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Article

First Report of Middle Eocene Micromorphic Brachiopods from Northeastern Libya: Taxonomy and Paleobiogeography Implications

by
Sayed M. Abd El-Aziz
1,
Ibrahim M. Abd El-Gaied
2,
Mansour H. Al-Hashim
3,
Maria Aleksandra Bitner
4,
Yasser F. Salama
5,
Petra Heinz
6 and
Mostafa M. Sayed
5,6,7,*
1
Department of Geology, Faculty of Science, Fayoum University, Fayoum 63511, Egypt
2
Faculty of Earth Science, Beni-Suef University, Beni-Suef 62511, Egypt
3
Department of Geology and Geophysics, College of Science, King Saud University, P.O. Box 2445, Riyadh 11451, Saudi Arabia
4
Institute of Paleobiology, Polish Academy of Sciences, 00-818 Warszawa, Poland
5
Geology Department, Faculty of Science, Beni-Suef University, Beni-Suef 62511, Egypt
6
Department of Palaeontology, Faculty of Earth Sciences, Geography and Astronomy, University of Vienna, 1090 Vienna, Austria
7
Department of Geology, Faculty of Earth Sciences, Geography and Astronomy, University of Vienna, 1090 Vienna, Austria
*
Author to whom correspondence should be addressed.
Diversity 2024, 16(11), 672; https://doi.org/10.3390/d16110672
Submission received: 14 August 2024 / Revised: 4 October 2024 / Accepted: 8 October 2024 / Published: 31 October 2024
Browse Figures
Versions Notes

Abstract

:
Two brachiopod species, Terebratulina tenuistriata (Leymerie, 1846) and Orthothyris pectinoides (von Koenen, 1984), have been recorded for the first time from the Middle Eocene (Late Lutetian) nummulitic limestone beds in the Darnah Formation at the Wadi Darnah area in Northeast Libya. These brachiopod species are associated here with Nummulites discorbinus (Schlotheim), Nummulites praelyelli (Boukhary and Kamal), and Nummulites bullatus (Schaub) and are widely distributed on this Middle Eocene Nummulites carbonate platform. The two recorded species are common in the Eocene rocks of Europe and the Arabian Gulf. In northern Africa, the brachiopod species Terebratulina tenuistriata (Leymerie) was only recorded from the Middle Eocene (Bartonian) of Egypt, while Orthothyris pectinoides (von Koenen) is firstly recorded from the Middle Eocene of the southern Tethyan Province (NE Libya) in the present work.

1. Introduction

The Middle Eocene rocks in North-east Libya are widely distributed along the northeastern flank of Al Gabal Al Akhdar (Figure 1a,b). The Darnah Formation is mainly composed of shallow marine algal, coral, and nummulitic limestone. The studied formation is conformably overlying the Apollonia Formation and is disconformably underlying the Al Bayda Formation. The larger benthic foraminifera (Nummulites spp.) represent the main constituent of the Middle Eocene deposits and are considered the important index fossils in these rocks. The fossil contents of the Darnah Formation were firstly reported by Gregory [1,2], where the nummulites, Operculina, pelecypods, gastropods, echinoids, and corals are considered the more common faunal assemblages. Since then, most of the previous studies were focused on the nummulites and other larger benthic foraminifera; however, no detailed studies concerning the occurrence of the Middle Eocene brachiopods of Libya have been performed. Bitner and Boukhary [3] recorded Terebratulina tenuistriata (Leymerie, 1846) from the Middle Eocene (Bartonian) of Egypt. In the Arabian Gulf, Bitner and Boukhary [4] recorded Terebratulina tenuistriata (Leymerie, 1846) and Orthothyris pectinoides (von Koenen, 1984) from the Upper Eocene Dammam Formation in the Gebel Hafit anticline, United Arab Emirates. Most records of Eocene brachiopod species are from the northern Tethyan localities. Bitner and Dieni [5] recognized five brachiopod species belonging to five genera. In NW Hungary, seven brachiopod species have been reported from the Middle–Upper Eocene deposits. Additionally, Bitner and Müller [6] recorded six brachiopod species from the Silberberg Formation (Late Eocene–Early Oligocene) in Central Germany. Moreover, Bitner and Müller [7] recorded eleven brachiopod species belonging to ten genera from the Priabonian deposits in eastern Ukraine. Goedert et al. [8] identified Argyrotheca sp. from the Humptulips Formation (Middle Eocene) in western North America. More recently, Terebratulina tenuistriata (Leymerie) was identified in the lower Oligocene deposits of Germany and Italy [9,10]. Sulser et al. [11] recognized a rhynchonellide brachiopod, Erymnaria Cooper [12], in northeastern Switzerland. This study aims to describe a new record of an Eocene brachiopod fauna from the Darnah Formation in the Wadi Darnah section at the southern side of Darnah City. The aim of the present study is to throw more light on the systematic classification of the recorded brachiopod species as well as their paleoecology and paleogeographical distribution.

