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New occurrence of the brachyuran crab Palaeocarpilius macrochelus (Desmarest, 1822) from the Oligocene of the United Arab Emirates
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New occurrence of the brachyuran crab Palaeocarpilius macrochelus(Desmarest, 1822) from the Oligocene of the United Arab Emirates Nouvelle occurrence du crabe brachyurien Palaeocarpilius macrochelus (Desmarest, 1822) dans l’Oligocène des Emirats Arabes Unis Mohamed Fouad Aly Department of Geology, Faculty of Science, Cairo University, Cairo, Egypt
a r t i c l e
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Article history: Received 24 September 2019 Accepted 8 November 2019 Available online xxx Keywords: Oligocene Brachyura Palaeocarpilius Reefs Paleobiogeography Tethys
a b s t r a c t A new occurrence of the Palaeocarpilius macrochelus (Desmarest, 1822) (Crustacea: Decapoda: Brachyura), from the Middle Oligocene rocks of the United Arab Emirates is systematically and paleogeographically described. The Oligocene Asmari Formation is well exposed on the flanks of Jabal Hafit anticline near Al Ain City, UAE, and represents one of the very interesting fringing reef complex outcrops at the western margin of the Northern Oman Mountains. A relatively well preserved single specimen was collected from the Asmari Formation of Jabal Hafit. The specimen is herein described and assigned to Palaeocarpilius macrocheles (Desmarest, 1822). The present record complements the information on the temporal and spatial distribution of decapods at the southern margin of the Tethyan Ocean during the Middle Oligocene times. © 2019 Elsevier Masson SAS. All rights reserved.
r é s u m é Mots clés : Oligocène Brachyoures Palaeocarpilius Récifs, Paleobiogéographie Téthys
Une nouvelle occurrence de Palaeocarpilius macrochelus (Desmarest, 1822) (Crustacea : Decapoda : Brachyura), provenant des dépôts de l’Oligocène moyen des Emirats Arabes Unis, est analysée du point de vue systématique et paléogéographique. La formation Oligocène Asmari affleure largement sur les flancs de l’anticlinal de Jabal Hafit, près de la ville d’Al Ain, et représente l’un des affleurements complexes de récifs frangeants à la limite ouest des montagnes du nord de l’Oman. Un seul spécimen relativement bien conservé a été recueilli dans la formation Asmari de Jabal Hafit. Le spécimen est décrit ici et attribué à Palaeocarpilius macrocheles (Desmarest, 1822). Le présent article complète notre connaissance de la distribution temporelle et spatiale des décapodes dans la marge sud de l’océan Téthysien pendant la période de l’Oligocène moyen. ´ ´ es. © 2019 Elsevier Masson SAS. Tous droits reserv
1. Introduction Tertiary fossil decapods from the Middle East have been poorly studied. The decapod crustacean fauna of Egypt attracted a great deal of interest in the second half of the 19th and early 20th cen˝ turies (Lorenthey, 1909; Glaessner, 1929; Cuvillier, 1930; Feldmann
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´ 2016 concluded that the Mediterranean marine et al., 2011). Hyˇzny, decapod taxa migrated in an easterly direction during the Oligocene and/or Miocene, establishing present-day decapod communities in the Indo-West Pacific. The most common and widely distributed Eocene to Miocene crabs belong to the genus Palaeocarpilius (A. Milne-Edwards, 1862). Palaeocarpilius is known to have ranged from the Lutetian (Middle Eocene) to the Lower Miocene of northwest Africa to Spain and from the Paris Basin to Zanzibar (Collins and Morris, 1973; Schweitzer et al., 2018). Stubblefield (1946) reported the Lutetian to the Oligocene Palaeocarpilius macrochelus
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Fig. 1. A. Geologic map for Jabal Hafit, UAE., showing the study area (modified after Whittle and Al Sharhan, 1994). B. Stratigraphic columnar section of the exposed early Middle Oligocene Mutaredh Member, at Jabal Hafit. A. Carte géologique de Jabal Hafit, Émirats Arabes Unis, montrant la zone d’étude (modifiée après Whittle et Al Sharhan 1994). B. Colonne stratigraphique du membre Mutaredh de l’Oligocène moyen ancien affleurant à Jabal Hafi.
(Desmarest) from northeast Africa (Egypt and the Somali Republic). Glaessner (1929) listed the Palaeocarpilius simplex Stoliczka, 1871 from the Upper Eocene rocks of Egypt. Also, Palaeocarpilius straeleni Remy and Tessier, 1954 was described from the lower Lutetian of Senegal (Karasawa and Schweitzer, 2006). The scope of the present short contribution is to describe a new finding of decapod crustaceans from the Middle Oligocene rocks of the United Arab Emirates and to enhance the knowledge of the paleogeographic distribution of the genus Palaeocarpilius on the southern margin of the Tethyan Ocean during the late phase of the Tertiary Period.
