Carboniferous brachiopod communities from Wadi Araba, western side of the Gulf of Suez, Egypt: Paleontology, paleoecology and paleobiogeography

Journal Pre-proof Carboniferous brachiopod communities from Wadi Araba, western side of the Gulf of Suez, Egypt: Paleontology, paleoecology and paleobiogeography Shahin Abd-Elhameed, Abdel Aziz Mahmoud, Yahia El kazzaz, Yasser Salama PII:

S1464-343X(19)30399-1

DOI:

https://doi.org/10.1016/j.jafrearsci.2019.103744

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AES 103744

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Journal of African Earth Sciences

Received Date: 31 August 2019 Revised Date:

3 November 2019

Accepted Date: 15 December 2019

Please cite this article as: Abd-Elhameed, S., Mahmoud, A.A., El kazzaz, Y., Salama, Y., Carboniferous brachiopod communities from Wadi Araba, western side of the Gulf of Suez, Egypt: Paleontology, paleoecology and paleobiogeography, Journal of African Earth Sciences (2020), doi: https:// doi.org/10.1016/j.jafrearsci.2019.103744. This is a PDF file of an article that has undergone enhancements after acceptance, such as the addition of a cover page and metadata, and formatting for readability, but it is not yet the definitive version of record. This version will undergo additional copyediting, typesetting and review before it is published in its final form, but we are providing this version to give early visibility of the article. Please note that, during the production process, errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. © 2019 Published by Elsevier Ltd.

Carboniferous brachiopod communities from Wadi Araba, western side of the Gulf of Suez, Egypt: paleontology, paleoecology and paleobiogeography Shahin Abd-Elhameed1, Abdel Aziz Mahmoud1, Yahia El kazzaz1 and Yasser Salama*2 1

Geology Department, Faculty of Science Helwan University

2

Geology Department, Faculty of Science, Beni-Suef University

*

[email protected]

Abstract Rod El-Hamal area was considered as one of the most important outcrops in the western side of the Gulf of Suez where the Carboniferous marine sediments are exposed. 12 brachiopod species fauna identified from the late Carboniferous Rod El-Hamal Formation in the study area. The identified brachiopod fauna is dominated by diverse species of Athyridida, Orthotetida, and Productida. Composita is the dominant brachiopod genus that represented by six species. The bryozoans are also diverse group including Cystoporata, Fistuliporoida, Trepostomata and Fenestrata, which are known from the limestone beds in Rod El-Hamal Formation. Grainstone and Rudstone are the main microfacies in the bryozoa-bearing limestone beds. The facies associations of Rod ElHamal Formation show fluctuation between the carbonate and siliciclastic inner shelf. The articulated shells of the Brachiopoda display damage resulting from predation. The paleobiogeography of the benthic assemblages including brachiopods and bryozoans was highlighted. Keywords: Carboniferous, brachiopod communities, paleontology, Gulf of Suez, Egypt 1. Introduction The late Paleozoic outcrops on the western side of the Gulf of Suez in Egypt have been the subject of sporadic literatures (Schweinfurth, 1883; Walther, 1890; Schellwien 1894; Said, 1962; Darwish, 1992; Abdel-Azeam, 1997; Abdel-Shafy et al., 2000). Abdallah and El-Adindani (1963) subdivided the late Carboniferous succession at Wadi Araba area into Abu Darag, Aheimer, and Rod El-Hamal formations. Klitzsch (1990) later confirmed the late Carboniferous age for Rod El-Hamal Formation. The main widely-distributed benthic communities of the Rod El-Hamal Formation are the brachiopods and bryozoans. Brachiopods are one of the most thrived benthic groups during the Paleozoic time (Shen, 2018) that live in fully oxygenated, normal saline waters (Doyle, 1996), inhabiting a variable water depths (Harper et al., 2008; Cusack et al., 2008). Throughout the Paleozoic, the bryozoan communities were globally distributed (Tuckey, 1990a, 1990b; Gilmour and Morozova, 1999; McCoy and Anstey, 2010). They inhabited different 1

marine environments, and were the significant components of the Paleozoic reefs (Cuffey, 2006; Smith et al., 2006). The main objectives of this work are to discuss the stratigraphy, paleontology and paleoenvironments of the Rod El-Hamal Formation and to highlight the paleoecology and paleobiogeography of the recorded benthic communities. 2. Geological Setting During the Paleozoic, Egypt was located between a large exposed continental landmass in the south and the Paleo-Tethys in the north (Guiraud et al., 2005). The Early Carboniferous witnessed a global rise in sea-level and warm temperatures in North Africa (Guiraud et al., 2005). A shallow-marine siliciclastic platform developed in Egypt, passing northwards into mixed and carbonate platforms (Fig. 1a). During the Bashkirian period, the sea-level fell and the marine conditions were restricted to northeastern Egypt. The sea-level rose again during the Moscovian period and a mixed carbonate-siliciclastic platform fringed the North African margin (Fig. 1b).The investigated area includes two important plateaus, Northern and Southern Galala plateaus, separated by Wadi Araba. Wadi Araba is a topographically low-area of easily eroded Carboniferous rocks. It represents a NE-SW doubly-plunging anticline, with an axis dipping at opposite directions (Farouk, 2014). The studied Rod El-Hamal section is located at Latitude 29° 08` 49`` N and Longitude 32° 27` 27`` E. This area can be reached through El-KorimatZaafarana asphaltic road (Fig. 2). 3. Materials, methods and depository The field work was the backbone of this work. A composite section from Rod El-Hamal Formation was studied in detail and brachiopod specimens were collected for systematic and paleoecological studies. The carbonate rocks were sampled, investigated and photographed using Polarizing Microscope for microfacies studies. The microfacies types of the carbonate rocks are classified following the nomenclature of Dunham (1962) with modifications of Embry and Klovan (1972) and Scholle (2003). All the materials are deposited in the Museum of the Geology Department, Helwan University, Egypt. 4. Lithostratigraphy The exposed late Carboniferous Rod El-Hamal Formation is composed mainly of colored sandstones, mudstones and thin carbonate interbeds (Fig.3). It is subdivided into five units (Abdallah and Adindani, 1963). These units from base to top are as follow: Unit I: It is well represented at the junction between Wadi Rod El-Hamal and Wadi Araba. The basal part is not exposed and the measured section attains 70 m thick. This 2

unit consists mainly of cross-bedded sandstone alternating with multi-colored clays (Fig.3g), and topped by two fossiliferous limestone beds (Abdallah and Adindani, 1963). Unit II: 60 m-thick succession facing Wadi Rod El-Hamal is measured (Fig.3f). The lower part is mainly made up of grey sandy marls overlain by fossiliferous limestone beds, containing bryozoan fragments, crinoid ossicles and few brachiopod shells. The top of this unit is composed of brown sandstone intercalated with shale (Abdallah and Adindani, 1963). Unit III: This 70 m-thick unit is best exposed at the entrance of Wadi Bekheit (Fig.3d). The lower part is distinguished by the presence of intercalations of grey to greenish sandstone and gypsiferous shale. This is followed by fossiliferous sandy limestone beds and capped by brown dolomitic limestone. This unit yields brachiopods (Fig.3e), bivalves, gastropod moulds with few crinoid fragments (Abdallah and Adindani, 1963). Unit IV: To the east of the junction between Wadi Araba and Wadi Rod El-Hamal, this unit is measured about 45 m. It mainly consists of sandstones and shales at the lower part, while green marls intercalated with thin fossiliferous limestone beds at the upper part (Fig.3b). The limestone beds yield well-preserved brachiopods (Fig.3c), gastropods, crinoids, bivalves, bryozoans and rare trilobite fragments (Abdallah and Adindani, 1963). Unit V: It was measured about 112 m at the upstream of Wadi Rod El-Hamal (Fig.3a). This unit is made up of shales and sandstones with minor limestone interbeds. It is characterized by the presence of a coral association at its top, a Rhynconella assemblage at the middle part and thick shale beds at the base. The fossiliferous horizons yield some corals, brachiopods, bivalves and gastropods (Abdallah and Adindani, 1963). 5. Macrofauna from Rod El-Hamal Formation: The identified brachiopoda species from the Rod El-Hamal Formation belong to 12 monospecific genera. Athyridida brachiopods are the most abundant that represented by seven species (Fig.4). The Orthotetida has three species of the brachiopod population. The order Productida includes two species that constitute a small portion of individual abundance (4.5% of the total faunal abundance). On the generic level, Composita is the most dominant that represented by six species such as C. ovata, C. subquadrata, C. subtilita, C. ambigua, C. trilobata, and C. trinuclea (Fig.5). Phlyum: Brachiopoda Dumeril, 1806 Class: Articulata Huxley, 1869 Order: Athyridida Boucot Johnson, and Staton, 1964 3

