encrusting foraminifera represented the most vital segment of a reefal system, without a framework, consisted mainly of embedded and massive limestone development. The recognized encrusting foraminifera assemblage was much abundant but not diverse. The special multitude was that of the planorbulinid species Tayamaia marianensis. Encrusting foraminifera associations were restricted only to certain level and/ or few levels inside each interval of the succession revealing (i) moderate variations in taxonomic composition and (ii) high variations in relative abundance of individuals within intervals. Such association succeeded to attribute the succession in Sinai to three different ecozones suggesting a wide range of paleoecological variations. Accordingly, the reefal system in Sinai represents three ecozones. In ecozone 1, the reef-flat conditions were identified reflecting upper photic zone conditions, not exceed 25 m depth. On the other hand, in the lower photic zone at 25–70 m depth represents fore-reef under low energy conditions (ecozone 2). Under high energy and near shore back-reef warm lagoonal conditions, the ecozone 3 was identified. At the same paleolatitudinal positions, the Late Oligocene to Early Miocene (Early Burdigalian) encrusting foraminifera in Sinai display similar diversity and confirm obvious faunal resemblance with those reefs of the W. Pacific islands. These observations between Sinai and W. Pacific encrusting foraminifera indicates that circulation across apparent connection extending north of India passing through east Iran and expanded directly across Asia to West Pacific occurred. The prior marine connection in the Early

Fig. 23. During the Oligocene-Early Miocene (Aquitanian), a marine connection, north of India, extended west-east direction from eastern Mediterranean passing through the east of Iran, then expanded directly eastward direction across Asia to W. Pacific permitting copious faunal exchange and warm temperate to tropical water inroad (modified after [100]).

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Oligocene is proposed to be the best direct and shortest water connection to the W. Pacific interpreting this strong, rigorous faunal similarity. Accordingly, it is thought that the water connection was still open during Aquitanian and it was ceased in the Early Burdigalian. As the Iranian central basins have provided evidence to be the proficient gateway of the Tethyan Sea to the Proto-Indopacific and at the same time, it is the gate to the W. Pacific across the proposed connection; then, the restriction of the Iranian basins during Early Burdigalian would cease connection also with W. Pacific. Similar situation in Sinai has asserted the closure time and confirmed the generation of the biogeographic land barrier. Regarding the above-mentioned results, pronounced integration of warm temperate and tropical water above and below the tropic of Cancer, respectively, was proved by typical indicators of fauna associated to encrusting foraminifera. Such subtropical, warm temperate and tropical fauna were found endemic in Sinai reflecting the predicted climatic criteria.

Superfamily ACERVULINACEA [103] Family ACERVULINIDAE [103] Genus BORODINIA [48] Borodinia septentrionalis [48] Genus DISCOGYPSINA [106] Discogypsina vesicularis [106] Genus GYPSINA [20] Gypsina mastelensis [18] Gypsina plana [19] Gypsina saipanensis [49] Genus LADORONIA [49] Ladoronia vermicularis [49] Genus SPHAEROGYPSINA [40] Sphaerogypsina globulus [96] Family HOMOTREMATIDAE [34] Genus SPORADOTREMA Hickson, 1911 Sporadotrema cylindricum [21]

Acknowledgements

Supplementary materials

I would like to thank Mohammad Abd-Elazez for graciously providing access to the rock material and generously allowing me to prepare the thin sections required for getting data of this publication and I am indebted to Ibrahim Ied and Ahmed Abd El-Shaheed Henaish for helping in final directing of figures.

Supplementary material associated with this article can be found, in the online version, at doi:10.1016/j.grj.2017.03.001.

Appendix Literature that rendered the description of the encrusting foraminifera either recent or fossil forms are generally few; most taxa have a confusing taxonomic history and thus actually difficult to recognize. The morphology and the chambers arrangement of most encrusting species vary in different stages of growth; this indeed, makes an accurate specific determination much critical especially for specimens available only in non-oriented thin sections. The acervulinids may also vary in form according to the configuration of the substratum that they encrust and the conditions of the surrounding environment [82,83]. In the present microscopic diagnosis, the shapes, interrelations, and arrangements of chambers in the different growth stages of the individuals are the differential features that the definition of taxa is based on; and in most instances, it is wrong to rely mainly on the external outline for specimens examined in random thin sections or for those which stretch and cover vast area of the substratum during the development of their growth. Many fossil genera of larger foraminifera including some of the encrusting forms have been abolished in modern taxonomic divisions and in the new databases of foraminifera on the websites; in addition, some generic names are used in systematic classifications other than foraminifera, e.g. the famous genus Borodinia of the acervulinids is newly accepted as a genus in the family Brassicaceae, in the major group Angiosperms (flowering plants). Taxonomic lists of encrusting foraminifera in Wadi Gharandal (Sinai). Order FORAMINIFERIDA Eichwald, 1830 Suborder ROTALIINA [35] Superfamily PLANORBULINACEA [104] Family PLANORBULINIDAE [104] Subfamily PLANORBULININAE [104] Genus NEOPLANORBULINELLA [74] Neoplanorbulinella saipanensis [74] Genus PLANOLINDERINA [39] Planolinderina escornebovensis [39] Genus TAYAMAIA [50] Tayamaia marianensis [49]

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