2. Geological Setting Al Ain City is located at the margin of the Oman Mountains and Jabal Hafit has been involved in a later stage of the mountain building process (Hunting, 1979). The sedimentary Hawasina and ophiolitic Semail rocks, of the pre-Maastrichtian age form an allochthonous sequence of an overthrust nappes on Oman Mountains and their western margin. The other exposed rocks were laid down after the emplacement of the nappes and are represented by the Maastrichtian to Neogene sediments (Glennie et al., 1974; Hunting, 1979; Aly and Gameil, 2000). The Paleogene rocks were well exposed at Al Ain City and were subdivided into different formal and informal rock units by many authors (Fig. 1A). Oligocene fringing reef complex outcrop along the northern part of Jabal Hafit, at the western margin of the Northern Oman Mountains. The Oligocene sediments of Jabal Hafit lithologically named the Asmari Formation, which subdivided by Hamdan and Bahr (1992) into three formal members from the base to the top are: Zakher, Mutaredh and Muwaiji members. The Mutaredh Member is
an early Middle Oligocene age (Hamdan and Bahr, 1992). Mutaredh Member is mainly formed of thickly bedded, algal coralline and foraminiferal limestone. The collected and described decapod comes from Mutaredh Member, which is approximately 90 m thick in the study area (Fig. 1B). Highly diversified and abundant corals associated with nummulitid foraminifers, coralline algae, bryozoan, gastropods and bivalves characterize the Mutaredh Member (Aly and Gameil, 2000; Gameil, 2001). These faunal communities growing under conditions of relatively pure carbonate sedimentation. The coral fauna associated with the recorded crab form dense-rigid framework, with massive coral colonies in point to point contact. The corals of the crab bed (Fig. 1B) are often encrusted by coralline algae, bryozoans and foraminifers and are affected by poring activity in parts. This coral fauna is represented by hemispherical and laminar poritids belonging to Goniopora and Actinacis. The massive corals Astreopora, Alveopora and Thamnastraea, and the meanderoid forms belonging to the coral genera: Leptoria, Meandrina, Oulophyllia and Platygyra are characteristic (Aly and Gameil, 2000). Gameil and Aly (2001) studied the systematic aspects and paleoecologic implications of the Asmari Oligocene corals of the United Arab Emirates.
3. Systematic Paleontology Morphological features, terminology employed, along with the systematic classification used herein follows Schweitzer et al. (2018). The synonymies are limited to a minimum (first reference, regional records, important revisions). Biometric measurements of carapace length (L1), length from anterior of carapace to position of maximum carapace width (L2), carapace width (W), frontal width (FW), fronto-orbital width (FOW), posterior width (PW) were
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1946 P. macrochelus (Desmarest) – Stubblefield, p. 513, t. 8, ff. 2–6 1962 Palaeocarpilius macrocheilus (Desmarest) – Piccoli and Mocellin, p. 38, 48, 78 1995 P. macrochelus (Desmarest) – De Angeli, p. 16 2006 P. macrochelus (Desmarest) – Beschin et al., p. 107, t. 3, ff. 3, 4a, b 2006 P. macrochelus (Desmarest) – Beschin and De Angeli, p. 13, fig. 2, t. 7, ff. 1, 2a, b (full synonymy). 2009 P. macrochelus (Desmarest) – De Angeli and Caporiondo, p. 30 Material. One imperfectly preserved carapace (ventral part of the carapace is not preserved) UAE/OLIGO/088/CUGM, the left chelipede partly preserved, was collected from the early Middle Eocene carbonates of the UAE, in association with Nummulites spp., corals, bryozoans and mollusks. Measurements (in mm).
Fig. 2. Composite line drawing of Palaeocarpilius macrochelus (Desmarest) showing the position and orientation of taken measurements. Dessin au trait composite de Palaeocarpilius macrochelus (Desmarest) montrant la position et l’orientation des mesures prises.