Suborder: Athyrididina Boucot Johnson, and Staton, 1964 Superfamily: Athyridoidea Davidson, 1881 Family: Athyrididae Davidson, 1881 Genus: Composita Brown, 1849 Composita ambigua (Sowerby, 1823) (Fig.6-9) 1823-1834 Spirifer ambigus, Sowerby, p. 105. 1890 Spirigera ambigua (Sowerby), Walther, p. 430, pl. 24, Figs. 1, 2, 4, and 5. 1965 Composita ambigua (Sowerby), Moore, p. H662, Fig. 537 (2). Material and occurrence: 12 specimens [PRB-(3-14)], Unit (II) and (III). Measurements: Dimensions in mm are given in Table.1. Diagnosis: Composita is the most common genus in the collected fauna. The shells of C. ambigua are suboval outline, biconvex, with smooth surfaces, bent beak, and small foramen. Composita subtilita (Hall, 1852) (Fig.10-13) 1852 Terebratula subtilita, Hall, p. 409, pl. 4, Figs. 1 and 2. 1931 Composita subtilita (Hall), Morse, p. 314, pl. 50, Fig. 6. Material and occurrence: 16 specimens [PRB-(15-30)], Unit (II) and (III). Measurements: Dimensions in mm are given in Table.2. Diagnosis: The shells are of medium size, subovate, with subequally-convex valves, small beak, and greatest thickness at the midlength. C. ambigua (Sowerby) differes from C. subtilita (Hall) by having only one fold in the anterior side, while the latter has two folds. Composita subquadrata (Hall, 1858) (Fig.14-17) 4

1858 Athyris subquadrata, Hall, p. 703, 708, pl. 27, figs. 2 a-d. 1914 Composita subquadrata (Hall), Weller, p. 489, 490, pl. 81, figs. 1-15. 1964 Composita subquadrata (Hall), Grinnell and Andrews, p. 232. Material and occurrence: 19 specimens [PRB-(31-49)], Unit (II) and (III). Measurements: Dimensions in mm are given in Table.3. Diagnosis: The shell of species is subquadrate outline and medium size. The anterior commissural line shows rounded deflection due to a projected anterior margin extending into a fold. Composita ovata Mather, 1915 (Fig.18-21) 1915 Composita ovata, Mather, p. 212, 213, pl. 14, figs. 6-6c. 1968 Composita ovata, Sturgeon and Hoare, p. 57, 58, pl. 18, figs. 11- 18 Material and occurrence: 24 specimens [PRB-(50-73)], Unit (II) and (III). Measurements: Dimensions in mm are given in Table.4. Diagnosis: The medium-sized shells of C. ovata can be confused with C. argentea (Shepard, 1838). However, dorsal outline in C. ovata is more oval. This species shows weak fold on the anterolateral margins, a rectimarginate anterior commissure, and a larger ventral beak. Composita trilobata Dunbar and Condra, 1932 (Fig.22) 1932 Composita trilobata, Dunbar and Condra, p. 372-373, pl. 43, fig. (25-31). 1964 Composita trilobata, Grinnell and Andrews, p. 236. Material and occurrence: Three specimens [PRB-(74-76)], Unit (II). Measurements: Dimensions in mm are given in Table.5. Diagnosis: This species is represented by subtriangular, small-sized shells, with a more convex dorsal valve. The distinctive trilobat character of C. trilobata is produced by a narrow, high fold, flanked by two well-developed sulci on the dorsal valve. 5

Composita trinuclea (Hall, 1856) (Fig.23a-e) 1856 Terebratula trinuclea, Hall, p. 7. 1906 Seminula trinuclea, Clark and Mathews, pl. 18, fig. 11. 1914 Composita trinuclea, Weller, p. 486-488, pl. 81, fig. (16-45). 1937 Athyris (Composita) trinuclea, Nalivkin, p. 124, pl. 37, fig. (1-7). 1983 Composita trinuclea, Abramov and Grigorjeva, p. 55, pl. 9, fig. 10. Material and occurrence: One specimen [PRB-77], Unit (II). Measurements: The recorded specimen is 21.5 mm long, 19.5 mm wide, and 13.7 mm thick. Diagnosis: The shell is subovate in outline, with two equally-convex valves. C. trinuclea can be confused with C. trilobata as both show a trilbate character. However, C. trinuclea differs in having suboval to subpentagonal outline, being more transverse, with wider dorsal fold, flanked by deeper sulci. Genus: Athyris M’Coy, 1844 Athyris sp. (Fig.23f-l) Material and occurrence: Three specimens [PRB-(78-80)], Unit (II). Measurements: Dimensions in mm of the 3 specimens are given in Table 6. Diagnosis: Small-sized, biconvex shells, with elongated to oval outline and rectimarginate anterior commissure line are assigned to genus Athyris, based on the outline of the shell. The collected specimens show some similarity to A. supervittata (Tien, 1938b), but the main difference is the longer than wider profile of A. supervittata. Order: Productida Sarytcheva and Sokolskaya, 1959 Family: Linoproductida Stehli, 1954 Genus: Linoproductus Chao, 1927 Linoproductus lineatus (Waagen, 1884) 6

(Fig.24a-c) 1884 productus lineatus, Waagen, p. 673, pl. 66, fig. 1, 2. 1927 Linoproductus lineatus, Chao, p. 129, pl. 15, fig. 27, 28. Material and occurrence: Four specimens [PRB-(87-90)], Unit (IV). Measurements: Dimensions in mm are given in Table.7. Diagnosis: This species is represented by various ventral valves. These valves are moderately-convex, subrectangular in outline, with strongly-incurved beak and illdefined median sulcus. Strong costellae and the growth rugae show resemblance to those of L. cora (d’Orbigny), but the rectangular outline and the less conspicuous median sulcus are characteristic of L. lineatus. Genus: Antiquatonia Miloradovich, 1945 Antiquatonia coloradoensis (Girty, 1903) (Fig.24d-i) 1903 Productus inflatus Girty, p. 359-361, pl. 3, figs. 1-3. 1910 Productus inflatus var. coloradoensis (Girty), Girty, p. 215. 1927 Productus coloradoensis (Girty), Girty, pl. 27, fig. 17. 1938 Dictyoclostus coloradoensis (Girty), Sutton, p. 563. 1954 Antiquatonia coloradoensis (Girty), Stehli, p. 317, pl. 26, figs. 9–11. Material and occurrence: Three specimens [PRB-(94-96)], Unit (IV). Measurements: Dimensions in mm are given in Table.8. Diagnosis: The shells are strongly concavo-convex, and subcircular in outline. The strong costellae are the main surface ornamentation. A. coloradoensis and A. morrowensis exhibit the same size and shape (Mather, 1915). However, the recorded specimens of A. coloradoensis show more convexity, deeper sulcus, and coarser radial ornamentation. Order: Orthotetida Waagen, 1884 Superfamily: Orthotetoidea Waagen, 1884 Family: Pulsiidae Cooper and Grant, 1974 7

