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Disponible en ligne sur ScienceDirect www.sciencedirect.com
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
a b s t r a c t
Article history: A new occurrence of the Palaeocarpilius macrochelus (Desmarest, 1822) (Crustacea: Decapoda: Brachyura),
Received 24 September 2019
from the Middle Oligocene rocks of the United Arab Emirates is systematically and paleogeographically
Accepted 8 November 2019
described. The Oligocene Asmari Formation is well exposed on the flanks of Jabal Hafit anticline near Al
Available online xxx
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
Keywords:
the Asmari Formation of Jabal Hafit. The specimen is herein described and assigned to Palaeocarpilius
Oligocene
macrocheles (Desmarest, 1822). The present record complements the information on the temporal and
Brachyura
spatial distribution of decapods at the southern margin of the Tethyan Ocean during the Middle Oligocene Palaeocarpilius
Reefs times.
Paleobiogeography © 2019 Elsevier Masson SAS. All rights reserved.
Tethys
r é s u m é
Mots clés : Une nouvelle occurrence de Palaeocarpilius macrochelus (Desmarest, 1822) (Crustacea : Decapoda :
Oligocène Brachyura), provenant des dépôts de l’Oligocène moyen des Emirats Arabes Unis, est analysée du point de
Brachyoures
vue systématique et paléogéographique. La formation Oligocène Asmari affleure largement sur les flancs
Palaeocarpilius
de l’anticlinal de Jabal Hafit, près de la ville d’Al Ain, et représente l’un des affleurements complexes de
Récifs, Paleobiogéographie
récifs frangeants à la limite ouest des montagnes du nord de l’Oman. Un seul spécimen relativement bien
Téthys
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 dis-
tribution temporelle et spatiale des décapodes dans la marge sud de l’océan Téthysien pendant la période
de l’Oligocène moyen.
© 2019 Elsevier Masson SAS. Tous droits reserv´ es.´
1. Introduction et al., 2011). Hyznˇ y,´ 2016 concluded that the Mediterranean marine
decapod taxa migrated in an easterly direction during the Oligocene
Tertiary fossil decapods from the Middle East have been poorly and/or Miocene, establishing present-day decapod communities in
studied. The decapod crustacean fauna of Egypt attracted a great the Indo-West Pacific. The most common and widely distributed
deal of interest in the second half of the 19th and early 20th cen- Eocene to Miocene crabs belong to the genus Palaeocarpilius
turies (Lorenthey,˝ 1909; Glaessner, 1929; Cuvillier, 1930; Feldmann (A. Milne-Edwards, 1862). Palaeocarpilius is known to have ranged
from the Lutetian (Middle Eocene) to the Lower Miocene of north-
west 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
E-mail address: [email protected]
https://doi.org/10.1016/j.annpal.2019.102383
0753-3969/© 2019 Elsevier Masson SAS. All rights reserved.
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
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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). an early Middle Oligocene age (Hamdan and Bahr, 1992). Mutaredh
Glaessner (1929) listed the Palaeocarpilius simplex Stoliczka, 1871 Member is mainly formed of thickly bedded, algal coralline and
from the Upper Eocene rocks of Egypt. Also, Palaeocarpilius straeleni foraminiferal limestone. The collected and described decapod
Remy and Tessier, 1954 was described from the lower Lutetian of comes from Mutaredh Member, which is approximately 90 m thick
Senegal (Karasawa and Schweitzer, 2006). in the study area (Fig. 1B). Highly diversified and abundant corals
The scope of the present short contribution is to describe a new associated with nummulitid foraminifers, coralline algae, bry-
finding of decapod crustaceans from the Middle Oligocene rocks of ozoan, gastropods and bivalves characterize the Mutaredh Member
the United Arab Emirates and to enhance the knowledge of the (Aly and Gameil, 2000; Gameil, 2001). These faunal communities
paleogeographic distribution of the genus Palaeocarpilius on the growing under conditions of relatively pure carbonate sedimen-
southern margin of the Tethyan Ocean during the late phase of the tation. The coral fauna associated with the recorded crab form
Tertiary Period. 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
2. Geological Setting
poring activity in parts. This coral fauna is represented by hemi-
spherical and laminar poritids belonging to Goniopora and Actinacis.
