Fayyum (fossil deposit)
Coordinates: 29° 22′ 0″ N , 30° 47′ 0″ E
The Fayyum is a region and an important fossil site in northern Egypt . The region includes the Fayyum Basin , which is used intensively for agriculture, and adjoining areas, the significant fossil sites are mostly exposed north and west of Lake Qarun . The south-western end is formed by the Wadi al-Hitan , known for numerous whale fossils and a UNESCO World Heritage Site since 2005 . The deposits of the Fayyum belong to different geological formations . They are mainly composed of limestone ,siltstone and sandstone together. The lower sections consist of marine deposits, the upper continental sediments are formed in a coastal landscape. The formation period ranges from the Middle to the Upper Eocene to the Lower Oligocene , which corresponds to an age of around 41 to 28 million years ago. The entire sedimentary complex is overlain by basalt that dates back to volcanic activity about 24 million years ago.
The very rich fossil evidence of the Fayyum includes plants , invertebrates and vertebrates , and numerous trace fossils have also survived. Among the vertebrates are fish , reptiles , birds and mammals . Some forms represent important evolutionary links in the development of individual lineages, particularly in the hyrax , the proboscidea , the manatee , and the whale. Among the fossil primates are early forms of Old World monkeys , among others . In addition, marsupials , tenrecs , even-toed ungulates , bats and rodents also occur. Some groups such as the giant, herbivorous Embrithopoda or the predatory Hyaenodonta or the probably insectivorous Ptolemaiida have no living descendants. Because of this diversity, the Fayyum represents the most important fossil site of the African Palaeogene . Based on the terrestrial vertebrates, a delta-like landscape criss-crossed by watercourses can be reconstructed near the sea coast under tropical climatic conditions .
The first fossil finds in Fayyum go back to the middle of the 19th century. A major phase of investigation took place at the transition from the 19th to the 20th century, when at times English, American, German and French researchers were working on site at the same time. Numerous fossils were recovered during this period; this led to some important discoveries that contributed to the nationwide fame of the Fayyum area. In addition, the first geological investigations were carried out . Modern research began in the 1960s and continues to this day. It is organized internationally. Analyzes of the find material lead to the description of new fossil forms from the Fayyum almost every year.
Geographical location
The Fayyum region covers an area of around 12,000 km² and is located around 70 km southwest of Cairo on the left-hand side
des Nils inmitten der Libyschen Wüste. Das Zentrum bildet das Fayyum-Becken, ein etwa 1500 km² großes Depressionsgebiet. Es weist eine im Umriss dreieckige Form auf mit nach Süden zeigender Spitze. Zentralort ist al-Fayyūm, der zudem auch die Hauptstadt des Gouvernements al-Fayyum bildet. Im Norden des Beckens liegt der etwa 230 km² große Qarun-See, ein abflussloser See, der eine Länge von 40 km von Nordost nach Südwest aufweist bei einer durchschnittlichen Tiefe von rund 5 m und einer maximalen von 7,5 m. Als einzelne Landmasse ragt die Insel Geziret el-Qorn aus dem Wasser. Der Qarun-See (deutsch für „See der Hörner“, nach den beiden spitzen Halbinseln, die vom Norden in den See hineinreichen) ist der größte See des nördlichen Afrikas, er stellt aber nur den Rest eines noch größeren Sees dar. Dieser bestand in prähistorischer und historischer Vergangenheit und trägt die Bezeichnung Moeris-See. Die Depression senkt sich von Süd nach Nord ab. Befindet sich der Zentralort al-Fayyūm noch bei etwa 24 m über dem Meeresspiegel, so liegt die Wasserfläche des Qarun-Sees bei etwa 45 m unter dem Meeresspiegel. Das gesamte Fayyum-Becken wird landwirtschaftlich genutzt, zwei Wassersammler (El Bats im Osten und El Wadi im Westen) drainieren das überschüssige Wasser in den Qarun-See.