Genus: Schellwienella Thomas, 1910 Schellwienella crenistria (Phillips, 1836) (Fig.24j) 1836 Spirifer crenistria, Phillips, pl. 9, Fig. 6. 1836 Leptaena senilis, Phillips, pl. 9, Fig. 5. 1844 Orthis bechei, M’Coy, pl. 22, Fig. 3. 1844 Orthis comate, M’Coy, pl. 22, Fig. 5. 1844 Orthis caduca, M’Coy, pl. 22, Fig. 6. 1861 Streptorhynchus crenistria (Phillips), Davidson, p. 124, pl. 26, Fig. 1. 1894 Derbyia aff. senilis Phillips, Schellwien, p. 71, pl. 7, Figs. 7 and 8. 1900 Orthothetes crenistria (Phillips), Frech and Arthaber, p. 200, pl. 16, Fig. 6. 2002 Schellwienella crenistria (Phillips), Haydukiewicz and Muszer, p. 25, Fig. 8 (A). Material and occurrence: Two specimens [PRB-(101-102)], Unit (III). Measurements: Dimensions in mm are given in Table.9. Diagnosis: The shells have subcircular to subpentagonal outline, with straight hinge line shorter than the greatest width of the shell. Strong costellae ornamente the surface. S. crenistria is similar in size and ornamentation to S. burlingtonensis (Weller, 1914). However, S. burlingtonensis has more rounded outline, and decidedly cardinal termination in both valves. Family: Derbyiidae Stehli, 1954 Genus: Derbyia Waagen, 1884 Derbyia robusta Hall, 1858 (Fig.24k) 1858 Orthis robusta, Hall, p. 713, 714, pl. 28, fig. 5a-d. 1892 Derbyia robusta Hall, Hall and Clarke, 1892, pt 10, fig.12-15. Material and occurrence: One specimen [PRB-103], Unit (II). 8

Measurements: The recorded specimen is 14 mm long, and 19 mm wide. Diagnosis: This species is represented by on pedicle valve. The recorded valve is elliptical in outline, with convex umbonal region. The external sculpture is represented by narrow costellae, separated by wide sulci.

Derbyia crassa Meek and Hayden, 1858 (Fig.24l) 1858 Orthisina crassa, Meek and Hayden, p. 261. 1858 Orthis crenistria, Marcou, p. 49. 1861 Streptorhynchus umbraculum, Newberry, pt. 3, p. 125. 1865 Orthis richmonda, McChesney, pl. 1, fig. 5a-c. 1872 Hemipronites crassus, Meek, p. 174, pl. 5, fig. 10a-c; pl. 8, fig. 1. 1884 Derbyia crassa, Waagen, p. 592. Material and occurrence: One specimen [PRB-104], Unit (II). Measurements: The recorded specimen is 21 mm long. Diagnosis: This species is only represented by a broken, approximately-convex pedicle valve, with a surface ornamented by regular, rounded costellae. 6. Microfacies and depositional environment interpretation Throughout the late Carboniferous Rod El-Hamal Formation, the clastic rock constituents are more dominant than carbonate (Fig.3). The clastic rocks have been previously studied by Darwish (1992). He described two main lithofacies, shale/mudstone facies and sandstone facies. The multicolored, fossiliferous, thin- and flat-bedded shale facies with rippled and bioturbated top surfaces reflect tidal flats to shallow sheltered subtidal environment, sometimes with sabkha conditions indicating open coasts of low relief and low energy. The sandstone facies are cross-bedded, rippled and partly bioturbated with fining upwards sandstone bodies reflecting tidal channel domains.

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The carbonate rocks are mainly distributed in the middle part of Unit II and the upper part of Unit III and IV of Rod El-Hamal Formation and are usually intercalated with greenish marls. The microfacies analysis of the fossil-bearing limestone beds allowed the identification of the grain types and textural features characterizing four microfacies types. SandyCrinoid-Bryozoan grainstone and Bryozoan grainstone microfacies are recorded from the limestone beds of Unit III, and Bioclastic Grainstone and Bryozoan-Crinoid rudstone microfacies are recorded from the limestone beds of Unit IV. The recognized microfacies types are compared with the standard microfacies types of Flügel (2004), and are related to the possible facies zones suggested by Wilson (1975). The identified microfacies are: 6. 1. Sandy-Crinoid-Bryozoan Grainstone (MFT-1) Description: The main skeletal components are represented by trepostome bryozoans, with rounded to oval zooecia (Fig.25a), and fenestrate bryozoans, with elongated zooecia, which are filled with sparite cement (Fig.25c). Crinoidal and echinoidal fragments are also present, with syntaxial cement, constituting a significant portion of this microfacies (Fig.25b). Relatively high amount of poorly-sorted, subangular to angular quartz grains of sand-sized are present (Fig.52d). The ground mass is represented by blocky sparite cement. Interpretation: The occurrence of bryozoans, crinoids and echinoids indicate open marine environment with normal oxygen and salinity conditions (Khan et al., 2014). Nevertheless, the abundance of the quartz grains reflects fair influx of terrigenous materials. Therefore, this microfacies suggests a depositional environment of a shelf lagoon with restricted water circulation (El-Sorogy et al., 2017; Torres-Martinez et al., 2017). It is correlated to the Standard Microfacies Type (SMF 10) of Flügel (2004) and Facies Zone (FZ 8), Restricted shelf interior, of Wilson (1975). 6. 2. Bryozoan Grainstone (MFT-2) Description: This microfacies is mainly composed of fenestrate bryozoan colonies with relatively oval zooecia, filled with sparite cement (Fig.25e). The trepostome bryozoans are partially subjected to dissolution and latter filled with precipitated calcite. Some fragments of brachiopod shells are also recorded (Fig.25f). The matrix is micrite which was recrystallized into neomorphosed sparry calcite cement with blocky appearance. Interpretation: The faunal assemblages in this microfacies are represented only by bryozoans and few brachiopod shells, which indicate an open marine inner shelf 10

environment (Torres-Martinez et al., 2017). This microfacies is similar to the Standard Microfacies Type (SMF 11) of Flügel (2004) and Facies Zone (FZ 7), Open-marine shelf interior, of Wilson (1975). 6. 3. Bioclastic Grainstone (MFT-3) Description: It consists of highly diversified biota. Fistuliporoid and fenestrate bryozoans, including bifoliates Stictopora are widely distributed (Fig.26a,b). Crinoid fragments with well-preserved alignment traces (Fig.26c), echinoid plates and spines with syntaxial cement (Fig.26d) are relatively abundant. Shells of bivalve with foliated structure (Plate 2, Fig. 1), gastropods (Fig.26e), brachiopod shells and fusulinid foraminifers are also present. Several trilobite fragments are recorded for the first time in thin sections from Rod El-Hamal Formation (Fig.26f). Few poorly-sorted, angular to subrounded quartz grains are recorded (Fig.26d). The ground mass is blocky sparry calcite cement. Interpretation: The occurrence of crinoids, echinoids, bryozoans, brachiopods, gastropods and trilobites indicate open marine environment of well-oxygenated, normal saline and highly nutrient-supplied water (Khan et al., 2014). The quartz grains reflect an influx of terrigenous materials. The mostly suggested depositional environment is an open-marine inner shelf environment (Torres-Martinez et al., 2017). This microfacies is similar to the Standard Microfacies Type (SMF 11) of Flügel (2004) and Facies Zone (FZ 7), Open-marine shelf interior, of Wilson (1975). 6. 4. Bryozoan-Crinoid Rudstone (MFT-4) Description: It is made up of a diverse biota. One of the most common constituents is fenestrate bryozoan colonies with sub-rectangular zooecia (Fig.27a). Trepostome bryozoans are also found (Fig.27b). Gastropod shells were dissolved and formed moldic porosity, filled with precipitated calcite (Fig.27c). Crinoidal particles are found in frequent amount, represented by ossicles and arm plates, some with micrite-filled borings (Fig.27d). Echinoid fragments and brachiopod shells are also present (Fig.27e). The bivalve shells with foliated structure are also recorded, some of which are encrusted by bryozoans (Fig.27f). The foraminifers are represented by planispirally-coiled shells. The foraminifera chambers are filled with microsparite. Terrigenous particles are few, fine to medium, subangular to angular quartz grains in the sand size. The rock matrix is a micrite that underwent recrystallization into sparry calcite cement. Interpretation: The diversified faunal content in this microfacies, represented by crinoids, echinoids, bryozoans, brachiopods, and gastropods, suggest an open marine 11

