During the latest Cretaceous, distinct dinosaur faunas were found in Laurasia and Gondwana. Tyrannosaurids, hadrosaurids, and ceratopsians dominated in North America and Asia, while abelisaurids and titanosaurids dominated in South America, India, and Madagascar. Little is known about dinosaur faunas from the latest Cretaceous of Africa, however. Here, a new abelisaurid theropod, Chenanisaurus barbaricus, is described from the upper Maastrichtian phosphates of the Ouled Abdoun Basin in Morocco, North Africa on the basis of a partial dentary and isolated teeth. Chenanisaurus is both one of the largest abelisaurids, and one of the youngest known African dinosaurs. Along with previously reported titanosaurid remains, Chenanisaurus documents the persistence of a classic Gondwanan abelisaurid-titanosaurid fauna in mainland Africa until just prior to the end-Cretaceous mass extinction. The animal is unusual both in terms of its large size and the unusually short and robust jaw. Although it resembles South American carnotaurines in having a deep, bowed mandible, phylogenetic analysis suggests that Chenanisaurus may represent a lineage of abelisaurids that is distinct from those previously described from the latest Cretaceous of South America, Indo-Madagascar, and Europe, consistent with the hypothesis that the fragmentation of Gondwana led to the evolution of endemic dinosaur faunas during the Late Cretaceous.
Abelisauridae Bonaparte and Novas (Bonaparte and Novas, 1985)
Chenanisaurus barbaricus new genus and species
Etymology. The genus name refers to the Sidi Chennane phosphate mines, the type locality, and Greek ‘sauros, ‘lizard’. The species name is derived from the Greek barbaros, “savage” or “barbaric” and also refers to Barbary, a name historically used for Morocco and the northwest coast of Africa.
Diagnosis. Large abelisaurid characterized by a proportionately deep mandible, dentary bowed in lateral view, lateral groove and associated foramina located high on dentary; anterodorsal margin of dentary downturned in lateral view; massive symphysis, with anterior margin of symphysis vertical in lateral view; tip of jaws broad, with dentaries meeting at an obtuse angle.
Holotype. OCP DEK-GE 772, anterior end of the left dentary ( Fig. 3 ; Fig. 4).
Horizon and Locality. The type specimen is reported as coming from the upper Maastrichtian (Couche III) of Sidi Chennane; the pale white color of the bone and the color of the associated matrix are consistent with the preservation seen in other material recovered from this locality. The isolated premaxillary teeth come from the upper Maastrichtian (Couche III) of Sidi Daoui. The maxillary tooth is of unknown provenance but likely comes from either Sidi Chennane or Sidi Daoui. All specimens come from the Ouled Abdoun Basin, Morocco North Africa (Fig. 1).
Fig. 9. Chenanisaurusbarbaricus shown to scale with the Couche III titanosaur.
Nicholas R. Longrich, Xabier Pereda-Suberbiola, Nour-Eddine Jalil, Fatima Khaldoune and Essaid Jourani. 2017. An Abelisaurid from the latest Cretaceous (late Maastrichtian) of Morocco, North Africa. Cretaceous Research. In Press. DOI: 10.1016/j.cretres.2017.03.021
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ملاحظة هامة: كل عمليات تنزيل، رفع، وتعديل المواضيع الجاهزة تتم بطريقة آلية، ونعتذر عن اي موضوع مخالف او مخل بالحياء مرفوع بالمدونات الجاهزة بآلاف المواضيع والمشاركات، ولكم ان تقوموا بحذف هذه المواضيع والمشاركات والطريقة بسيطة وسهلة. ــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــسلامـ.
Abelisaurid theropods are well-know from the Cretaceous of several parts of the Southern Hemisphere, including South America, Madagascar, and Africa, but also in India and Europe. Abelisaurids are high-diverse among other theropods with several cervicocephalic specializations reaching medium/large sizes. In the present contribution, we describe a new abelisaurid (Thanos simonattoi, gen. et sp. nov.) from the São José do Rio Preto Formation, Bauru Group, Brazil (Upper Cretaceous). Thanos differs from other theropods by having a well-developed keel becoming wider and deeper posteriorly on the ventral surface; two lateral small foramina separated by a relative wide wall on each lateral surface of the centrum, and well-developed and deep prezygapophyseal spinodiapophyseal fossae. The closed sutures between the axis and odontoid suggest that Thanos had reached a subadult/adult stage before death. Thanos is phylogenetically related to Brachyrostra abelisaurid. The keel on the ventral axial centrum in abelisauroids is here interpreted as a homoplastic condition that became more pronounced towards the phylogeny. The presence of well-developed keel in Thanos suggests that this taxon could be more derived than other abelisaurids. Finally, even though abelisaurids could reach large sizes, Thanos shared the environment with a larger theropod that was probably close to Megaraptora.