Al Ain City is located at the margin of the Oman Mountains
The massive corals Astreopora, Alveopora and Thamnastraea, and the
and Jabal Hafit has been involved in a later stage of the moun-
meanderoid forms belonging to the coral genera: Leptoria, Mean-
tain building process (Hunting, 1979). The sedimentary Hawasina
drina, Oulophyllia and Platygyra are characteristic (Aly and Gameil,
and ophiolitic Semail rocks, of the pre-Maastrichtian age form an
2000). Gameil and Aly (2001) studied the systematic aspects and
allochthonous sequence of an overthrust nappes on Oman Moun-
paleoecologic implications of the Asmari Oligocene corals of the
tains and their western margin. The other exposed rocks were laid
United Arab Emirates.
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 3. Systematic Paleontology
well exposed at Al Ain City and were subdivided into different for-
mal and informal rock units by many authors (Fig. 1A). Morphological features, terminology employed, along with the
Oligocene fringing reef complex outcrop along the northern systematic classification used herein follows Schweitzer et al.
part of Jabal Hafit, at the western margin of the Northern Oman (2018). The synonymies are limited to a minimum (first reference,
Mountains. The Oligocene sediments of Jabal Hafit lithologically regional records, important revisions). Biometric measurements
named the Asmari Formation, which subdivided by Hamdan and of carapace length (L1), length from anterior of carapace to posi-
Bahr (1992) into three formal members from the base to the top are: tion of maximum carapace width (L2), carapace width (W), frontal
Zakher, Mutaredh and Muwaiji members. The Mutaredh Member is width (FW), fronto-orbital width (FOW), posterior width (PW) were
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
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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 che-
lipede 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).
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
Fig. 2. Composite line drawing of Palaeocarpilius macrochelus (Desmarest) showing
the position and orientation of taken measurements.
Description. Relatively large sized, triangular front, trans-
Dessin au trait composite de Palaeocarpilius macrochelus (Desmarest) montrant la
position et l’orientation des mesures prises. versely triangular to sub-ovate carapace. It is wider than long;
carapace length being about two-third of width (73.2% of max-
imum width); carapace maximum length is only 82 mm, while
the maximum width is about 112 mm. Carapace is strongly arched
obtained using a Vernier Caliper and are given in mm (Fig. 2). The
anteriorly and less arched, strongly inclined posteriorly. Carapace
collected specimen is housed at the Paleontological Museum of the
is almost smooth, with fine closely spaced dots characterizing the
Geology Department, Cairo University, and stored with the collec-
dorsal surface. Seven strong rounded projections distinguish the
tion number UAE/OLIGO/088/CUGM.
strongly convex anterolateral margin, the three anterior tubercles
Infraorder Brachyura Latreille, 1802
are well-developed more rounded and more prominent, remaining
Section Heterotremata Guinot, 1977
tubercles become smaller, most posterior tubercle being stronger
Superfamily Carpilioidea Ortmann, 1893
and acute. Posterolateral margin is smooth, short and deeply
Family Carpiliidae Ortmann, 1893
concave, one-third the maximum carapace width. Posterolateral
Palaeocarpilius
Genus A. Milne-Edwards, 1862 ◦
margin is composed of two segments, initially at about 82 angle,
Type Species. Cancer macrochelus Desmarest, 1817, by subse- ◦
then becoming more gentler, at about 58 . The fronto-orbital width
quent designation of Glaessner (1929, p. 291).
represents about 62.50% of the maximum carapace width. The
Included Species. Palaeocarpilius aquitanicus A. Milne- Edwards,
orbits hemicircular, entire and rimmed open to the front, with
1862; P. ignotus A. Milne-Edwards, 1862 (finger only); P. inter-
thickened upper orbital margin. The propodus are large, strong,
medius Stubblefield, 1927; P. Laevis Imaizumi, 1939; P. macrochelus
and triangular in shape; the left one is well preserved than the
(Desmarest, 1822); P. mississippiensis (Rathbun, 1935); P. rugifer
right one. Eight projecting spines are well developed on the left
Stoliczka, 1871; P. valrovinensis (De Gregorio, 1895).
cheliped.