environment, with well-oxygenation and normal salinity conditions, most probably connected to a shallow lagoon. Bryozoan encrustation on the bivalve shells and the boring activity in the shell fragments reflect quiet environment with low rate of sedimentation, and the quartz grains reflect an influx of terrigenous detritus. The most possible depositional environment is inner shelf environment, with restricted water circulation (El-Sorogy et al., 2017). This microfacies is similar to the Standard Microfacies Type (SMF 12) of Flügel (2004) and Facies Zone (FZ 8), Restricted shelf interior, of Wilson (1975). 7. Predation on brachiopod shells Predation in the fossil record can indicate the behaviors and relationships between predator and prey (Kowalewski et al., 1997), evolutionary processes and the adaptive differences between them (Leighton, 2003a). In particular, Biting and crushing predation is considered as a major source of mortality on shelled marine invertebrates throughout the Phanerozoic (Vermeij, 1987; Leighton, 2003b). The brachiopoda predation in the fossil records is mostly preserved as borings and scars produced by the different predators, e.g. gastropods. Such borings and scars on brachiopods have been documented during the whole Paleozoic (Kaplan and Baumiller, 2000). Brachiopods have been the prey for various predators, including crustaceans, gastropods, and cephalopods (Baumiller et al., 2003; Harper et al., 2009; Taddei Ruggiero et al., 2006) which could drill the brachiopod shells (Baumiller and Bitner, 2004; Delance and Emig, 2004). Biting and crushing scars on both pedicle and brachial valves of Athyridida and Orthotetida brachiopod shells are recognized in the studied material. One bite, two bite injuries or complete crushing of both valves are detected (Fig.28). Such damages were most probably the result of some attacks by predators capable of causing a series damage to one or both valves simultaneously. The modern brachiopods show similar shell damage, which is mostly attributed to crustaceans. However, conchostracan arthropods or nautiloids are the most likely predators for the Pennsylvanian brachiopods (Bartlett and Elliott, 2010). 8. Taphonomy of brachiopods The taphonomic analysis applied to the present study includes the disarticulation and size frequency distribution. Disarticulation can reflect the burial rate. This study follows the disarticulation ratio (N/No) of Boucot et al. (1958), where, N is the number of articulated shells and N0 is the original number of all shells present before disarticulation. 8.1. Disarticulation

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Most of brachiopod shells are articulated, including 83 articulated shells (80%) in contrast to 21 disarticulated shells. Shells of Composita ovata, with ventral and dorsal valve similar in size, thickness, and convexity, are mostly articulated. This reflects similar occurrence frequency for both ventral and dorsal valves, which is typical of a normal community with in-situ preservation (Zuschin et al., 2002). This reflects that the studied shells haven’t underwent neither a great transportation nor sorting by waves 8.2. Size-frequency distribution The size frequency distribution of a total 24 shells of Composita ovata was statistically analyzed. Of these, individuals with shell widths between 13.5 and 21 mm are the most frequently counted, up to 80%; while small shells of 8.5–13.5mm in width, probably representing juveniles or subadults, are less frequent. The majority of the 19 measured Composita subquadrata shells (79%) are small, with shell widths of 11.5–16 mm, and they are more numerous than large shells, 16–19 mm in width. For Composita subtilita, 16 shells were analyzed. Individuals with shell widths between 13 and 16 mm are the most frequently counted, up to 63%; while small shells of 12–13 mm in width, are less numerous than large shells, 16–19 mm in width. The size frequency distribution of a total 12 shells of Composita ambigua was analyzed. Such analysis indicated that individuals with shell widths between 14.5 and 17.5 mm are the most frequently counted, up to 58%; while small shells of 12–14.5 mm in width, probably representing juveniles or subadults, are more numerous than the large shells of 17.5–19.5 mm in width (Fig.29). The bell-shaped frequency distribution reflects size-sorting during transportation (Tomašových, 2006). The juvenile and adult shells have almost the same occurrence frequency. Such bell-shaped distributions may be due to preferential destruction of small and more fragile shells than the adult ones (Tomašových, 2004). 9. Paleoecology of bryozoans Bryozoans are sessile colonial metazoans, usually with low-magnesium calcitic skeletons, which are largely resistant against dissolution and diagenesis, occurring most commonly on stable bottom sediments and living only in normal sea water with salinity of 35‰ (Smith et al., 2006). They have been recorded since the Early Ordovician, with fluctuated diversity throughout the Paleozoic (Tuckey, 1990a, 1990b; Gilmour and Morozova, 1999; McCoy and Anstey, 2010). The identified bryozoans from Rod ElHamal Formation show high diversity, represented by fenestrate, trepostome, fistuliporoid and bifoliate types. The high diversity of bryozoans acquire low sedimentation rates and moderate water energy of the shelf, in order to provide a continuous food supply and remove small sediment particles to enhance bryozoan growth 13

(Ziko and Hamza, 1987). Bryozoans colonize flexible and hard substrates, with higher diversity on the hard ones. In conclusion, a diverse bryozoan community indicative to the temperate shelf normal marine environment, most probably with restricted water circulation, as shown by the encrusting bryozoans on some bivalve shells (Fig.6a). 10. Paleobiogeography of brachiopods Some of the common late Carboniferous brachiopod genera from Wadi Rod El-Hamal were selected for palaeobiogeographic study. The tolerant brachiopod genus Derbyia was widespread, with discoveries recorded in various Carboniferous collections all over the world, e.g. Bolivia, Canada, North and South China, Iran, the United Kingdom, the western United States, the North American Midcontinent region, Italy, Spain, South Korea, Libya, Morocco, and Egypt (Fig.30) (Schellwien, 1894; McGugan, 1963; Fantini Sestini, 1966; Sutherland and Harlow, 1967; Smith and Xu, 1988; Kora, 1995; Wendt et al., 2001; Kues, 2002; Lee et al., 2010; Tawadros, 2011; Brand, 2011; Badyrka et al., 2013). The genus Composita was also widely distributed, as recorded from Australia, Belgium, Bolivia, Brazil, Canada, Mexico, Peru, the United Kingdom, the western United States, the North American Midcontinent region, Pakistan, Russia, China, Iran, Libya, and Egypt (Fig.30) (Newell, 1953; Easton et al., 1958; Gehrig, 1958; McGugan, 1963; Gaetani, 1965; Roberts et al., 1976; Kora, 1995; Angiolini et al., 1999; Kabanov, 2003; Chen, 2004; Chen et al., 2005; Chevalier and Aretz, 2005; Bassett and Bryant, 2006; Badyrka et al., 2013). A more or less similar distribution characterizes the genus Linoproductus, which has been reported from Australia, Bolivia, Brazil, Canada, Mexico, Peru, North and South China, Ireland, Italy, Spain, South Korea, Russia, the United Kingdom, the western United States, the North American Midcontinent region, and Morocco (Fig.31) (Schellwien, 1894; Newell, 1953; Gehrig, 1958; Raasch, 1958; Hudson et al., 1966; Barskov and Morozov, 1996; Wendt et al., 2001; Chen, 2004; Chen et al., 2004; ElAlbani et al., 2005; Lee et al., 2010; Brand, 2011; Badyrka et al., 2013). The genus Antiquatonia has been recorded from Australia, Canada, China, Ireland, Spain, Russia, the United Kingdom, the western United States, the North American Midcontinent region, and Egypt (Fig.31) (Hudson et al., 1966; Brew and Beus, 1976; Roberts et al., 1976; Kora, 1995; Barskov and Morozov, 1996; Carter and Poletaev, 1998; Chen, 2004; El-Albani et al., 2005; Brand, 2011). 11. Paleobiogeography of bryozoans