The area surrounding the Fayyum Basin consists largely of desert-like terrain. In the east, a 5 to 12 km wide ridge separates the depression from the Nile valley , its top is about 30 to 90 m above sea level. These edge mountains show an approximately 1.5 km wide opening through which the Bahr Yusuf also runs. The canal connects the Nile with the Fayyum Depression over a length of 270 km. It ends at al-Fayyūm and here it branches out into numerous small arms. To the south follows the small basin of el-Gharaq , which has a maximum depth of 4 m below the sea surface. It is joined to the southwest and west of the Fayyum Basin by Wadi el-Rayyan . This is an approximately 700 km² large depression with the deepest point at about 42 m below sea level. The wadi is fed by its own springs and is now partly filled by Lake el-Fayyum , a two-part artificial lake of 51 and 62 km² respectively. However, the northern boundary of the Fayyum Basin, where several escarpments rise, is significant. They run from southwest to northeast over a length of around 70 km and are separated from each other by plateau-like plains. The terrain thus rises from Lake Qarun at its lowest point to an average of 350 m above sea level. The main escarpments are, from south to north, the Birket Qarun Escarpment , the Qasr el-Sagha Escarpment and the Gebel Qatrani Escarpment . In between there are still smaller escarpments such as the el-Ekhwat Escarpment and the Talata Escarpment .
The south-west end of the escarpment marks the Wadi el-Hitan (“Valley of the Whales”, also Zeuglodon valley ), around 50 km from Lake Qarun. The area has been a UNESCO World Heritage Site since 2005 . It is a wide, flat valley some 10 km long, which is not an actual wadi in the geomorphological sense. It is bounded on the northwest by the Birket Qarun Escarpment . In the southwest, on the other hand, a series of flat-topped hills forms the conclusion (the so-called "boundary hills"). Due to these two geographic borders, the valley narrows from about 4 km in width in the north-east to about 0.5 km in the south-west. About 5 km to the north-east towards the Fayyum Basin extends another group of hills, below them Garet Gehannam rises about 120 m above the surrounding area as an important landmark.
geology
General and underground
The geological situation of the Fayyum region has been intensively researched since the end of the 19th century. Significant work on this was published in 1905 by Hugh John Llewellyn Beadnell , 1988 by Thomas M. Bown and Mary J. Kraus , and 1992 by Philip D. Gingerich . The Fayyum Basin represents a tectonically engineered structure dug into the Triassic subsoil. However , rock layers from this period, such as the Bahariya Formation , only appear on the outermost edges of the region, for example in the north-east. The main faults run southwest-northeast and belong to the Syrian arc fold belt , which runs in an S-arc from the Levant across the Sinai Peninsula to North Africa . They also determine the formation of the large escarpments north of Lake Qarun. Subordinate fault zones also occur from north-west to south-east and from east to west. They frame the Fayyum Basin on all sides. It can therefore be assumed that the basin was formed by tectonic forces. These shaped the depression in the transition from the Oligocene to the Miocene , possibly in connection with local volcanism . In the region itself, deposits from the Eocene and Oligocene are mainly formed, with a total thickness of up to 800 m.
Eocene to Oligocene sedimentary sequence
Mokattam group
The footwall of the Fayyum region is generally assigned to the Mokattam Group (also Wadi-el-Rayyan series), named after the Gebel Mokattam east of Cairo. It is marine in origin and consists of four rock units, the Muweilih Formation , the Midawara Formation , the Sath-el-Hadid Formation and the Gharaq Formation . All four formations are composed of limestones , which differ in their structure and in the content of nummulites and glauconites . The upper end of the Mokattam Group is defined by a rapid decline in numulites and an increase in glauconites. The Gharaq Formation is the most widely exposed unit of the Mokattam Group in the region and is found mostly in the southwestern Fayyum area from Wadi el-Rayyan to the southern part of Wadi el-Hitan and as far as the Garet Gehannam .
Maadi group
The Gehannam Formation (also Ravine beds ) is the base member of the Maadi Group. It mainly consists of marine clays , silts and sandy silts, in which marly and clayey limestones are incorporated. Their total thickness is about 46 m. The base consists of glauconite clay / siltstones that are finely laminated and merge into calcareous sandstones at the top . This is followed by silt, limestone and marlstone . In some outcrops, limestone forms the upper end. In some cases, individual layers of gypsum are embedded in the upper sections of the Gehannam Formation . The Gehannam Formation in the western Fayyum area at Wadi el-Hitan and at Garet Gehannam is exposed particularly well . It also forms the bedrock of most of the cultivated land in the basin.