Discussion. Feldmann et al., 2011 reported that the examina-
Remarks. P. macrochelus is firstly reported by Desmarest (1822)
tion of the species currently referred to Palaeocarpilius (Schweitzer
based on a specimen (MNHN R03830) from? China, preserved in
et al., 2010) indicated that the range of variation within the genus
the National Museum of Natural History of Paris. The Ratios of
has gotten very broad. Schweitzer et al. (2018) concluded that
measurements (L1/L2, L1/W, F/W, FOW/W), carapace shape and
genus Palaeocarpilius and its allied species have length/width
outline, anterolateral and posterolateral margins, in the present
ratios of 65–78% and nearly straight posterolateral margins. On
work, are closely resemble to data given for Palaeocarpilius A. Milne-
the other hand, Feldmann et al., 2011 restricted Palaeocarpilius to
Edwards, 1862, in the literatures (Schweitzer, 2003; Feldmann
those species with a triangular front, at least seven anterolateral
et al., 2011). The specimen described herein from Jabal Hafit,
spines, and a length to width ratio of about 75%.
UAE corresponds well in carapace morphology to P. macrochelus
(Desmarest, 1822; p. 91, t. 7, ff. 1–2), which figured and described
P.
macrochelus (Desmarest, 1822)
in (Nöetling, 1885; Stubblefield, 1927; Beschin et al., 2006). The
Fig. 3A–D
described specimens from Middle Eocene of British Somaliland by
1822 Cancer macrochelus n. sp. Desmarest, p. 91, taf. VII, figs.
Stubblefield (1946) are ranging in length between 50 to 30 mm,
1–2.
while the present specimen from UAE is about 58 mm length.
1862 Palaeocarpilius macrocheilus (Desmarest) – A. Milne
On the other hand, the upper marginal edge of the hand of the
Edwards, p. 186, t. 1, f. 2, t. 2, fig. 1, t. 3, f. 1
Oligocene specimen from UAE is characterized by eight tuber-
1885 Palaeocarpilius macrocheilus (Desmarest) – Nöetling, p.
cles, where the P. macrochelus described by Stubblefield ( 1946)
487, 489, t. 4, f. 2
has six to eight tubercles. Beschin and De Angeli (2006) described
1886 Palaeocarpilius macrocheilus var. coronatea; Bittner, p. 44,
twenty-one specimens of P. macrochelus (Desmarest) from the
t. 1, f. 1
Upper Eocene (Priabonian) from Italy, their maximum width of
1895 Harpactocarcinus supragigas n. sp. – De Gregorio, p. 13, t.
carapace ranges between 130 to 58 mm, while their maximum
6, ff. 1–3
length of carapace ranges between 110 to 42.5 mm. The Beschin
1929 Palaeocarpilius macrocheilus (Desmarest) – Lorenthey˝ and
and De Angeli (2006) specimens from Italy are characterized by
Beurlen, p. 222 (full synonymy).
eight rounded tubercles on the cheliped upper lateral margin like
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
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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 eight counted for the anterolateral margin Palaeocarpilius rugifer
A. Milne Edwards, 1862 has only seven tubercles. Moreover, P. sim- Stoliczka, 1871, and therefore, the number of tubercles should be
plex (= P. anodon Bittner, 1875) is distinguished by the anterolateral seven.
margins of the carapace and the upper edge of the smooth claws, Age and Distribution. P. macrochelus (Desmarest) is recorded
without lobes (Beschin et al., 2012). Francisco et al. (2010) con- from the Middle Oligocene Asmari Formation of Jabal Hafit, UAE; it
cluded that Stoliczka included the outer orbital spine as one of the 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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the Oligocene of the United Arab Emirates. Annales de Paléontologie (xxxx), https://doi.org/10.1016/j.annpal.2019.102383
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1915; De Angeli and Caporiondo, 2009), Hungary (Lorenthey˝ and Cuvillier, J., 1930. Révision du Nummulitique égyptien. Mémoires de l’Institut
d’Egypte 16, 1–372.