14

Bryozoan distribution patterns during the Carboniferous show widespread dispersal and highly fluctuated diversity. During the Early Carboniferous, warm climatic conditions prevailed and carbonate-producing organisms displayed a great diversity, especially bryozoans. Tournaisian bryozoans showed vast distribution as described from Kazakhstan, Mongolia, the Russian Platform, Eastern Transbaikalia, Ireland, Germany, the Donetz Basin, Poland, and China (Yang et al., 1988; Tolokonnikova, 2012). Visean and Serpukhovian bryozoans had a greater diversity and broader distribution than those of Tournaisian time, such fauna were recorded from Kazakhstan, Transbaikalia, Mongolia, Japan, the Donetz Basin, Russian Platform, and Spain (Sakagami, 1962, 1963; Ernst, 1998; Ernst and Rodriguez, 2013). The late Carboniferous time witnessed a fluctuating generic diversity of the bryozoan assemblages, represented in the study area by different species of the Orders Cystoporata (Astrova, 1964), Fistuliporoida, Trepostomata and Fenestrata (Ulrich, 1882). The dispersal of such fauna throughout different parts of the world is represented in (Fig.32). During the Bashkirian, the world climate witnessed a rapid cooling which led to a marked depletion in the generic diversity of bryozoan assemblages. However, many families were still represented as documented from the Russian platform, Northwestern Europe (including United Kingdom, Belgium, Germany, and Poland), Kazakhstan, Jaban, and northern Gondwanan margin (Sugiyama and Nagai, 1994; Aretz and Herbig, 2003a; Cózar et al., 2016). Resumption in the generic diversity characterized the Moscovian and Kasimovian bryozoans, many of them were cosmopolitan. Such fauna were recorded from the Russian platform, the Donetz basin, the Frankilian shelf (Greenland), Southwestern shelves of the United States, West Texas, Kansas, Andean Sea, North America Cordillera, Argentina, South China, northern Gondwana margin (Engel, 1975; Ernst and Minwegen, 2006; Ahlborn and Stemmerik, 2015; Yao and Wang, 2016; Cózar et al., 2016). During the Gzhelian, bryozoan assemblages showed further increase in the generic diversity, especially in the second half of this stage. They were identified from the Russian platform, the Donetz basin, Southwestern shelves of the United States, West Texas, the Carnic Alps, Thailand, Malaysia, South Wales, South China, and the Norwegian Barents Sea (Engel, 1975; Alipour et al., 2012; Yao and Wang, 2016; Di Lucia et al., 2017). Conclusion 1. Based on lithostratigraphic criteria, the Rod El-Hamal Formation at Wadi Araba area, on the western side of the Gulf of Suez, Egypt, is divided into five rock units (I–V). The 15

collected brachiopod specimens are assigned to 12 monospecific genera, belonging to the orders of Athyridida, Orthotetida, and Productida, in descending order of individual abundance. 2. The carbonates of the Rod El-Hamal Formation are distinguished into grainstone and rudstone. The most common bioclasts belong to bryozoans, crinoids, echinoids, brachiopods, molluscs, foraminifers and trilobites. Microfacies analysis revealed a deposition in a restricted to open-marine inner shelf environment for the studied carbonates. 3. Some athyridida and orthotetida brachiopod shells had biting and crushing scars on both valves. Such damage was the result of some predation attempts, which are mostly attributed to conchostracan arthropods or nautiloids that were the most common predators for the Pennsylvanian brachiopods (Bartlett and Elliott, 2010). 4. The taphonomic analysis included the disarticulation and size frequency distribution. The disarticulation ratio (80%) indicate a normal community with in-situ preservation (Zuschin et al., 2002), which hasn’t underwent neither a great transportation nor sorting by waves. The bell-shaped size-frequency distribution reflects size-sorting, which may be due to preferential destruction of small and more fragile shells than the adult ones (Tomašových, 2004). 5. Some common late Carboniferous brachiopod genera were selected from the identified specimens for palaeobiogeographic study, including Derbyia, Composita, Linoproductus, and Antiquatonia. Such study revealed their distribution all over the world, e.g. Australia, Belgium, Bolivia, Canada, North and South China, Iran, the United Kingdom, the western United States, the North American Midcontinent region, Italy, Spain, South Korea, Libya, Morocco, and Sinai, Egypt. 6. The paleobiogeographic study of the late Carboniferous bryozoans revealed a great variation in the generic diversity. The rapid cooling of the world during the Bashkirian led to a marked depletion in the generic diversity of bryozoan assemblages. However, many families were still represented. Such reduction was followed during the Westphalian by a resumption in the generic diversity. During the Stephanian, bryozoan assemblages showed further increase in the generic diversity, especially in the second half of this stage.

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Roberts, J., Hunt, J. W., and Thompson, D. M., 1976. Late Carboniferous marine invertebrate zones of eastern Australia. Alcheringa 1:197-225. Said, R., 1962. The Geology of Egypt. Elsevier, p. 377. Sakagami, S., 1962. Lower Carboniferous Bryozoa from the Omi limestone, Japan. Part 1. Discovery of the Profusulinella zone, and descriptions of Profusulinella, Cyclostomata, Trepostomata and Fenestella. Trans. Proc. Paleontol. Soc. Jpn 48, 321–330. Sakagami, S., 1963. Lower Carboniferous Bryozoa from the Omi limestone, Japan. Part 2. Successive description of Cryptostomata. Trans. Proc. Paleontol. Soc. Jpn 49, 15–24. Sarycheva, T. G., and Sokolskaya, A. N., 1959. O klassifikatsin lozh- noporistykh brakhiopod [On the classification of pseudopunctate brachiopods]. Akademiya Nauk SSSR, Doklady (Moscow), 125(1):181- 184. (In Russian) Schellwien, E., 1894. Uber eine angebliche Kohlen-Kalkfauna aus der Agyptischenarabischen Wuste. Zeitschrift der Deutschen Geologischen Gesellschaft. Berlin 46, 68–78. Scholle, P. A., 2003. A Color Guide to the Petrography of Carbonate Rocks: Grains, textures, porosity, diagenesis. American Association of Petroleum Geologists Memoir 77, 470 pp. Schweinfurth, G., 1883. Sur la decouverte d’une faune Paleozoique dans le gres d’ Egypte. Bulletin Institute de l’Egypte 6 (2), 239–255. Shen, S., 2018. Global Permian brachiopod biostratigraphy: an overview. Geological Society, London, Special Publications, 450, 289-320. Shepard, C. V., 1838. Geology of upper Illinois. Americal Journal of Science. ser. 1, vol. 34, no. 1, p. 134-161. Smith, A. B., and Xu, J., 1988. Paleontology of the 1985 Geotraverse, Lhasa to Golmud. Philosophical Transactions of the Royal Society of London A 327: 53-105. Smith, A. M., Key, M. M., and Gordon, D. P., 2006. Skeletal mineralogy of bryozoans: taxonomic and temporal patterns. Earth-Science Reviews 78(3–4): 287–306. Sowerby, G. B., 1823-1834. The genera of recent and fossil shells, for the use of students. In: Concology and Geology, Commenced by J. Sowerby and continued by G.B. Sowerby, 2 vols., London.