obtained using a Vernier Caliper and are given in mm (Fig. 2). The collected specimen is housed at the Paleontological Museum of the Geology Department, Cairo University, and stored with the collection number UAE/OLIGO/088/CUGM. Infraorder Brachyura Latreille, 1802 Section Heterotremata Guinot, 1977 Superfamily Carpilioidea Ortmann, 1893 Family Carpiliidae Ortmann, 1893 Genus Palaeocarpilius A. Milne-Edwards, 1862 Type Species. Cancer macrochelus Desmarest, 1817, by subsequent designation of Glaessner (1929, p. 291). Included Species. Palaeocarpilius aquitanicus A. Milne- Edwards, 1862; P. ignotus A. Milne-Edwards, 1862 (finger only); P. intermedius Stubblefield, 1927; P. Laevis Imaizumi, 1939; P. macrochelus (Desmarest, 1822); P. mississippiensis (Rathbun, 1935); P. rugifer Stoliczka, 1871; P. valrovinensis (De Gregorio, 1895). Discussion. Feldmann et al., 2011 reported that the examination of the species currently referred to Palaeocarpilius (Schweitzer et al., 2010) indicated that the range of variation within the genus has gotten very broad. Schweitzer et al. (2018) concluded that genus Palaeocarpilius and its allied species have length/width ratios of 65–78% and nearly straight posterolateral margins. On the other hand, Feldmann et al., 2011 restricted Palaeocarpilius to those species with a triangular front, at least seven anterolateral spines, and a length to width ratio of about 75%. P. macrochelus (Desmarest, 1822) Fig. 3A–D 1822 Cancer macrochelus n. sp. Desmarest, p. 91, taf. VII, figs. 1–2. 1862 Palaeocarpilius macrocheilus (Desmarest) – A. Milne Edwards, p. 186, t. 1, f. 2, t. 2, fig. 1, t. 3, f. 1 1885 Palaeocarpilius macrocheilus (Desmarest) – Nöetling, p. 487, 489, t. 4, f. 2 1886 Palaeocarpilius macrocheilus var. coronatea; Bittner, p. 44, t. 1, f. 1 1895 Harpactocarcinus supragigas n. sp. – De Gregorio, p. 13, t. 6, ff. 1–3 ˝ 1929 Palaeocarpilius macrocheilus (Desmarest) – Lorenthey and Beurlen, p. 222 (full synonymy).
L
L2
L2/L1
W
L1/W
FW
FW/W
FOW
FOW/W
PW
82
58
70.73
112
73.2
50
44.64
70
62.50
55
Description. Relatively large sized, triangular front, transversely triangular to sub-ovate carapace. It is wider than long; carapace length being about two-third of width (73.2% of maximum width); carapace maximum length is only 82 mm, while the maximum width is about 112 mm. Carapace is strongly arched anteriorly and less arched, strongly inclined posteriorly. Carapace is almost smooth, with fine closely spaced dots characterizing the dorsal surface. Seven strong rounded projections distinguish the strongly convex anterolateral margin, the three anterior tubercles are well-developed more rounded and more prominent, remaining tubercles become smaller, most posterior tubercle being stronger and acute. Posterolateral margin is smooth, short and deeply concave, one-third the maximum carapace width. Posterolateral margin is composed of two segments, initially at about 82◦ angle, then becoming more gentler, at about 58◦ . The fronto-orbital width represents about 62.50% of the maximum carapace width. The orbits hemicircular, entire and rimmed open to the front, with thickened upper orbital margin. The propodus are large, strong, and triangular in shape; the left one is well preserved than the right one. Eight projecting spines are well developed on the left cheliped. Remarks. P. macrochelus is firstly reported by Desmarest (1822) based on a specimen (MNHN R03830) from? China, preserved in the National Museum of Natural History of Paris. The Ratios of measurements (L1/L2, L1/W, F/W, FOW/W), carapace shape and outline, anterolateral and posterolateral margins, in the present work, are closely resemble to data given for Palaeocarpilius A. MilneEdwards, 1862, in the literatures (Schweitzer, 2003; Feldmann et al., 2011). The specimen described herein from Jabal Hafit, UAE corresponds well in carapace morphology to P. macrochelus (Desmarest, 1822; p. 91, t. 7, ff. 1–2), which figured and described in (Nöetling, 1885; Stubblefield, 1927; Beschin et al., 2006). The described specimens from Middle Eocene of British Somaliland by Stubblefield (1946) are ranging in length between 50 to 30 mm, while the present specimen from UAE is about 58 mm length. On the other hand, the upper marginal edge of the hand of the Oligocene specimen from UAE is characterized by eight tubercles, where the P. macrochelus described by Stubblefield ( 1946) has six to eight tubercles. Beschin and De Angeli (2006) described twenty-one specimens of P. macrochelus (Desmarest) from the Upper Eocene (Priabonian) from Italy, their maximum width of carapace ranges between 130 to 58 mm, while their maximum length of carapace ranges between 110 to 42.5 mm. The Beschin and De Angeli (2006) specimens from Italy are characterized by eight rounded tubercles on the cheliped upper lateral margin like
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Fig. 3. Palaeocarpilius macrochelus (Desmarest, 1822) from the Asmari Formation, UAE, Middle Oligocene, UAE/OLIGO/088/CUGM. A. Anterior view. B. Dorsal view. C. Left lateral view. D. Right lateral view. Scal bars: 1.3 cm. Palaeocarpilius macrochelus (Desmarest, 1822) de la formation d’Asmari, Émirats Arabes Unis, Oligocène moyen, UAE/OLIGO/088/CUGM. A. Vue antérieure. B. Vue dorsale. C. Vue latérale gauche. D. Vue latérale droite. Barres d’échelle : 1,3 cm.