Figure 4. An interpretative picture of the paleoenvironment of São José do Rio Preto Formation (Santonian, Upper Cretaceous). The picture shows two specimens of Thanos simonattoi (on the right side) and titanosaur sauropods (on the mid and left sides).
Art by Deverson da Silva (Pepi).
Systematic palaeontology
Dinosauria Owen, 1842
Theropoda Marsh, 1881
Ceratosauria Marsh 1884
Abelisauroidea Bonaparte and Novas, 1985
Abelisauridae Bonaparte and Novas, 1985
Brachyrostra Canale et al. 2009
Thanos simonattoi gen. et sp. nov.
Etymology: Genus name from the Greek thánato, meaning death and from the Marvel’s character Thanos, the Conquer, created by Jim Starlin. Species name in honor of Sérgio Simonatto, the discoverer of the specimen.
.....
Conclusion:
In the present contribution, we have described a new abelisaurid from the São José do Rio Preto Formation, Upper Cretaceous of Brazil. Thanos simonattoi is distinguished from other abelisaurids by having a combination of features: a well-developed keel on the ventral surface becoming wider and deeper posteriorly; two lateral small foramina separated by a relative wide wall and two welldeveloped and deep prezygapophyseal spinodiapophyseal fossae. The keel on the ventral axial centrum of Thanos is interpreted as a homoplastic condition that became more pronounced towards the phylogeny, suggesting that this taxon could be more derived than other abelisaurids. The presence of a medium-sized abelisaurid in the São José do Rio Preto Formation increases the record of theropods, suggesting a complex carnivorous fauna during the Upper Cretaceous of Bauru Group. More findings are needed to better understand the morphology and the internal relationships of Thanos simonattoi.
Rafael Delcourt and Fabiano Vidoi Iori. 2019. A New Abelisauridae (Dinosauria: Theropoda) from São José do Rio Preto Formation, Upper Cretaceous of Brazil and Comments on the Bauru Group Fauna. Historical Biology: An International Journal of Paleobiology. DOI: 10.1080/08912963.2018.1546700
Méndez AH, Novas FE and Iori FV. 2014. New Record of Abelisauroid Theropods from the Bauru Group (Upper Cretaceous), São Paulo State, Brazil. Revista Brasileira de Paleontologia. 17(1). Citeseer: 23–32.
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رابط مدونة منتدى مدونات بلوجر جاهزة بآلاف المواضيع والمشاركات في أي وقت حــــتى لو تم حذفها من هنا شاهد صفحة منتدى مدونات بلوجر جاهزة بألاف المواضيع والمشاركات على الفيس بوك لمزيد من الشرح من هنا
تعرف على ترتيب مواضيع منتدى مدونات بلوجر جاهزة بآلاف المواضيع والمشاركات (حتى لا تختلط عليك الامور) من هنا
ملاحظة هامة: كل عمليات تنزيل، رفع، وتعديل المواضيع الجاهزة تتم بطريقة آلية، ونعتذر عن اي موضوع مخالف او مخل بالحياء مرفوع بالمدونات الجاهزة بآلاف المواضيع والمشاركات، ولكم ان تقوموا بحذف هذه المواضيع والمشاركات والطريقة بسيطة وسهلة. ــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــسلامـ.