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Stehli, F. G., 1954. Lower Leonardian Brachiopoda of the Sierra Diablo: Am. Mus. Nat. History Bull, v. 105, art. 3, 358 p., pls. 17-27, 55 figs. Sturgeon, M. T., and Hoare, R. D., 1968. Pennsylvanian brachiopods of Ohio: Ohio, Div. Geol. Survey Bull. 63, 95 p., 22 pls., 14 figs. Sugiyama, T., and Nagai, K., 1994. Reef facies and paleoecology of reef-building corals in the lower part of the Akiyoshi Limestone Group (Carboniferous), Southwest Japan. Courrier Forschunginstitut Seckenberg 172, 231–240. Sutherland, P. K., and Harlow, F. H., 1967. Late Pennsylvanian Brachiopods from NorthCentral New Mexico. Journal of Paleontology, 41: 1065-1089. Sutton, A. H., 1938. Taxonomy of Mississippian Productidae: Jour. Paleontology, v. 12, no. 6, p. 537-569, pls. 62-66, 2 figs. Taddei Ruggiero, E., Buono, G., and Raia, P., 2006. Bioerosion on brachiopod shells of a thanatocoenosis of Alboràn Sea (Spain). Ichnos 13, 175–184. Tawadros, E., 2011. Geology of North Africa. CRC Press. Tien, C. C., 1938. Devonian brachiopod of Human. Paleontologia Sinica, N. S., ser. B, no. 4. Thomas, I., 1910. The British Carboniferous Orthotetinae. Great Britain Geological Survey Memoir, 1: 83-134. Tolokonnikova, Z., 2012. Early Carboniferous bryozoans fromWestern Siberia, Russia. In: Ernst, A., Schäfer, P., Scholz, J. (Eds.), Bryozoan Studies 2010Lecture Notes in Earth System Sciences 143. Springer-Verlag, Berlin Heidelberg, pp. 385-399. Tomašových, A., 2004. Postmortem durability and population dynamics affecting the fidelity of brachiopod size-frequency distributions. Palaios 19, 477-496. Tomašových, A., 2006. Linking taphonomy to community-level abundance: insights into compositional fidelity of the Upper Triassic shell concentrations (Eastern Alps). Palaeogeography, Palaeoclimatology, Palaeoecology 235, 355-381. Torres-Martínez, M. A., Barragan, R., Sour-Tovar F., and Gonzalez-Mora F., 2017. Depositional paleoenvironments of the Lower Permian (upper Cisuralian) carbonate succession of Paso Hondo Formation in Chiapas State, southeastern Mexico. Paleobiogeographical implications. J. S. Am. Earth Sci. 79, 254-263.

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Tables and Figures captions: Table 1: The dimensions in mm for the 12 specimens of Composita ambigua Table 2: The dimensions in mm for the 16 specimens of Composita subtilita Table 3: The dimensions in mm for the 19 specimens of Composita subquadrata Table 4: The dimensions in mm for the 24 specimens of Composita ovata Table 5: The dimensions in mm for the 3 specimens of Composita trilobata Table 6: The dimensions in mm for the 3 specimens of Athyris sp. Table 7: The dimensions in mm for the 4 specimens of Linoproductus lineatus. Table 8: The dimensions in mm for the 3 specimens of Antiquatonia coloradoensis. Table 9: The dimensions in mm for the 2 specimens of Schellwienella crenistria.

Fig. 1: Carboniferous paleogeographic maps of Egypt, (a) during latest Tournaisian– earliest Visean (~342 Ma); (b) late Moscovian (~305 Ma), modified after (Guiraud and Bosworth, 1999; Guiraud et al., 2005). Fig. 2: Geological map of the study area, modified after Conoco, 1987. Fig. 3: Composite section of the five units comprising the Rod El-Hamal Formation. (a) General view showing the complete section of Unit (V). (b, c) General view, and close view showing the fossiliferous horizon of Unit (IV). (d) Panoramic view of Unit (III), photo looking NE. (e) Fossiliferous limestone of Unit (III) with well-preserved brachiopod shells. (f) Panoramic view of Unit (II), photo looking SE. (g) Field photograph showing the varicolored shales, alternating with ferruginous sandstones of Unit (I). The arrows point at the brachiopod horizons. Fig. 4: Brachiopod composition of orders based on species richness (a), and individual abundance (b). Fig. 5: Composita composition of species based on the individual abundance. Fig. 6: (a) Scatter diagram with Width plotted against Length for Composita ambigua, showing the increase in width during the growth of the shell. (b) Width/Thickness ratio versus Length diagram of Composita ambigua, showing that the width increases faster than thickness during the growth of the shell. 28

Fig. 7: (a) Plot of Width/Surface length ratio against Surface length for Composita ambigua. (b) Plot of Width/Length ratio against Length for Composita ambigua. Fig. 8: Plot of Thickness/Length ratio against Length for Composita ambigua. Fig. 9: Composita ambigua (Sowerby, 1823), (a-e) PRB-3; (a) dorsal view, (b) ventral view, (c) lateral view, (d) posterior view, (e) anterior view. (f-j) PRB-7; (f) dorsal view, (g) ventral view, (h) lateral view, (i) posterior view, (j) anterior view. (k, l) PRB-10; (k) dorsal view. (l) lateral view. Scale bar= 1 cm. Fig. 10: (a) Scatter diagram with Width plotted against Length for Composita subtilita, showing a regular increase in width during the growth of the shell. (b) Width/Thickness ratio versus Length diagram of Composita subtilita, showing that the width increases faster than thickness during the growth of the shell. Fig. 11: (A) Plot of Width/Surface length ratio against Surface length for Composita subtilita. (B) Plot of Width/Length ratio against Length for Composita subtilita. Fig. 12: Plot of Thickness/Length ratio against Length for Composita subtilita. Fig. 13: Composita subtilita (Hall, 1852), (a-e) PRB-18; (a) dorsal view, (b) ventral view, (c) lateral view, (d) posterior view, (e) anterior view. (f-j) PRB-21; (f) dorsal view, (g) ventral view, (h) lateral view, (i) posterior view, (j) anterior view. (k, l) PRB-25; (k) dorsal view. (l) posterior view. Scale bar= 1 cm. Fig. 14: (a) Scatter diagram with Width plotted against Length for Composita subquadrata, showing the increase in width during the growth of the shell. (b) Width/Thickness ratio versus Length diagram of Composita subquadrata, showing that the width increases faster than thickness during the growth of the shell. Fig. 15: (a) Plot of Width/Surface length ratio against Surface length for Composita subquadrata. (b) Plot of Width/Length ratio against Length for Composita subquadrata. Fig. 16: Plot of Thickness/Length ratio against Length for Composita subquadrata. Fig. 17: Composita subquadrata (Hall, 1858), (a-e) PRB-34; (a) dorsal view, (b) ventral view, (c) lateral view, (d) posterior view, (e) anterior view. (f-j) PRB-38; (f) dorsal view, (g) ventral view, (h) lateral view, (i) posterior view, (j) anterior view. (k, l) PRB-41; (k) dorsal view. (l) lateral view. Scale bar= 1 cm. Fig. 18: (a) Scatter diagram with Width plotted against Length for Composita ovata, showing a regular increase in width during the growth of the shell. (b) Width/Thickness 29