Beurlen, 1929), Egypt (Nöetling, 1885; Lorenthey,˝ 1909) and Soma-
Dainelli, G., 1915. L’Eocene Friulano - Monografi a geologica e paleontologica. Editrici
lia (Stubblefield, 1946). Beschin and De Angeli (2006) mentioned
le “Memorie Geografiche”, Firenze, pp. 1–72.
that many records of P. macrochelus (Desmarest) in literatures are De Angeli, A., 1995. Crostacei dell’Eocene superiore di “Fontanella” di Grancona
doubtful. (Vicenza - Italia settentrionale). Studi e Ricerche - Associazione Amici Museo
Civico “G. Zannato”, Montecchio Maggiore (Vicenza), pp. 7–24.
De Angeli, A., Caporiondo, F., 2009. Crostacei Decapodi del Priaboniano di
Sossano (Monti Berici, Vicenza - Italia Settentrionale). Associazione Amici
4. Conclusions
Museo Civico “G. Zannato”, Montecchio Maggiore (Vicenza), pp. 23–33
(16).
De Gregorio, A., 1895. Description des faunes tertiaires de la Vénétie. Note sur certain
The present work documents the first occurrence and record
Crustacés (Brachioures) Éocéniques (avec un catalogue de tous les Crustacés
of P. macrochelus (Desmarest) from the Arabian Peninsula. The de la Vénétie cités par les auteurs). Annales de Géologie et de Paléontologie,
Palermo 18, 1–22.
described specimen originates from the topmost irregularly bed-
Desmarest, A.G., 1817. Crustacés fossiles. In: Nouveau dictionnaire d’histoire
ded, massive and hard coralline Middle Oligocene Mutaredh
naturelle, appliquée aux arts, à l’agriculture, à l’économie rurale et domestique,
limestones of the eastern flank of Jabal Hafit, UAE. These lime- à la médecine, etc. par une société de naturalistes et d’agriculteurs, 2nd éd. 8.
stones are represented coral reef facies, which is highly diversified Deterville, Paris, pp. 495–519.
Desmarest, A.G., 1822. Histoire naturelle des Crustacés fossiles. Les Crustacés pro-
with gastropods, bivalves, and coral rubble and heads up to 50 cm
prement dits. Levrault, Paris, pp. 67–154.
in diameter with abundant Nummulites spp. The Oligocene reefal
Gameil, M., Aly, M.F., 2001. Paleontological studies on some Oligocene colonial corals
limestones of Jabal Hafit represent shelf margin prograding com- from Gabal Hafit (Al Ain area, UAE). M.E.RC. Ain Shams University, Earth Science
Series 15, 61–78.
plex related to the major Middle Oligocene sea lowstand level. In
Glaessner, M.F., 1929. Crustacea Decapoda. In: Pompeckj, F.J. (Ed.), Fossilium Cata-
general, the genus Palaeocarpilius A. Milne-Edwards, 1862 exhibits
logus, 1: Animalium 41. W. Junk, Berlin (464 p.).
a Tethyan distribution (Fig. 4). The described fauna from the Glennie, K.W., Boeuf, M.G., Hughes-Clarke, M.H.W., Moody-Stuart, M.,
Pilaar, W.F., Reinhardt, B., 1974. Geology of the Oman Mountains. Ver-
Middle Oligocene Mutaredh Member, along with the described
handelingen Koninklijk, The Nederlands (Geologisch Mijnbouwkundig
P. macrochelus (Desmarest) reflecting southern Tethyan faunal
Genootschap no. 31).
affinities and reef front facies with water depth exceeded 10 m Guinot, D., 1977. Propositions pour une nouvelle classification des Crustacés
Décapodes Brachyoures. Comptes rendus hebdomadaires des séances de
depth, high energy, hard substrate and normal saline water sett-
l’Académie des sciences (Paris) 285, 1049–1052.
ings.