the described specimen herein, where Palaeocarpilius aquitanicus A. Milne Edwards, 1862 has only seven tubercles. Moreover, P. simplex (= P. anodon Bittner, 1875) is distinguished by the anterolateral margins of the carapace and the upper edge of the smooth claws, without lobes (Beschin et al., 2012). Francisco et al. (2010) concluded that Stoliczka included the outer orbital spine as one of the
eight counted for the anterolateral margin Palaeocarpilius rugifer Stoliczka, 1871, and therefore, the number of tubercles should be seven. Age and Distribution. P. macrochelus (Desmarest) is recorded from the Middle Oligocene Asmari Formation of Jabal Hafit, UAE; it is also recorded from the Eocene-Oligocene rocks of Italy (Dainelli,
Fig. 4. Palaeogeographic distribution of Palaeocarpilius macrochelus (Desmarest, 1822) (map modified after: http://www.geologypage.com/wp-content/uploads/2014/04/ Paleogene.jpg). Répartition paléogéographique de Palaeocarpilius macrochelus (Desmarest, 1822) (carte modifiée d’après : http://www.geologypage.com/wp-content/uploads/2014/04/Paleogene. jpg).
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˝ 1915; De Angeli and Caporiondo, 2009), Hungary (Lorenthey and ˝ 1909) and SomaBeurlen, 1929), Egypt (Nöetling, 1885; Lorenthey, lia (Stubblefield, 1946). Beschin and De Angeli (2006) mentioned that many records of P. macrochelus (Desmarest) in literatures are doubtful. 4. Conclusions The present work documents the first occurrence and record of P. macrochelus (Desmarest) from the Arabian Peninsula. The described specimen originates from the topmost irregularly bedded, massive and hard coralline Middle Oligocene Mutaredh limestones of the eastern flank of Jabal Hafit, UAE. These limestones are represented coral reef facies, which is highly diversified with gastropods, bivalves, and coral rubble and heads up to 50 cm in diameter with abundant Nummulites spp. The Oligocene reefal limestones of Jabal Hafit represent shelf margin prograding complex related to the major Middle Oligocene sea lowstand level. In general, the genus Palaeocarpilius A. Milne-Edwards, 1862 exhibits a Tethyan distribution (Fig. 4). The described fauna from the Middle Oligocene Mutaredh Member, along with the described P. macrochelus (Desmarest) reflecting southern Tethyan faunal affinities and reef front facies with water depth exceeded 10 m depth, high energy, hard substrate and normal saline water settings. Disclosure of interest The author declares that he has no competing interest. Acknowledgments The editor (Didier Néraudeau; Université de Rennes), the referees (Sylvian Charbonnier; Muséum National d’Histoire Naturelle, and Denis Audo; Yunnan University) are thanked for careful and constructive reviews. Many hanks are addressed to Rodney M. Feldmann (Kent State University), A. De Angeli (Vicenza, Italy) for their encouragement, support and providing the author with their valuable literatures. I’m greatly indebted to Luc Bulot (Cerege, AixMarseille University and NARG, Manchester University) for revising and the critical reading of the final manuscript for submission. I would like also to thank Prof. Mohamed El Sharkawi (Cairo University) for his comments and corrections to improve the final manuscript language. References Aly, M.F., Gameil, M., 2000. Oligocene reef coral biofacies of Jabal Hafit, Al Ain, United Arab Emirates. Geology of the Arab World, 5th International Conference, Cairo, Egypt 3, 1497–1508. Beschin, C., De Angeli, A., 2006. Il genere Palaeocarpilius A. Milne Edwards, 1862 (Decapoda, Brachyura, Carpiliidae) nel Terziario del Vicentino (Italia settentrionale). Studi e Ricerche - Associazione Amici Museo Civico “G. Zannato”, Montecchio Maggiore (Vicenza) 13, 11–23. Beschin, C., De Angeli, A., Checchi, A., Mietto, P.G., 2006. Crostacei del Priaboniano di Priabona (Vicenza–Italia settentrionale). Lavori, Società Veneziana di Scienze Naturali 31, 95–112. Beschin, C., De Angeli, A., Checchi, A., Zarantonello, G., 2012. Crostacei del Giacimento Eocenico di Grola Presso Spagnago di Cornedo Vicentino (Vicenza, Italia Settentrionale) (Decapoda, Stomatopoda, Isopoda). Museo di Archeologia e Scienze Naturali “G. Zannato”, Vicenza, Italy (104 p.). Bittner, A., 1886. Neue Brachyuren des Eocäens von Verona. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften in Wien 94, 44–55. Collins, J.S.H., Morris, S.F., 1973. A new crab from the Middle Eocene of Libya. Palaeontology 16 (2), 283–292.
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Further reading Geologypagecom http://www.geologypage.com/wp-content/ uploads/2014/04/Paleogene.jpg.