The homology of the tridactyl hand of birds is a still debated subject, with both paleontological and developmental evidence used in support of alternative identity patterns in the avian fingers. With its simplified phalangeal morphology, the Late Jurassic ceratosaurian Limusaurus has been argued to support a II–III–IV digital identity in birds and a complex pattern of homeotic transformations in three-fingered (tetanuran) theropods. We report a new large-bodied theropod,Saltriovenator zanellai gen. et sp. nov., based on a partial skeleton from the marine Saltrio Formation (Sinemurian, lowermost Jurassic) of Lombardy (Northern Italy). Taphonomical analyses show bone bioerosion by marine invertebrates (first record for dinosaurian remains) and suggest a complex history for the carcass before being deposited on a well-oxygenated and well-illuminated sea bottom. Saltriovenator shows a mosaic of features seen in four-fingered theropods and in basal tetanurans. Phylogenetic analysis supports sister taxon relationships between the new Italian theropod and the younger Early Jurassic Berberosaurus from Morocco, in a lineage which is the basalmost of Ceratosauria. Compared to the atrophied hand of later members of Ceratosauria, Saltriovenator demonstrates that a fully functional hand, well-adapted for struggling and grasping, was primitively present in ceratosaurians. Ancestral state reconstruction along the avian stem supports 2-3-4-1-X and 2-3-4-0-X as the manual phalangeal formulae at the roots of Ceratosauria and Tetanurae, confirming the I–II–III pattern in the homology of the avian fingers. Accordingly, the peculiar hand of Limusaurus represents a derived condition restricted to late-diverging ceratosaurians and cannot help in elucidating the origin of the three-fingered condition of tetanurans. The evolution of the tridactyl hand of birds is explained by step-wise lateral simplification among non-tetanuran theropod dinosaurs, followed by a single primary axis shift from digit position 4 to 3 at the root of Tetanurae once the fourth finger was completely lost, which allowed independent losses of the vestigial fourth metacarpal among allosaurians, tyrannosauroids, and maniraptoromorphs. With an estimated body length of 7.5 m, Saltriovenator is the largest and most robust theropod from the Early Jurassic, pre-dating the occurrence in theropods of a body mass approaching 1,000 Kg by over 25 My. The radiation of larger and relatively stockier averostran theropods earlier than previously known may represent one of the factors that ignited the trend toward gigantism in Early Jurassic sauropods.
Figure 1: Fossil location and geological setting. (A–C) Outline maps of Italy, Lombardy, Varese Province, and Saltrio Municipality; (D) satellite view of the Saltrio area, with map marker indicating the Saltrio quarry; (E) map marker indicating the stratigraphic log in the Saltrio quarry; (F) the ammonite Paracoroniceras cf. gmuendense and (G) the nautiloid Cenoceras striatum, both found associated in the layer containing the dinosaur bones; (H) glauconite present as accessory mineral in block C (counterpart of block A of Fig. 2); (I) the discordance between the Dolomia Principale Fm. and the Saltrio Fm.; (J) thin sections of the layer embedding the dinosaur bones; (K) stratigraphic log of the Saltrio quarry, based on Croce (2005), with geological time scale and ammonites zones based on Sacchi Vialli (1964) and Ogg & Hinnov (2012). Abbreviations: c, crinoids; f, foraminifers; g, gastropods; o, ostracods. Scale bars equal 200 km in (A), 30 km in (B), six km in (C), one km in (D), one mm in (K), and 150 cm in (L). Photos by F. Berra, G. Bindellini, M. Croce, and G. Pasini; drawings by M. Croce and S. Maganuco.
Figure 1: Fossil location and geological setting. (A–C) Outline maps of Italy, Lombardy, Varese Province, and Saltrio Municipality.
Scale bars equal 200 km in (A), 30 km in (B), six km in (C).
Figure 1: Fossil location and geological setting. (D) satellite view of the Saltrio area, with map marker indicating the Saltrio quarry; (E) map marker indicating the stratigraphic log in the Saltrio quarry; (F) the ammonite Paracoroniceras cf. gmuendense and (G) the nautiloid Cenoceras striatum, both found associated in the layer containing the dinosaur bones; (H) glauconite present as accessory mineral in block C (counterpart of block A of Fig. 2); (I) the discordance between the Dolomia Principale Fm. and the Saltrio Fm.; (J) thin sections of the layer embedding the dinosaur bones; (K) stratigraphic log of the Saltrio quarry, based on Croce (2005), with geological time scale and ammonites zones based on Sacchi Vialli (1964) and Ogg & Hinnov (2012). Abbreviations: c, crinoids; f, foraminifers; g, gastropods; o, ostracods.