2. Geologic Setting and Stratigraphy

The study area lies at the northeastern part of the Al Jabal Al Akhdar anticline in Wadi Darnah, at the southern part of Darnah City; coordinates are lat. 32°44′45″ and long. 22°37′26″ (Figure 1a,b). The area under study was subjected to a major tectonic event in the Late Cretaceous and reached its maximum intensity in the Early Paleocene [13,14]. The studied succession in this area is represented by the Darnah Formation, which forms the lower coastal scarp of the Al Jabal Al Akhdar area (Figure 1a,b). Gregory [1] named the Darnah Formation “Derna Limestone” at Darnah escarpment along the district from Darnah in the east to Tubruk at the west. The term Darnah Formation was also introduced by Zert [15] for the carbonate sequence of about 140 m thick at Wadi Darnah, south of Darnah City. It is composed mainly of gray limestones rich in nummulites assemblages of all sizes at the basal part, followed upward by a light gray nummulitic limestone with dolomitic limestone intercalations. In this work, the Darnah Formation is 80 m thick, overlying the Apollonia Formation limestones, and is overlain by the Al Bayda Formation (Figure 2 and Figure 3A). The lower part of this rock unit consists mainly of fossiliferous limestone flooded with echinoids, bivalves, gastropods, nummulites, and micromorphic brachiopods (Figure 3B,C). This part is distinguished by grayish-white dolomitic limestone beds flooded with large amounts of nummulites (Figure 3D,E). While the upper part of Darnah Formation consists mainly of bedded limestone, rich in nummulites, algae, corals, Carolia placunoides, Ostrea multicostata, Turritella spp., Echinolampas spp., and brachiopods (Figure 3F,G). In previous studies, the Darnah Formation was considered to be of Late Lutetian–Priabonian age based on its microfaunal contents [13,16,17,18,19,20,21]. However, based on the Nummulites contents, the Darnah Formation was assigned to the Late Lutetian age where the Bartonian-Priabonian age is absent [22].
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Figure 1. (a) Location of the study area (after El Mehaghag and Ashahomi, [23]); (b) Geologic map of Gabal Al Akhdar (after: Klen, [24]; Rohlich, [16]; and Zert, [15]).
Figure 1. (a) Location of the study area (after El Mehaghag and Ashahomi, [23]); (b) Geologic map of Gabal Al Akhdar (after: Klen, [24]; Rohlich, [16]; and Zert, [15]).
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3. Materials and Methods

A complete section of the Darnah Formation is studied carefully to distinguish the occurrence of brachiopod species. The studied section was sampled for the micromorphic brachiopod analysis. Twenty-four samples were washed over a 200 μm mesh sieve. The residue was dried, and then the brachiopod species were hand-picked and described using a stereo-binocular microscope (OPTIKA, Via Rigla, 30, Bergamo, Italy). One hundred twenty-two brachiopod specimens have been investigated, including 104 articulated and 18 inarticulated valves. The brachiopod specimens were photographed at Assiut University using the scanning electron microscope (JSM 5400 LD). The studied specimens were kept in the Faculty of Earth Science, Beni-Suef University.

4. Results

Two brachiopod species have been identified for the first time from the Middle Eocene (Late Lutetian) nummulitic limestone beds of the Darnah Formation at the Wadi Darnah area in North-east Libya. The identified species are vertically distributed along the whole studied successions (Figure 4). These species were systematically described as follows:

Systematic Palaeontology

Phylum Brachiopoda Duméril, 1805
Subphylum: Rhynchonelliformea Williams, Carlson, Brunton, Holmer, and Popov, 1996
Class: Rhynchonellata Williams, Carlson, Brunton, Holmer, and Popov, 1996
Order: Terebratulida Waagen, 1883
Superfamily: Cancellothyridoidea Thomson, 1926
Family: Cancellothyrididae Thomson, 1926
Subfamily: Cancellothyridinae Thomson, 1926
Genus: Terebratulina d’Orbigny, 1847
Type species: Anomia retusa Linnaeus, 1758
Terebratulina tenuistriata (Leymerie, 1846)
(Figure 5. 1–12)
Ref. [25] Terebratula tenuistriata Leymerie, p. 363, pl. 15, Figure 11.
Ref. [26] Terebratulina tenuistriata (Leymerie)—Bitner, pp. 118–120, Figures 2, 3, 4A–F and 5A–G (cum syn.)
Ref. [27] Terebratulina tenuistriata (Leymerie)—Bitner and Dulai, p. 33, Figures 4.1–8.
Ref. [3] Terebratulina tenuistriata (Leymerie)—Bitner and Boukhary, p. 396, Figure 3a–f.
Ref. [28] Terebratulina tenuistriata (Leymerie)—Dulai et al., p. 185, pl. 3, Figures 1–11.
Ref. [29] Terebratulina tenuistriata (Leymerie)—Dulai, pp. 299–300, Figure 4.
Ref. [30] Terebratulina tenuistriata (Leymerie)—Bitner et al., pp. 122–124, Figure 3a–c.
Ref. [4] Terebratulina tenuistriata (Leymerie)—Bitner and Boukhary, Figure 2a,b.
Ref. [6] Terebratulina tenuistriata (Leymerie)—Bitner and Müller, pp. 677–678, Figure 4a–f.
Ref. [31] Terebratulina tenuistriata (Leymerie)—Bitner et al., p. 3, Figures 2 and 4–6.
Ref. [7] Terebratulina tenuistriata (Leymerie)—Bitner and Müller, p. 214, Figure 3.
Ref. [32] Terebratulina tenuistriata (Leymerie)—Astibia et al., p. 10, Figure 3a–d.
Ref. [10] Terebratulina tenuistriata (Leymerie)—Bitner et al., p. 171, Figure 4B–G.
Ref. [33] Terebratulina tenuistriata (Leymerie)—Dulai et al., p. 58, Figures 34–36.
Ref. [9] Terebratulina tenuistriata (Leymerie)—Bitner and Müller, p. 92, Figure 4H–J
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Figure 4. Distribution chart for the Nummulite and Brachiopod species in the Wadi Darnah section.
Figure 4. Distribution chart for the Nummulite and Brachiopod species in the Wadi Darnah section.
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  • Material: Forty articulated specimens and six separate valves have been collected from the Darnah Formation at the Wadi Darnah area in Northeast Libya.
  • Remarks: This species is relatively common in the studied specimens and is represented by juvenile individuals measured 1.3 mm to 4 mm and 1.3 mm to 4 mm. The specimens are well preserved and consistent with those hitherto described by [3,4,6,7,9,26,27,31]. Its shell is small, with the maximum observed length of about 5 mm. It is elongate oval to subtriangular in outline, nearly equally biconvex, ornamented by numerous coarse radial ribs and fine, concentric growth lines. The subcircular foramen is bordered by two small, triangular deltidial plates. The internal structures were not investigated.
  • Occurrence: Terebratulina tenuistriata recorded from the Middle Eocene Darnah Formation. This species is distributed in the Eocene of Europe: Great Britain [34], Belgium [35,36], France [25,37,38], Spain [26,31,32], Italy [35,39], Austria [29,33], Poland [40,41], Hungary [27,30,42], Romania [33,43,44], Bulgaria [45], Ukraine [7,46]; Africa: Egypt [3] and Asia: the United Arab Emirates [4]; also, it has been reported from the Oligocene of Italy and Germany [6,9,10].
Family: Chlidonophoridae Muir-Wood, 1959
Subfamily: Orthothyridinae Muir-Wood, 1965
Genus: Orthothyris Cooper, 1955
Type species: Orthothyris radiata Cooper, 1955
Orthothyris pectinoides (von Koenen, 1984)
(Figure 6. 1–12)
Ref. [47] Terebratulina pectinoides von Koenen, pp. 1354—1355, pl. 99, Figures 8 and 9.
Ref. [46] Terebratulina pectinoides von Koenen—Zelinskaya, pp. 116–118, pl. 13, Figures 5–19.
Ref. [5] Orthothyris pectinoides (von Koenen)—Bitner and Dieni, pp. 108–109, Figures 5 and 6B–N.
Ref. [27] Orthothyris pectinoides (von Koenen)—Bitner and Dulai, p. 35, Figures 4.9–16.
Ref. [30] Orthothyris pectinoides (von Koenen)—Dulai et al., p. 186, pl. 2, Figure 3.
Ref. [29] Orthothyris pectinoides (von Koenen)—Dulai, pp. 303–304, Figure 7a–n.
Ref. [4] Orthothyris pectinoides (von Koenen)—Bitner and Boukhary, Figure 2c,d.
Ref. [48] Orthothyris pectinoides (von Koenen)—Müller et al., p. 92, pl. 1, Figure 10.
Ref. [6] Orthothyris pectinoides (von Koenen)—Bitner and Müller, pp. 681–684, Figures 8, 9a–p and 10a–k.
  • Material: Sixty-four articulated specimens and 12 separate valves collected from the Darnah Formation at the Wadi Darnah area in Northeast Libya.
  • Remarks: Orthothyris pectinoides is the most common in the studied material, measuring 1.66 mm to 6.25 mm and 1.35 mm to 6.25 mm. It can be easily distinguished from Terebratulina tenuistriata, differing in many characters. The shell of O. pectinoides is biconvex and subcircular in outline; the ventral valve is more convex and ornamented by coarse, radial ribs varying in number from 12 to 18. The hinge line is wide and straight to slightly curved. The beak is high, with sharp ridges and a small, triangular, hypothyroid foramen bordered by elevated deltidial plates. The specimens from the Darnah Formation are similar in size and agree well with those previously described [4,5,6,27,29].
  • Occurrence: Orthothyris pectinoides recorded from the Middle Eocene Darnah Formation in this work. This species is recorded in the Eocene of Europe: Germany [47], Austria [29,30], Ukraine [46], Italy [5], Hungary [27], and Asia: The United Arab Emirates [4]. It has also been found in the Oligocene of Germany [6].
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Figure 5. (1,6–8) Ventral views of the complete specimens; (2–5,10–12) dorsal views of the complete specimens; (9) lateral view of complete specimen.
Figure 5. (1,6–8) Ventral views of the complete specimens; (2–5,10–12) dorsal views of the complete specimens; (9) lateral view of complete specimen.
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Figure 6. (1,3,5–7,9–11) Dorsal views of the complete specimens; (2,4,8,12) ventral views of the complete specimens.
Figure 6. (1,3,5–7,9–11) Dorsal views of the complete specimens; (2,4,8,12) ventral views of the complete specimens.
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5. Discussion