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Original article
New occurrence of the brachyuran crab Palaeocarpilius macrochelus(Desmarest, 1822) from the Oligocene of the United Arab Emirates Nouvelle occurrence du crabe brachyurien Palaeocarpilius macrochelus (Desmarest, 1822) dans l’Oligocène des Emirats Arabes Unis Mohamed Fouad Aly Department of Geology, Faculty of Science, Cairo University, Cairo, Egypt
a r t i c l e
i n f o
Article history: Received 24 September 2019 Accepted 8 November 2019 Available online xxx Keywords: Oligocene Brachyura Palaeocarpilius Reefs Paleobiogeography Tethys
a b s t r a c t A new occurrence of the Palaeocarpilius macrochelus (Desmarest, 1822) (Crustacea: Decapoda: Brachyura), from the Middle Oligocene rocks of the United Arab Emirates is systematically and paleogeographically described. The Oligocene Asmari Formation is well exposed on the flanks of Jabal Hafit anticline near Al Ain City, UAE, and represents one of the very interesting fringing reef complex outcrops at the western margin of the Northern Oman Mountains. A relatively well preserved single specimen was collected from the Asmari Formation of Jabal Hafit. The specimen is herein described and assigned to Palaeocarpilius macrocheles (Desmarest, 1822). The present record complements the information on the temporal and spatial distribution of decapods at the southern margin of the Tethyan Ocean during the Middle Oligocene times. © 2019 Elsevier Masson SAS. All rights reserved.
r é s u m é Mots clés : Oligocène Brachyoures Palaeocarpilius Récifs, Paleobiogéographie Téthys
Une nouvelle occurrence de Palaeocarpilius macrochelus (Desmarest, 1822) (Crustacea : Decapoda : Brachyura), provenant des dépôts de l’Oligocène moyen des Emirats Arabes Unis, est analysée du point de vue systématique et paléogéographique. La formation Oligocène Asmari affleure largement sur les flancs de l’anticlinal de Jabal Hafit, près de la ville d’Al Ain, et représente l’un des affleurements complexes de récifs frangeants à la limite ouest des montagnes du nord de l’Oman. Un seul spécimen relativement bien conservé a été recueilli dans la formation Asmari de Jabal Hafit. Le spécimen est décrit ici et attribué à Palaeocarpilius macrocheles (Desmarest, 1822). Le présent article complète notre connaissance de la distribution temporelle et spatiale des décapodes dans la marge sud de l’océan Téthysien pendant la période de l’Oligocène moyen. ´ ´ es. © 2019 Elsevier Masson SAS. Tous droits reserv
1. Introduction Tertiary fossil decapods from the Middle East have been poorly studied. The decapod crustacean fauna of Egypt attracted a great deal of interest in the second half of the 19th and early 20th cen˝ turies (Lorenthey, 1909; Glaessner, 1929; Cuvillier, 1930; Feldmann
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´ 2016 concluded that the Mediterranean marine et al., 2011). Hyˇzny, decapod taxa migrated in an easterly direction during the Oligocene and/or Miocene, establishing present-day decapod communities in the Indo-West Pacific. The most common and widely distributed Eocene to Miocene crabs belong to the genus Palaeocarpilius (A. Milne-Edwards, 1862). Palaeocarpilius is known to have ranged from the Lutetian (Middle Eocene) to the Lower Miocene of northwest Africa to Spain and from the Paris Basin to Zanzibar (Collins and Morris, 1973; Schweitzer et al., 2018). Stubblefield (1946) reported the Lutetian to the Oligocene Palaeocarpilius macrochelus
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Fig. 1. A. Geologic map for Jabal Hafit, UAE., showing the study area (modified after Whittle and Al Sharhan, 1994). B. Stratigraphic columnar section of the exposed early Middle Oligocene Mutaredh Member, at Jabal Hafit. A. Carte géologique de Jabal Hafit, Émirats Arabes Unis, montrant la zone d’étude (modifiée après Whittle et Al Sharhan 1994). B. Colonne stratigraphique du membre Mutaredh de l’Oligocène moyen ancien affleurant à Jabal Hafi.
(Desmarest) from northeast Africa (Egypt and the Somali Republic). Glaessner (1929) listed the Palaeocarpilius simplex Stoliczka, 1871 from the Upper Eocene rocks of Egypt. Also, Palaeocarpilius straeleni Remy and Tessier, 1954 was described from the lower Lutetian of Senegal (Karasawa and Schweitzer, 2006). The scope of the present short contribution is to describe a new finding of decapod crustaceans from the Middle Oligocene rocks of the United Arab Emirates and to enhance the knowledge of the paleogeographic distribution of the genus Palaeocarpilius on the southern margin of the Tethyan Ocean during the late phase of the Tertiary Period.