ratio versus Length diagram of Composita ovata, showing that the width increases faster than thickness during the growth of the shell. Fig. 19: (a) Plot of Width/Surface length ratio against Surface length for Composita ovata. (b) Plot of Width/Length ratio against Length for Composita ovata. Fig. 20: Plot of Thickness/Length ratio against Length for Composita ovata. Fig. 21: Composita ovata (Mather, 1915), (a-e) PRB-50; (a) dorsal view, (b) ventral view, (c) lateral view, (d) posterior view, (e) anterior view. (f, g) PRB-63; (f) lateral view, (g) posterior view. (h-l) PRB-69; (h) dorsal view, (i) ventral view, (j) lateral view, (k) posterior view. (l) anterior view. Scale bar= 1 cm. Fig. 22: Composita trilobata (Dunbar and Condra, 1932), (a-d) PRB-74; (a) dorsal view, (b) ventral view, (c) lateral view, (d) posterior view. (e-h) PRB-75; (e) dorsal view, (f) ventral view, (g) lateral view, (h) posterior view. (i-l) PRB-76; (i) dorsal view, (j) ventral view, (k) lateral view, (l) posterior view. Scale bar= 1 cm. Fig. 23: (a-e) Composita trinuclea (Hall, 1856), PRB-77; (a) dorsal view, (b) ventral view, (c) lateral view, (d) posterior view, (e) anterior view. (f-l) Athyris sp., (f, g) PRB-78; (f) dorsal view, (g) ventral view. (h-l) PRB-79; (h) dorsal view, (i) ventral view, (j) lateral view, (k) posterior view, (l) anterior view. Scale bar= 1 cm. Fig. 24: (a-c) Linoproductus lineatus (Waagen, 1884), PRB-(87-90), pedicle valves. (d-i) Antiquatonia coloradoensis (Stehli, 1954), (d-f) PRB-94; (d) pedicle valve, (e) brachial valve, (f) posterior view. (g-i) PRB-95; (g) pedicle valve, (h) brachial valve, (i) posterior view. (j) Schellwienella crenistria (Phillips, 1836); PRB-101, pedicle valve. (k) Derbyia robusta (Hall, 1858), PRB-103, pedicle valve. (l) Derbyia crassa (Meek and Hayden, 1858), PRB-104, pedicle valve. Scale bar= 1 cm. Fig. 25: represents Sandy-Crinoid-Bryozoan Grainstone: (a) Transverse section through a trepostome bryozoan colony filled with sparry calcite cement. (b) Tangential longitudinal section through fenestrate bryozoans; the fenestrules are filled with sparry calcite cement. (c) Crinoid ossicles, the centre is filled with the same material of the ground mass. (d) poorly-sorted, sub-angular to angular quartz grains. (e & f) represent Bryozoan Packstone: (e) Circular and elongated fenestrate bryozoa filled with sparite cement. (f) Longitudinal section through a brachiopod fragment with precipitated phosphate. Fig. 26: represents Bioclastic Grainstone: (a) Fenestrate bryozoans (Br) encrusting a bivalve shell fragment (Bs). (b) Tangential section through fistuliporoid bryozoans shows 30

large circular zooecia filled with micrite and surrounded by completely spar-filled cystopores. (c) Crinoid ossicles, the centre is filled with the same material of the ground mass. (d) Echinoid spine filled with microcrystalline calcite (Es) and Poorly sorted angular to sub-rounded quartz grains (Qz). (e) Longitudinal section through an originally aragonitic gastropod shell, later inverted to calcite, and filled with micrite. (f) A trilobite fragment (Tr) shows characteristic complex curvature of the shell with shape like shepherd’s crooks. Fig. 27: represents Bryozoan-Crinoid Rudstone: (a) Tangential longitudinal section through fenestrate bryozoans. (b) Transverse section through a trepostome bryozoan colony. (c) Transverse section through a single, originally aragonitic gastropod with recognizable “baby-bottom structure” and micrite filling. (d) Crinoid fragment with micrite-filled borings. (e) Longitudinal section through a brachiopod shell wall with lowangle fibrous structure. (f) Bivalve shell fragment (Bs) encrusted by bryozoans (Br) and planisrprally-coiled foraminiferal shell (Fs). Fig. 28: Biting injuries (black arrows). (a-c) in Composita subtilita (Hall, 1852), traces on (a) brachial valve, and (b, c) pedicle valves. (d-f) in Composita subquadrata (Hall, 1858), traces on (d, f) pedicle valves, and (e) brachial valves.(g, h) in Schellwienella crenistria (Phillips, 1836), traces on pedicle valves.(i) in Composita ovata (Mather, 1915), on brachial valve. Scale bar= 1 cm.

Fig. 29: Size frequency distributions of four dominant species. (N) is the total specimens measured in the species. Fig. 30: Global paleobiogeography of brachiopod genera Derbyia and Composita during the late Carboniferous. The base map is modified from https://commons.wikimedia.org/ Fig. 31: Global paleobiogeography of brachiopod genera Linoprodustus and Antiquatonia during the late Carboniferous. The base map is modified from https://commons.wikimedia.org/ Fig. 32: Global Bryozoan paleobiogeography during the late Carboniferous. The base map is modified from https://commons.wikimedia.org/

31

Table.1. The dimensions in mm for the 12 specimens of Composita ambigua

Specimens

Length (L)

PRB-3 PRB-4 PRB-5 PRB-6 PRB-7 PRB-8 PRB-9 PRB-10 PRB-11 PRB-12 PRB-13 PRB-14

19 18 18 19.5 21.5 17 16 16.5 17.5 18 10 19

Surface Length (SL) 27 27.5 26.5 26 32 22 19 22 25 23 19 27

Width (W)

Thickness (D)

W/L

W/SL W/D

D/L

16.5 18.5 17.5 17 18 14 14.5 15.5 15 15.5 12 16

12.5 11.5 12.5 13 13.5 10.5 10 10.2 12.5 10.5 9 12.2

0.87 1.03 0.97 0.87 0.84 0.82 0.91 0.94 0.86 0.86 1.20 0.84

0.61 0.67 0.66 0.65 0.56 0.64 0.76 0.70 0.60 0.67 0.63 0.59

0.66 0.64 0.69 0.67 0.63 0.62 0.63 0.62 0.71 0.58 0.90 0.64

1.32 1.61 1.40 1.31 1.33 1.33 1.45 1.52 1.20 1.48 1.33 1.31

Table.2. The dimensions in mm for the 16 specimens of Composita subtilita

Specimens

Length (L)

PRB-15 PRB-16 PRB-17 PRB-18 PRB-19 PRB-20 PRB-21 PRB-22 PRB-23 PRB-24 PRB-25 PRB-26 PRB-27 PRB-28 PRB-29 PRB-30

18.5 18.5 14 15 16.5 15 16 20 17.5 20 14 17.5 16 16.5 14 17.5

Surface Length (SL) 26 24 20 19 26 --23 27.5 22.5 31 17 24 21 23 17 22.5

Width (W)

Thickness (D)

W/L

W/SL W/D

D/L

14.5 17 14.5 14 13.5 --15 17 16 19 12 15.5 14 15.5 12 14.5

11.5 13 11 10 13 --10 13.2 9 15 9 11.5 10 11 9 12

0.78 0.92 1.04 0.93 0.82 --0.94 0.85 0.91 0.95 0.86 0.89 0.88 0.94 0.86 0.83

0.56 0.71 0.73 0.74 0.52 --0.65 0.62 0.71 0.61 0.71 0.65 0.67 0.67 0.71 0.64

0.62 0.70 0.79 0.67 0.79 --0.63 0.66 0.51 0.75 0.64 0.66 0.63 0.67 0.64 0.69

1.26 1.31 1.32 1.40 1.04 --1.50 1.29 1.78 1.27 1.33 1.35 1.40 1.41 1.33 1.21

Table.3. The dimensions in mm for the 19 specimens of Composita subquadrata Specimens