The micromorphic brachiopods (Terebratulina tenuistriata and Orthothyris pectinoides) thrive in the Eocene Darnah Formation. This rock unit was deposited in the inner to middle shelf based on the lithological characteristics as well as the micro- and macrofaunal components. The occurrence of nummulitid assemblages indicates inner to middle shelf environments [13], while the recorded macrofossils from the limestone beds, such as Wakullina lefevrei, Carolia spp., Ostrea spp., and Turritella spp., indicate shallow inner to middle shelf environments of normal marine salinity [49]. The occurrence of macrofossils such as Carolia placunoides, Ostrea multicostata, Turritella spp., and Echinolampas spp. indicates shallow marine water depth [50]; the recent oysters occur at depths of 1–3 m in embayments [51]; the recent genus Turritella is common in tropical, shallow environments on different substrates [52,53,54]. Moreover, the presence of Nummulites spp. suggests an inner to middle neritic environment [55,56]. The repeated banks of Nummulites gizehensis and other Nummulites spp. indicate deposition under shallow marine environmental conditions [55,56,57].
Living brachiopods have very wide depth ranges [58], so they cannot be considered as good paleobathymetric indicators. Recent species of Terebratulina are known from very shallow water to up to 4000 m [58], although they are found at depths of 100 to 500 m [59]. The extinct species Orthothyris pectinoides belongs to the family Chlidonophoridae, of which representatives are deeper-water species, very rarely occurring in shallower water than 300 m [58]. Both species depicted here have a functional pedicle opening; thus, they were attached to the substrate by a pedicle. Modern Terebratulina species often have a pedicle split into rootlets [60,61], which permits them the utilization of a wide range of substrate types. Terebratulina tenuistriata is abundant in the marly deposits. However, in the hard substrate, this species is uncommon [3,26,27,30,34]. Thus, most probably T. tenuistriata was attached to the fine sediments by a pedicle.
In conclusion, all the above-mentioned environmental parameters indicate that the Darnah Formation at Wadi Darnah was deposited in inner to middle neritic environments. This conclusion is in accordance with the opinions of Al Menoufy et al. [22].
Both species identified in the studied material have a very wide geographical distribution (Figure 7). The species Terebratulina tenuistriata is known throughout the whole of Europe, as well as from northern Africa and eastern Asia.
Orthothyris pectinoides has a very similar distribution, although its distribution in Europe is more limited, noted from Italy and Germany to Austria, Hungary, and Ukraine. Such wide distribution should not be surprising, as in the Eocene, those regions were a part of the western Tethys Region with connections to the Indian and Atlantic oceans [62,63]. During the Early Eocene, the high sea level, caused by the sea floor spreading [62], made the free migration of faunas possible.