Feldmann, R.M., Schweitzer, C.E., Bennett, O., Frant¸escu, O., Resar, N., Trudeau, A.,
2011. Decapod crustaceans from the Eocene of Egypt. Neues Jahrbuch für Geolo-
gie und Paläontologie, Abhandlung 262, 323–353.
Disclosure of interest
Hamdan, A.A., Bahr, S.A., 1992. Lithostratigraphy of the Paleogene Succession of
Northern Jabal Hafit., Al-Ain Area, UAE. M.E.R.C., Ain Shams University. Earth
Science 6, 201–224.
The author declares that he has no competing interest. Hyznˇ y,´ M., 2016. Diversity and distribution patterns of the Oligocene and Miocene
decapod crustaceans (Crustacea: Malacostraca) of the Western and Central
Paratethys. Geologica Carpathica 67 (5), 471–494.
Acknowledgments Hunting, A., 1979. Geology and Geophysics Ltd., Report on Mineral Survey of the
U.A.E., Al-Ain Area., pp. 29 (9).
Imaizumi, R., 1939. Palaeocarpilius laevis sp. nov. from the Tappocho Limestone of
Saipan Mariana Group. Jubilee Publication in the Commemoration of Professor
The editor (Didier Néraudeau; Université de Rennes), the refer-
H. Yabe, M.I. A. Sixtieth Birthday, pp. 221–226 (1).
ees (Sylvian Charbonnier; Muséum National d’Histoire Naturelle,
Karasawa, H., Schweitzer, E., 2006. A new classification of the Xanthoidea sensu
and Denis Audo; Yunnan University) are thanked for careful and lato (Crustacea: Decapoda: Brachyura) based on phylogenetic analysis and
constructive reviews. Many hanks are addressed to Rodney M. Feld- traditional systematics and evaluation of all fossil Xanthoidea sensu lato. Con-
tributions to Zoology 75 (1/2), 23–73.
mann (Kent State University), A. De Angeli (Vicenza, Italy) for
Latreille, P.A., 1802–1803. Histoire naturelle, générale et particulière, des crustacés
their encouragement, support and providing the author with their
et des insectes 3. Dufart, Paris (468 p.).
valuable literatures. I’m greatly indebted to Luc Bulot (Cerege, Aix- Lorenthey,˝ E., 1909. Beiträge zur Kenntnis der Eozänen Dekapodenfauna Ägyptens.
Mathematische und Naturwissenschaftliche Berichte aus Ungarn 24, 106–152
Marseille University and NARG, Manchester University) for revising
(Hungarian version published in 1907).
and the critical reading of the final manuscript for submission. I
Lorenthey,˝ E., Beurlen, K., 1929. Die fossilen Decapoden der Länder der Ungarischen
would like also to thank Prof. Mohamed El Sharkawi (Cairo Uni- Krone. Geologica Hungarica, Series Palaeontologica 3, 1–421.
Milne-Edwards, A., 1862. Monographie des Crustacés de la famille Cancériens.
versity) for his comments and corrections to improve the final
Annales des Sciences Naturelles (Zoologie) 18 (4), 31–85.
manuscript language.
Nöetling, F., 1885. Ueber Crustaceen aus dem Tertiär Ägyptens. Sitzungs-
berichte der Preussischen Akademie der Wissenschaften, Berlin, pp.
References 487–501.
Ortmann, A.E., 1893. Abtheilung: Brachyura (Brachyura genuina Boas), II. Unter-
abtheilung: Cancroidea, 2. Section: Cancrinea, 1. Gruppe: Cyclometopa. Die
Aly, M.F., Gameil, M., 2000. Oligocene reef coral biofacies of Jabal Hafit, Al Ain, United
Decapoden-Krebse des Strassburger Museums, mit besonderer Berücksich-
Arab Emirates. Geology of the Arab World, 5th International Conference, Cairo,
tigung der von Herrn Dr. Döderlein bei Japan und bei den Liu-Kiu-Inseln
Egypt 3, 1497–1508.
gesammelten und zur Zeit im Strassburger Museum aufbewahrten Formen, VII.