Scale bars equal one mm in (K), and 150 cm in (L). Photos by F. Berra, G. Bindellini, M. Croce, and G. Pasini; drawings by M. Croce and S. Maganuco.
Figure 2: Taphonomy of the Saltrio theropod (block A). Bones of Saltriovenator mapped in temporal sequence (A–C), gradually emerging from the embedding rock during acid preparation of block A. Numbers refer to each fragment, not to a specific anatomical position. The latter is reported in other figures, for fragments that were later reconnected into more complete bones. Abbreviations as in text, and as follows: ind, indeterminate bone; ir, indeterminate rib; l (left) and r (right) are specified for fragments of paired bones certainly (appendicular elements) or tentatively (ribs) positioned in the skeleton. Macroborings facing front, side and back are mapped respectively with yellow circles, semicircles, and hatched circles. Scale bars equal 10 cm. Photos by G. Bindellini and C. Dal Sasso.
Figure 4: Selected elements used in the diagnosis of Saltriovenator zanellai n. gen. n. sp. Right humerus in medial (A), frontal (B) and distal (C) views; (D) left scapula, medial view; (E) right scapular glenoid and coracoid, lateral view; (F) furcula, ventral view; tooth, labial (G) and apical (H) views; (I) left humerus, medial view; right second metacarpal in dorsal (J), lateral (L) and distal (N) views; first phalanx of the right second digit in dorsal (K), lateral (M) and proximal (O) views; (P–T) right third digit in proximal, dorsal and lateral views; (U) right distal tarsal IV, proximal view; third right metatarsal in proximal (V) and frontal (X) views; second right metatarsal, proximal (W) and frontal (Y) views; (Z) reconstructed skeleton showing identified elements (red).
Abbreviations as in text, asterisks mark autapomorphic traits.
Scale bars: 10 cm in (A)–(E), (I), and (U)–(Y); two cm in (F), and (J)–(T); one cm in (G).
Photos by G. Bindellini, C. Dal Sasso and M. Zilioli; drawing by M. Auditore.
Figure 5: Cranio-mandibular fragments, tooth, and ribs of Saltriovenator zanellai. Indeterminate cranial fragment (A–B); right splenial in lateral (C), rostral (D) and ventral (E) views; right prearticular in lateral (F) and rostral views (G); sketch of the right prearticular of MOR 693 (Allosaurus fragilis) with virtual cross-section (H) diagnostic for G, also confirmed by CT slicing of the left side element of MOR 693 (I); splenial and prearticular in medial view, positioned in a reconstructed right lower jawof Saltriovenator (J). Maxillary or dentary tooth in labial (K) and apical (L) views; close-up of the distal carina and denticles in lingual (M) and distal (N) views. Left cervical rib (O) in craniolateral view; fragmentary right (P) and left (Q) dorsal ribs in craniolateral view.
Abbreviations as in text, ribs labeled as in Fig. 2 maps and caption. Scale bars equal two cm in (A)–(I), five cm in (J), one cm in (K), five mm in (L), one mm in (M)–(N), five cm in (O)–(Q).
Photos by G. Bindellini, C. Dal Sasso, and M. Zilioli; drawing by C. Dal Sasso.
Systematic Paleontology
DINOSAURIA Owen, 1842
THEROPODA Marsh, 1881
NEOTHEROPODA Bakker, 1986
CERATOSAURIA Marsh, 1884
Saltriovenator zanellai gen. et sp. nov.
Etymology. Saltrio, Italian toponym name, from the locality where the holotype was found; venator, Latin word for hunter, it also refers to a type of Roman gladiator; zanellai, Latin genitive dedicated to Angelo Zanella, who discovered the fossil.
Holotype. MSNM V3664, very fragmentary and disarticulated skeleton (Figs. 4–13), represented by the following elements (among brackets, number of fragments per bone): partial right splenial (2) and right prearticular (1); cervical (1) and dorsal (9) ribs; furcula (1), incomplete left scapula (16), right scapular glenoid (1), partial right coracoid (5), fragmentary right sternal plate (2); right humerus (2), and proximal half of left humerus (2); ?right ?distal carpal, right metacarpal II, right phalanx II-1, fragmentary right phalanx II-2, and tip of the ?second right ungual phalanx; complete third right manual digit (phalanges III-1 to III-4); right distal tarsals III and IV, proximal portions of right metatarsals II, III, IV, and V(2).