2. Geological Setting Al Ain City is located at the margin of the Oman Mountains and Jabal Hafit has been involved in a later stage of the mountain building process (Hunting, 1979). The sedimentary Hawasina and ophiolitic Semail rocks, of the pre-Maastrichtian age form an allochthonous sequence of an overthrust nappes on Oman Mountains and their western margin. The other exposed rocks were laid down after the emplacement of the nappes and are represented by the Maastrichtian to Neogene sediments (Glennie et al., 1974; Hunting, 1979; Aly and Gameil, 2000). The Paleogene rocks were well exposed at Al Ain City and were subdivided into different formal and informal rock units by many authors (Fig. 1A). Oligocene fringing reef complex outcrop along the northern part of Jabal Hafit, at the western margin of the Northern Oman Mountains. The Oligocene sediments of Jabal Hafit lithologically named the Asmari Formation, which subdivided by Hamdan and Bahr (1992) into three formal members from the base to the top are: Zakher, Mutaredh and Muwaiji members. The Mutaredh Member is
an early Middle Oligocene age (Hamdan and Bahr, 1992). Mutaredh Member is mainly formed of thickly bedded, algal coralline and foraminiferal limestone. The collected and described decapod comes from Mutaredh Member, which is approximately 90 m thick in the study area (Fig. 1B). Highly diversified and abundant corals associated with nummulitid foraminifers, coralline algae, bryozoan, gastropods and bivalves characterize the Mutaredh Member (Aly and Gameil, 2000; Gameil, 2001). These faunal communities growing under conditions of relatively pure carbonate sedimentation. The coral fauna associated with the recorded crab form dense-rigid framework, with massive coral colonies in point to point contact. The corals of the crab bed (Fig. 1B) are often encrusted by coralline algae, bryozoans and foraminifers and are affected by poring activity in parts. This coral fauna is represented by hemispherical and laminar poritids belonging to Goniopora and Actinacis. The massive corals Astreopora, Alveopora and Thamnastraea, and the meanderoid forms belonging to the coral genera: Leptoria, Meandrina, Oulophyllia and Platygyra are characteristic (Aly and Gameil, 2000). Gameil and Aly (2001) studied the systematic aspects and paleoecologic implications of the Asmari Oligocene corals of the United Arab Emirates.
3. Systematic Paleontology Morphological features, terminology employed, along with the systematic classification used herein follows Schweitzer et al. (2018). The synonymies are limited to a minimum (first reference, regional records, important revisions). Biometric measurements of carapace length (L1), length from anterior of carapace to position of maximum carapace width (L2), carapace width (W), frontal width (FW), fronto-orbital width (FOW), posterior width (PW) were
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1946 P. macrochelus (Desmarest) – Stubblefield, p. 513, t. 8, ff. 2–6 1962 Palaeocarpilius macrocheilus (Desmarest) – Piccoli and Mocellin, p. 38, 48, 78 1995 P. macrochelus (Desmarest) – De Angeli, p. 16 2006 P. macrochelus (Desmarest) – Beschin et al., p. 107, t. 3, ff. 3, 4a, b 2006 P. macrochelus (Desmarest) – Beschin and De Angeli, p. 13, fig. 2, t. 7, ff. 1, 2a, b (full synonymy). 2009 P. macrochelus (Desmarest) – De Angeli and Caporiondo, p. 30 Material. One imperfectly preserved carapace (ventral part of the carapace is not preserved) UAE/OLIGO/088/CUGM, the left chelipede partly preserved, was collected from the early Middle Eocene carbonates of the UAE, in association with Nummulites spp., corals, bryozoans and mollusks. Measurements (in mm).
Fig. 2. Composite line drawing of Palaeocarpilius macrochelus (Desmarest) showing the position and orientation of taken measurements. Dessin au trait composite de Palaeocarpilius macrochelus (Desmarest) montrant la position et l’orientation des mesures prises.
obtained using a Vernier Caliper and are given in mm (Fig. 2). The collected specimen is housed at the Paleontological Museum of the Geology Department, Cairo University, and stored with the collection number UAE/OLIGO/088/CUGM. Infraorder Brachyura Latreille, 1802 Section Heterotremata Guinot, 1977 Superfamily Carpilioidea Ortmann, 1893 Family Carpiliidae Ortmann, 1893 Genus Palaeocarpilius A. Milne-Edwards, 1862 Type Species. Cancer macrochelus Desmarest, 1817, by subsequent designation of Glaessner (1929, p. 291). Included Species. Palaeocarpilius aquitanicus A. Milne- Edwards, 1862; P. ignotus A. Milne-Edwards, 1862 (finger only); P. intermedius Stubblefield, 1927; P. Laevis Imaizumi, 1939; P. macrochelus (Desmarest, 1822); P. mississippiensis (Rathbun, 1935); P. rugifer Stoliczka, 1871; P. valrovinensis (De Gregorio, 1895). Discussion. Feldmann et al., 2011 reported that the examination of the species currently referred to Palaeocarpilius (Schweitzer et al., 2010) indicated that the range of variation within the genus has gotten very broad. Schweitzer et al. (2018) concluded that genus Palaeocarpilius and its allied species have length/width ratios of 65–78% and nearly straight posterolateral margins. On the other hand, Feldmann et al., 2011 restricted Palaeocarpilius to those species with a triangular front, at least seven anterolateral spines, and a length to width ratio of about 75%. P. macrochelus (Desmarest, 1822) Fig. 3A–D 1822 Cancer macrochelus n. sp. Desmarest, p. 91, taf. VII, figs. 1–2. 1862 Palaeocarpilius macrocheilus (Desmarest) – A. Milne Edwards, p. 186, t. 1, f. 2, t. 2, fig. 1, t. 3, f. 1 1885 Palaeocarpilius macrocheilus (Desmarest) – Nöetling, p. 487, 489, t. 4, f. 2 1886 Palaeocarpilius macrocheilus var. coronatea; Bittner, p. 44, t. 1, f. 1 1895 Harpactocarcinus supragigas n. sp. – De Gregorio, p. 13, t. 6, ff. 1–3 ˝ 1929 Palaeocarpilius macrocheilus (Desmarest) – Lorenthey and Beurlen, p. 222 (full synonymy).