Length (L)

PRB-31 PRB-32 PRB-33 PRB-34 PRB-35 PRB-36 PRB-37 PRB-38 PRB-39 PRB-40 PRB-41 PRB-42 PRB-43 PRB-44 PRB-45 PRB-46 PRB-47 PRB-48 PRB-49

18.5 19 16 16.5 15.5 13 16 17 14.5 13.5 19 20.5 16 15 16 15.5 16 14 13.5

Surface Length (SL) 24.5 26 25 21 18.5 16 20.5 24.5 19 17 25 25.5 20 18.5 23.5 19 17 18 13

Width (W)

Thickness (D)

W/L

W/SL W/D

D/L

18 19 15.5 15 14 12 14.5 15 12.5 12 16.5 17 13 13.5 15 16 14.5 11.5 12.5

11.5 10.2 11 9 9 8 9 12.2 8.5 8.5 11.5 11 9 9 8.5 9.5 7.5 8 9

0.97 1.00 0.97 0.91 0.90 0.92 0.91 0.88 0.86 0.89 0.87 0.83 0.81 0.90 0.94 1.03 0.91 0.82 0.93

0.73 0.73 0.62 0.71 0.76 0.75 0.71 0.61 0.66 0.71 0.66 0.67 0.65 0.73 0.64 0.84 0.85 0.64 0.96

0.62 0.54 0.69 0.55 0.58 0.62 0.56 0.72 0.59 0.63 0.61 0.54 0.56 0.60 0.53 0.61 0.47 0.57 0.67

1.57 1.81 1.41 1.67 1.56 1.50 1.61 1.23 1.47 1.41 1.43 1.55 1.44 1.50 1.76 1.68 1.93 1.44 1.39

Table.4. The dimensions in mm for the 24 specimens of Composita ovata

Specimens

Length (L)

PRB-50 PRB-51 PRB-52 PRB-53 PRB-54 PRB-55 PRB-56 PRB-57 PRB-58 PRB-59 PRB-60 PRB-61 PRB-62 PRB-63 PRB-64 PRB-65 PRB-66 PRB-67 PRB-68 PRB-69 PRB-70 PRB-71 PRB-72 PRB-73

12.5 22 13 15.5 21.5 13 19.5 17 16.5 20.5 20.5 8.5 15 15.5 21 14.5 18 10 16 20.2 19.5 18 17.2 16

Surface Length (SL) 16 31 16 22.5 29 15.5 27 21.5 22 27 27.5 11 19 22 32.5 23 25 12.5 18.5 28 27 24 24.5 21.5

Width (W)

Thickness (D)

W/L

W/SL W/D

D/L

12 21 11.5 14 19.5 12 18 14.5 15.5 20 20.5 8.5 14 18 20 15.5 17 11 14.5 20 20 17 18 14

6 12.5 8 9 14 7.5 12 10 10.5 11 13.5 5.5 9.5 9 14.5 9 11 5 9.5 13.2 12.5 10.5 10.5 10

0.96 0.95 0.88 0.90 0.91 0.92 0.92 0.85 0.94 0.98 1.00 1.00 0.93 1.16 0.95 1.07 0.94 1.10 0.91 0.99 1.03 0.94 1.05 0.88

0.75 0.68 0.72 0.62 0.67 0.77 0.67 0.67 0.70 0.74 0.75 0.77 0.74 0.82 0.62 0.67 0.68 0.88 0.78 0.71 0.74 0.71 0.73 0.65

0.48 0.57 0.62 0.58 0.65 0.58 0.62 0.59 0.64 0.54 0.66 0.65 0.63 0.58 0.69 0.62 0.61 0.50 0.59 0.65 0.64 0.58 0.61 0.63

2.00 1.68 1.44 1.56 1.39 1.60 1.50 1.45 1.48 1.82 1.52 1.55 1.47 2.00 1.38 1.72 1.55 2.20 1.53 1.52 1.60 1.62 1.71 1.40

Table.5. The dimensions in mm for the 3 specimens of Composita trilobata

Specimens

Length (L)

PRB-74 PRB-75 PRB-76

19.5 16.5 17

Surface Length (SL) 25.5 22 24.5

Width (W)

Thickness (D)

W/L

W/SL W/D

D/L

16 13 17

11 9.5 10

0.82 0.79 0.69

0.63 0.59 0.69

0.56 0.58 0.59

1.45 1.37 1.70

Table.6. The dimensions in mm for the 3 specimens of Athyris sp.

Specimens

Length (L)

PRB-78 PRB-79 PRB-80

15 17.5 13.5

Surface Length (SL) 24 25 20

Width (W)

Thickness (D)

W/L

W/SL W/D

D/L

17 18 17

11.5 13 10.5

1.13 1.03 1.26

0.71 0.72 0.85

0.77 0.74 0.78

1.48 1.38 1.62

Table.7. The dimensions in mm for the 4 specimens of Linoproductus lineatus.

Specimens

Length (L)

Surface Length (SL)

Width (W)

Thickness (D)

W/L W/SL W/D

PRB-87 PRB-88 PRB-89 PRB-90

53.5 43 41.5 46.5

84 56.5 52 66.5

49.5 49.5 46.5 ---

41 --22.5 21.5

0.93 1.15 1.12 ---

0.59 0.88 0.89 ---

1.20 --2.07 ---

No. of D/L costae in 10 mm 0.77 10 --10 0.54 9 0.46 9

No. of costae in 5 mm 6 5 4 4

Table.8. The dimensions in mm for the 3 specimens of Antiquatonia coloradoensis.

Specimens

Length (L)

Surface Length (SL)

Width (W)

Thickness (D)

W/L W/SL W/D

PRB-94 PRB-95 PRB-96

40 27 41

63 45 72.5

50 41.5 56.5

19 17.5 28.2

1.25 1.54 1.38

0.79 0.92 0.78

2.63 2.37 2

No. of D/L costae in 10 mm 0.48 10 0.65 14 0.70 9

No. of costae in 5 mm 5 6 4

Table.9. The dimensions in mm for the 2 specimens of Schellwienella crenistria. Specimens PRB-101 PRB-102

Length (L) 20 14

Width (W) 19 17

Thickness (D) -----

No. of costae in 10 mm 15 18

No. of costae in 5 mm 7 6

•

Paleoecology of Carboniferous brachiopod communities from Wadi Araba area

•

Taxonomy of Carboniferous brachiopoda

•

Paleobiogeography of Carboniferous brachiopod communities

Yasser F Salama (Geology Department, Faculty of Science, Beni-Suef University) Oct 15, 2019

Dear Editor-in-Chief: Journal of African Earth sciences On behalf of my co-authors Abdel Aziz A Mahmoud, Yahia A. El kazzaz and Shahin Abd-Elhameed (Helwan University, Faculty of Science, Geology Department), I confirmed that there are no known conflicts of interest associated with this publication and there has been no significant financial support for this work that could have influenced its outcome. The revised manuscript No. AES7882 " Carboniferous brachiopod communities from Wadi Araba, western side of the Gulf of Suez, Egypt: paleontology, paleoecology and paleobiogeography". Sincerely Yasser Salama PhD (Corresponding author) E-mail: [email protected] Tel: +20-1286331445 Postal address: Geology Department, Faculty of Science,Beni-Suef University, Salah Salim Street, 62514, Beni-Suef, Egypt