Referred material. MSNM V3659, one maxillary or dentary tooth (Figs. 4 and 5).
Comments. As noted above, the discovery of all skeletal elements at the same time in a very restricted spot, the fact that all of them are of matching size, and that fragmentary and anatomically adjacent elements are of matching morphology, leave no doubt that all bones referred to the holotype come from the same individual. We prudentially exclude from the holotype the single tooth, which was found relatively associated to the bones but lacking its root and any jaw bone connection, thus raising the doubt that it might represent a shed tooth.
Type locality. “Salnova” quarry, Saltrio, Varese Province, Lombardy (northern Italy).
Horizon and Age. Saltrio Fm. (sensu Gnaccolini, 1964), bucklandi Zone, early Sinemurian (199.3–197.5 mya) (Ogg & Hinnov, 2012).
Diagnosis. Mid-to-large sized ceratosaurian characterized by the following unique combination of anatomical features (autapomorphies marked by asterisk—see also Fig. 4): humerus with deltopectoral crest protruding craniomedially for more than twice the shaft diameter, with distal lamina forming an abrupt corner (about 90°) with the proximodistal axis of the humeral shaft; metacarpal II with hypertrofied semicircular extensor lip protruding over the condylar level* and bordering dorsolaterally a very deep and wide extensor pit; phalanx II-1 with flexor palmar groove which is deep and narrow*, and bearing a distinct bump distal to the dorsal extensor process*; manual ungual III with prominent flexor tubercle which is distinctly separated from articular facet by a concave cleft.
.....
Simplified evolutionary tree of predatory dinosaurs (theropods). Saltriovenator predates the massive meat-eating dinosaurs by over 25 million years: it is the oldest known ceratosaurian, and the world's largest predatory dinosaur from the Lower Jurassic. During the Jurassic, the three- fingered tetanuran theropods appeared, which gave rise to birds.
Conclusions
Saltriovenator zanellai gen. et sp. nov. is a new theropod dinosaur from the Lower Jurassic of Northern Italy. It represents the third named species of non-avian dinosaur from Italy, the first of Jurassic age. Saltriovenator shows a combination of ceratosaurian and tetanuran features, supporting close relationships between the two averostran lineages with the exclusion of coelophysoid-grade theropods. It also represents the first skeletal material supporting the occurrence of large and robustly-built predatory dinosaurs just at the aftermath of the Triassic–Jurassic boundary extinction events. Accordingly, the Italian ceratosaurian fills a stratigraphic and ecomorphological gap between the relatively more gracile coelophysoid-grade neotheropods (known from the Late Triassic to the Early Jurassic) and the large-bodied averostrans that occupied the majority of the apex predatory roles in the terrestrial ecosystems between the Middle Jurassic and the end of the Cretaceous.
The phylogenetic framework integrated with the new combination of features present in Saltriovenator dismisses the “II–III–IV homology pattern” in the interpretation of the tetanuran (and avian) hand, and suggests a complex process leading to the atrophied forelimb of later ceratosaurians. The evolution of a stocky and robust hand occurred in ceratosaurians before the relative shortening and the loss of predatory function: such a step-wise scenario raises intriguing perspectives on what adaptive and developmental factors led from a “Saltriovenator-like” condition to the aberrant condition present in Limusaurus and abelisaurids.