L
L2
L2/L1
W
L1/W
FW
FW/W
FOW
FOW/W
PW
82
58
70.73
112
73.2
50
44.64
70
62.50
55
Description. Relatively large sized, triangular front, transversely triangular to sub-ovate carapace. It is wider than long; carapace length being about two-third of width (73.2% of maximum width); carapace maximum length is only 82 mm, while the maximum width is about 112 mm. Carapace is strongly arched anteriorly and less arched, strongly inclined posteriorly. Carapace is almost smooth, with fine closely spaced dots characterizing the dorsal surface. Seven strong rounded projections distinguish the strongly convex anterolateral margin, the three anterior tubercles are well-developed more rounded and more prominent, remaining tubercles become smaller, most posterior tubercle being stronger and acute. Posterolateral margin is smooth, short and deeply concave, one-third the maximum carapace width. Posterolateral margin is composed of two segments, initially at about 82◦ angle, then becoming more gentler, at about 58◦ . The fronto-orbital width represents about 62.50% of the maximum carapace width. The orbits hemicircular, entire and rimmed open to the front, with thickened upper orbital margin. The propodus are large, strong, and triangular in shape; the left one is well preserved than the right one. Eight projecting spines are well developed on the left cheliped. Remarks. P. macrochelus is firstly reported by Desmarest (1822) based on a specimen (MNHN R03830) from? China, preserved in the National Museum of Natural History of Paris. The Ratios of measurements (L1/L2, L1/W, F/W, FOW/W), carapace shape and outline, anterolateral and posterolateral margins, in the present work, are closely resemble to data given for Palaeocarpilius A. MilneEdwards, 1862, in the literatures (Schweitzer, 2003; Feldmann et al., 2011). The specimen described herein from Jabal Hafit, UAE corresponds well in carapace morphology to P. macrochelus (Desmarest, 1822; p. 91, t. 7, ff. 1–2), which figured and described in (Nöetling, 1885; Stubblefield, 1927; Beschin et al., 2006). The described specimens from Middle Eocene of British Somaliland by Stubblefield (1946) are ranging in length between 50 to 30 mm, while the present specimen from UAE is about 58 mm length. On the other hand, the upper marginal edge of the hand of the Oligocene specimen from UAE is characterized by eight tubercles, where the P. macrochelus described by Stubblefield ( 1946) has six to eight tubercles. Beschin and De Angeli (2006) described twenty-one specimens of P. macrochelus (Desmarest) from the Upper Eocene (Priabonian) from Italy, their maximum width of carapace ranges between 130 to 58 mm, while their maximum length of carapace ranges between 110 to 42.5 mm. The Beschin and De Angeli (2006) specimens from Italy are characterized by eight rounded tubercles on the cheliped upper lateral margin like
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Fig. 3. Palaeocarpilius macrochelus (Desmarest, 1822) from the Asmari Formation, UAE, Middle Oligocene, UAE/OLIGO/088/CUGM. A. Anterior view. B. Dorsal view. C. Left lateral view. D. Right lateral view. Scal bars: 1.3 cm. Palaeocarpilius macrochelus (Desmarest, 1822) de la formation d’Asmari, Émirats Arabes Unis, Oligocène moyen, UAE/OLIGO/088/CUGM. A. Vue antérieure. B. Vue dorsale. C. Vue latérale gauche. D. Vue latérale droite. Barres d’échelle : 1,3 cm.
the described specimen herein, where Palaeocarpilius aquitanicus A. Milne Edwards, 1862 has only seven tubercles. Moreover, P. simplex (= P. anodon Bittner, 1875) is distinguished by the anterolateral margins of the carapace and the upper edge of the smooth claws, without lobes (Beschin et al., 2012). Francisco et al. (2010) concluded that Stoliczka included the outer orbital spine as one of the
eight counted for the anterolateral margin Palaeocarpilius rugifer Stoliczka, 1871, and therefore, the number of tubercles should be seven. Age and Distribution. P. macrochelus (Desmarest) is recorded from the Middle Oligocene Asmari Formation of Jabal Hafit, UAE; it is also recorded from the Eocene-Oligocene rocks of Italy (Dainelli,
Fig. 4. Palaeogeographic distribution of Palaeocarpilius macrochelus (Desmarest, 1822) (map modified after: http://www.geologypage.com/wp-content/uploads/2014/04/ Paleogene.jpg). Répartition paléogéographique de Palaeocarpilius macrochelus (Desmarest, 1822) (carte modifiée d’après : http://www.geologypage.com/wp-content/uploads/2014/04/Paleogene. jpg).