Beschin, C., De Angeli, A., 2006. Il genere Palaeocarpilius A. Milne Edwards, 1862
Theil. Zoologische Jahrbücher, Systematik. Geographie und Biologie der Thiere
(Decapoda, Brachyura, Carpiliidae) nel Terziario del Vicentino (Italia setten-
7, 411–495.
trionale). Studi e Ricerche - Associazione Amici Museo Civico “G. Zannato”,
Piccoli, G., Mocellin, L.G., 1962. Studi sulla macrofauna priaboniana di Priabona.
Montecchio Maggiore (Vicenza) 13, 11–23.
Memorie degli Istituti di Geologia e Mineralogia dell’ Universita di Padova 23,
Beschin, C., De Angeli, A., Checchi, A., Mietto, P.G., 2006. Crostacei del Priaboniano
3–120.
di Priabona (Vicenza–Italia settentrionale). Lavori, Società Veneziana di Scienze
Rathbun, M.J., 1935. Fossil Crustacea of the Atlantic and Gulf Coastal Plain. Geological
Naturali 31, 95–112.
Society of America, Pemba Special Papers 2 i–viii, 1–160.
Beschin, C., De Angeli, A., Checchi, A., Zarantonello, G., 2012. Crostacei del Giacimento
Remy, J.M., Tessier, F., 1954. Décapodes nouveaux de la partie ouest du Sénégal.
Eocenico di Grola Presso Spagnago di Cornedo Vicentino (Vicenza, Italia Setten-
Bulletin de la Société Géologique de France 6 (4), 185–191.
trionale) (Decapoda, Stomatopoda, Isopoda). Museo di Archeologia e Scienze
Schweitzer, C.E., Feldmann, R.M., Garassino, A., Karasawa, H., Schweigert, G., 2010.
Naturali “G. Zannato”, Vicenza, Italy (104 p.).
Systematic list of fossil decapod crustacean species. Crustaceana Monographs
Bittner, A., 1886. Neue Brachyuren des Eocäens von Verona. Sitzungsberichte der
10, 1–222 (Leiden [Brill]).
Kaiserlichen Akademie der Wissenschaften in Wien 94, 44–55.
Schweitzer, C.E., Feldmann, R.M., Karasawa, H., 2018. Part R, Revised, Volume 1,
Collins, J.S.H., Morris, S.F., 1973. A new crab from the Middle Eocene of Libya.
Chapter 8T2: Systematic descriptions: Superfamily Carpilioidea. Treatise Online
Palaeontology 16 (2), 283–292.
112, 1–22.
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
G Model
ANNPAL-102383; No. of Pages 6 ARTICLE IN PRESS
6 M.F. Aly / Annales de Paléontologie xxx (xxxx) xxx
Stoliczka, F., 1871. Observations of fossil crabs from Tertiary deposits in Sind and Whittle, L., Al Sharhan, A., 1994. Dolomitization and chertification of the Early Eocene
Kutch. Memoirs of the Geological Survey of India. Palaeontologica Indica 1 (1), Rus Formation in Abu Dhabi, United Arab Emirates. Sedimentary Geology 92
1–16 ([7] [14]). (3–4), 273–328.
Stubblefield, C.J., 1927. Lower Miocene Crustacea from Pemba Island.
Further reading
Reports on the Paleontology of the Zanzibar Protectorate 22 (23),
118–120.
Stubblefield, C.J., 1946. Some decapoden Crustacea from the Middle Eocene of British
Geologypagecom http://www.geologypage.com/wp-content/
Somaliland. The Annals and Magazine of Natural History, Eleventh Series 13 uploads/2014/04/Paleogene.jpg.
(104), 505–519.
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