Cristiano Dal Sasso, Simone Maganuco and Andrea Cau. 2018. The Oldest Ceratosaurian (Dinosauria: Theropoda), from the Lower Jurassic of Italy, Sheds Light on the Evolution of the Three-fingered Hand of Birds. PeerJ. 6:e5976. DOI: 10.7717/peerj.5976
كيف تحصل على مدونة جاهزة بآلاف المواضيع والمشاركات من هنا
شاهد قناة منتدى مدونات بلوجر جاهزة بألاف المواضيع والمشاركات على اليوتيوب لمزيد من الشرح من هنا
رابط مدونة منتدى مدونات بلوجر جاهزة بآلاف المواضيع والمشاركات في أي وقت حــــتى لو تم حذفها من هنا شاهد صفحة منتدى مدونات بلوجر جاهزة بألاف المواضيع والمشاركات على الفيس بوك لمزيد من الشرح من هنا
تعرف على ترتيب مواضيع منتدى مدونات بلوجر جاهزة بآلاف المواضيع والمشاركات (حتى لا تختلط عليك الامور) من هنا
ملاحظة هامة: كل عمليات تنزيل، رفع، وتعديل المواضيع الجاهزة تتم بطريقة آلية، ونعتذر عن اي موضوع مخالف او مخل بالحياء مرفوع بالمدونات الجاهزة بآلاف المواضيع والمشاركات، ولكم ان تقوموا بحذف هذه المواضيع والمشاركات والطريقة بسيطة وسهلة. ــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــسلامـ.
The hypothetical phylogenetic relationships of ceratosaurs based on current topologies.
The main source is from Wang et al. (2016).
Hypothetical reconstruction of two abelisaurids showing the soft tissues on the head inferred from osteological morphology of the skull. On the top, Carnotaurus; on the bottom, Pycnonemosaurus. Art by Maurilio Oliveira.
Abstract Ceratosaur theropods ruled the Southern Hemisphere until the end of the Late Cretaceous. However, their origin was earlier, during the Early Jurassic, a fact which allowed the group to reach great morphological diversity. The body plans of the two main branches (Noasauridae and new name Etrigansauria: Ceratosauridae + Abelisauridae) are quite different; nevertheless, they are sister taxa. Abelisaurids have lost the ability to grasp in the most derived taxa, but the reduced forelimb might have had some display function. The ontogenetic changes are well known in Limusaurus which lost all their teeth and probably changed the dietary preference at maturity. The results presented here suggest that abelisaurids had different soft tissues on the skull. These tissues might have been associated with evolution of a strong cervicocephalic complex and should have allowed derived taxa (e.g. Majungasaurus and Carnotaurus) to have low-displacement headbutting matches. The ability to live in different semi-arid environment plus high morphological disparity allowed the ceratosaurs to become an evolutionary success.
Figure 1 The hypothetical phylogenetic relationships of ceratosaurs based on current topologies. The main source is from Wang et al. (2016). The phylogenetic position of Chenanisaurus is from Longrich et al.(2016) and the Ligabueino, Austrocheirus, Majungasaurinae and Brachyrostra are from Filippi et al.(2011).
Figure 4 Skin structures inferred for abelisaurids. Dorsal surface of the skull of (A) Rugops (MNN IGU1), (C) Carnotaurus (MACN-CH 894) and dorsal surface of the fused nasal of (B) Abelisaurus (MPCA 11908). Scales bar: 5 cm.
Figure 6 Hypothetical reconstruction of two abelisaurids showing the soft tissues on the head inferred from osteological morphology of the skull. On the top, Carnotaurus; on the bottom, Pycnonemosaurus.
Art by Maurilio Oliveira.
Rafael Delcourt. 2018. Ceratosaur Palaeobiology: New Insights on Evolution and Ecology of the Southern Rulers. Scientific Reports. 8, 9730. DOI: 10.1038/s41598-018-28154-x
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شاهد قناة منتدى مدونات بلوجر جاهزة بألاف المواضيع والمشاركات على اليوتيوب لمزيد من الشرح من هنا
رابط مدونة منتدى مدونات بلوجر جاهزة بآلاف المواضيع والمشاركات في أي وقت حــــتى لو تم حذفها من هنا شاهد صفحة منتدى مدونات بلوجر جاهزة بألاف المواضيع والمشاركات على الفيس بوك لمزيد من الشرح من هنا
تعرف على ترتيب مواضيع منتدى مدونات بلوجر جاهزة بآلاف المواضيع والمشاركات (حتى لا تختلط عليك الامور) من هنا
ملاحظة هامة: كل عمليات تنزيل، رفع، وتعديل المواضيع الجاهزة تتم بطريقة آلية، ونعتذر عن اي موضوع مخالف او مخل بالحياء مرفوع بالمدونات الجاهزة بآلاف المواضيع والمشاركات، ولكم ان تقوموا بحذف هذه المواضيع والمشاركات والطريقة بسيطة وسهلة. ــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــــسلامـ.