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˝ 1915; De Angeli and Caporiondo, 2009), Hungary (Lorenthey and ˝ 1909) and SomaBeurlen, 1929), Egypt (Nöetling, 1885; Lorenthey, lia (Stubblefield, 1946). Beschin and De Angeli (2006) mentioned that many records of P. macrochelus (Desmarest) in literatures are doubtful. 4. Conclusions The present work documents the first occurrence and record of P. macrochelus (Desmarest) from the Arabian Peninsula. The described specimen originates from the topmost irregularly bedded, massive and hard coralline Middle Oligocene Mutaredh limestones of the eastern flank of Jabal Hafit, UAE. These limestones are represented coral reef facies, which is highly diversified with gastropods, bivalves, and coral rubble and heads up to 50 cm in diameter with abundant Nummulites spp. The Oligocene reefal limestones of Jabal Hafit represent shelf margin prograding complex related to the major Middle Oligocene sea lowstand level. In general, the genus Palaeocarpilius A. Milne-Edwards, 1862 exhibits a Tethyan distribution (Fig. 4). The described fauna from the Middle Oligocene Mutaredh Member, along with the described P. macrochelus (Desmarest) reflecting southern Tethyan faunal affinities and reef front facies with water depth exceeded 10 m depth, high energy, hard substrate and normal saline water settings. Disclosure of interest The author declares that he has no competing interest. Acknowledgments The editor (Didier Néraudeau; Université de Rennes), the referees (Sylvian Charbonnier; Muséum National d’Histoire Naturelle, and Denis Audo; Yunnan University) are thanked for careful and constructive reviews. Many hanks are addressed to Rodney M. Feldmann (Kent State University), A. De Angeli (Vicenza, Italy) for their encouragement, support and providing the author with their valuable literatures. I’m greatly indebted to Luc Bulot (Cerege, AixMarseille University and NARG, Manchester University) for revising and the critical reading of the final manuscript for submission. I would like also to thank Prof. Mohamed El Sharkawi (Cairo University) for his comments and corrections to improve the final manuscript language. References Aly, M.F., Gameil, M., 2000. Oligocene reef coral biofacies of Jabal Hafit, Al Ain, United Arab Emirates. Geology of the Arab World, 5th International Conference, Cairo, Egypt 3, 1497–1508. Beschin, C., De Angeli, A., 2006. Il genere Palaeocarpilius A. Milne Edwards, 1862 (Decapoda, Brachyura, Carpiliidae) nel Terziario del Vicentino (Italia settentrionale). Studi e Ricerche - Associazione Amici Museo Civico “G. Zannato”, Montecchio Maggiore (Vicenza) 13, 11–23. Beschin, C., De Angeli, A., Checchi, A., Mietto, P.G., 2006. Crostacei del Priaboniano di Priabona (Vicenza–Italia settentrionale). Lavori, Società Veneziana di Scienze Naturali 31, 95–112. Beschin, C., De Angeli, A., Checchi, A., Zarantonello, G., 2012. Crostacei del Giacimento Eocenico di Grola Presso Spagnago di Cornedo Vicentino (Vicenza, Italia Settentrionale) (Decapoda, Stomatopoda, Isopoda). Museo di Archeologia e Scienze Naturali “G. Zannato”, Vicenza, Italy (104 p.). Bittner, A., 1886. Neue Brachyuren des Eocäens von Verona. Sitzungsberichte der Kaiserlichen Akademie der Wissenschaften in Wien 94, 44–55. Collins, J.S.H., Morris, S.F., 1973. A new crab from the Middle Eocene of Libya. Palaeontology 16 (2), 283–292.
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Whittle, L., Al Sharhan, A., 1994. Dolomitization and chertification of the Early Eocene Rus Formation in Abu Dhabi, United Arab Emirates. Sedimentary Geology 92 (3–4), 273–328.
Further reading Geologypagecom http://www.geologypage.com/wp-content/ uploads/2014/04/Paleogene.jpg.
Please cite this article in press as: M.F. Aly, New occurrence of the brachyuran crab Palaeocarpilius macrochelus (Desmarest, 1822) from the Oligocene of the United Arab Emirates. Annales de Paléontologie (xxxx), https://doi.org/10.1016/j.annpal.2019.102383