6. Conclusions

The present work concerns the investigation of brachiopod assemblages from the Middle Eocene (Late Lutetian) in the Darnah Formation, Northeast Libya, based on micro-morphic analysis. Two brachiopod species (Terebratulina tenuistriata and Orthothyris pectinoides) have been recorded for the first time from this rock unit. The ecological preferences and morphological features of the identified species, aligned with the associated large benthic foraminifera (Nummulites spp.), indicate that the deposition of the Darnah Formation took place in inner to middle neritic settings. The recorded species show wide geographic distribution through Europe and the southern Tethys regions.

Author Contributions

Conceptualization, S.M.A.E.-A. and I.M.A.E.-G.; methodology, S.M.A.E.-A., Y.F.S., M.H.A.-H. and I.M.A.E.-G.; validation, M.M.S., Y.F.S., S.M.A.E.-A., M.A.B., P.H. and I.M.A.E.-G.; formal analysis, S.M.A.E.-A., M.A.B. and I.M.A.E.-G.; investigation, M.A.B., Y.F.S., M.M.S. and I.M.A.E.-G.; data curation, M.A.B., P.H., Y.F.S., M.M.S. and I.M.A.E.-G.; writing—original draft preparation, S.M.A.E.-A., M.M.S., Y.F.S., M.A.B. and I.M.A.E.-G.; writing—review and editing, M.A.B., P.H. and M.M.S.; visualization, M.A.B., Y.F.S., M.H.A.-H., P.H., M.M.S. and I.M.A.E.-G.; supervision, M.A.B., P.H. and I.M.A.E.-G. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

No new data were created or analyzed in this study. Data sharing is not applicable to this article.

Acknowledgments

The authors would like to extend their sincere appreciation to the Researchers Supporting Project number (RSPD2024R781), King Saud University, Riyadh, Saudi Arabia. Open Access Funding by the University of Vienna is also acknowledged. We are grateful to two anonymous reviewers for their helpful comments and suggestions.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 2. Lithology of the studied succession at the Wadi Darnah area.
Figure 2. Lithology of the studied succession at the Wadi Darnah area.
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Figure 3. (A) Rock units of the study section; (B,C) gray nummulitic limestone at the lower part; (D,E) yellowish gray nummulitic beds at the middle part; (F,G) light gray nummulitic limestone with large-sized bivalves at the upper part.
Figure 3. (A) Rock units of the study section; (B,C) gray nummulitic limestone at the lower part; (D,E) yellowish gray nummulitic beds at the middle part; (F,G) light gray nummulitic limestone with large-sized bivalves at the upper part.
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Figure 7. Paleobiogeographic distribution of Terebratulina tenuistriata and Orthothyris pectinoides during the Middle Eocene [62] (The map is modified from http://scotese.com).
Figure 7. Paleobiogeographic distribution of Terebratulina tenuistriata and Orthothyris pectinoides during the Middle Eocene [62] (The map is modified from http://scotese.com).
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Abd El-Aziz, S.M.; Abd El-Gaied, I.M.; Al-Hashim, M.H.; Bitner, M.A.; Salama, Y.F.; Heinz, P.; Sayed, M.M. First Report of Middle Eocene Micromorphic Brachiopods from Northeastern Libya: Taxonomy and Paleobiogeography Implications. Diversity 2024, 16, 672. https://doi.org/10.3390/d16110672

AMA Style

Abd El-Aziz SM, Abd El-Gaied IM, Al-Hashim MH, Bitner MA, Salama YF, Heinz P, Sayed MM. First Report of Middle Eocene Micromorphic Brachiopods from Northeastern Libya: Taxonomy and Paleobiogeography Implications. Diversity. 2024; 16(11):672. https://doi.org/10.3390/d16110672

Chicago/Turabian Style

Abd El-Aziz, Sayed M., Ibrahim M. Abd El-Gaied, Mansour H. Al-Hashim, Maria Aleksandra Bitner, Yasser F. Salama, Petra Heinz, and Mostafa M. Sayed. 2024. "First Report of Middle Eocene Micromorphic Brachiopods from Northeastern Libya: Taxonomy and Paleobiogeography Implications" Diversity 16, no. 11: 672. https://doi.org/10.3390/d16110672

APA Style

Abd El-Aziz, S. M., Abd El-Gaied, I. M., Al-Hashim, M. H., Bitner, M. A., Salama, Y. F., Heinz, P., & Sayed, M. M. (2024). First Report of Middle Eocene Micromorphic Brachiopods from Northeastern Libya: Taxonomy and Paleobiogeography Implications. Diversity, 16(11), 672. https://doi.org/10.3390/d16110672

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