All other formations are widely distributed in the bluffs north of Lake Qarun. The Birket Qarun Formation forms the lower escarpment immediately north of Lake Qarun, the Birket Qarun Escarpment . It presents itself as a rock unit up to 85 m thick, the main components of which are made up of fine to very fine sands and silts, which are sometimes very calcareous. Basal is a sandstone rich in sea urchins called the Schizaster lybica sandstone . Gypsum deposits have formed in individual areas. In general, the deposits go back to a coastal sea. In the outcrops north of Lake Qarun, they appear as a uniform sedimentary complex that fans out westward towards Wadi el-Hitan and interlocks with the Gehannam Formation in its lower and middle sections. Here, the Birket-Qarun Formation can be subdivided into four individual bodies, which can usually be clearly separated from the Gehannam Formation. A series of narrow, block-like sandstone layers, some only 30 cm thick, appear as a conspicuous phenomenon, the uppermost of which is referred to as the Camp White Layer . The latter is very distinctive because, in addition to numerous whale bones, it contains conspicuous vertical structures that are interpreted either as roots of mangroves or burial tracks of the track genus Thalassinoides (in some older works, the Camp White Layer is considered to be the boundary layer between the Gehannam and Birket Qarun formation, but more recent studies place it within the upper rock unit).
The Garet-el-Naqb formation was only defined as a possible rock formation in 2014, but it was noticed even before that as a special formation. It is a dark gray clay that increases in thickness from south to north. It reaches its greatest thickness north of the Wadi el-Hitan with around 25 m. The rock unit occurs largely only in the western Fayyum area, north of Lake Qarun it is not documented. In the western Fayyum outcrops, the Garet-el-Naqb Formation divides the Birket-Qarun Formation as a prominent dark horizon or directly overlies the Gehannam Formation.
The Qasr-el-Sagha formation (also Carolia beds ) completes the Maadi group. It is exposed at the Qasr el-Sagha Escarpment and is up to 200 m thick. The rock unit consists of sandstones, sandy clay/siltstones and calcareous stratified rocks with layers of limestone in between. Four subunits can be distinguished. At the bottom is the Umm Rigl Member with a 30 to 65 m thick sequence of sandy, partly bioclastic, hard limestone, which alternates with gypsum and calcareous, finely layered clay and siltstone. This is followed by the Harab Member , which is a 30 to 40 m thick package of brown, structureless clays. The Temple Member , in turn, is similar in structure to the Umm Rigl Member . The overhanging Dir Abu Lifa Member is up to 77 m thick and is characterized by an alternation of cross-layered sandstones and fine-grained, gypsum-bearing clay/siltstones and sandstones. The deposits go back to a shallow, coastal sea. The base of the formation has not been judged consistently in the past. Some authors pinned it with the first appearance of the Carolia shell , others with an argillaceous limestone a few meters higher. The upper transition to the next rock unit is discontinuous.
Gebel Qatrani Formation
The Gebel-Qatrani Formation (also fluvio-marine series ) lies unconformably on the Qasr-el-Sagha Formation . It reaches a thickness of 340 m and forms the highest escarpment with the Gebel Qatrani Escarpment . To the west, the formation thins out significantly and is then sometimes less than 100 m thick. It stands out from the Qasr-el-Sagha formation due to its composition of various fine to coarser sandstones as well as clay/siltstones. In principle, three zones can be distinguished: a lower zone of coarse, loose sandstone (the lower fossil wood zone , thickness 153 m), a middle zone of banked, medium to coarse-grained and partly extremely gypsum-containing sandstone as a marker horizon (thickness 7 to 10 m) and an upper one consisting of an alternation of sandstone, sandy mudstone/siltstone with incorporated lenses of coarse sandstone and limestone (the upper fossil wood zone , 190 m thick). The designations lower and upper fossil wood zone go back to the fossilized tree trunks found here. Both the lower and the upper series can be broken down into numerous subunits, of particular note are several sequences of colored banded sedimentary sequences that are distributed over the entire Gebel-Qatrani Formation and are referred to as variegated sequences . Especially in the upper fossil wood zone , they grow up to 70 m thick. The bands go back to fossil soil formation processes and the associated shifting and accumulation of soluble iron compounds . In addition, there is other evidence of paleosoils such as clay deposits. The soils show various stages of development from initial ( inceptisols ) to developed soil formations ( spodosols and ultisols ). Overall, the secondary soil overprints indicate a clearly terrestrial depositional environment of the Gebel-Qatrani Formation. It can therefore be viewed as a continental fluvial sedimentary sequence formed by the action of numerous rivers and watercourses.
Widan-el-Faras basalt
The upper end of the Eocene-Oligocene series is marked by the Widan-el-Faras basalt , separated from the underlying Gebel-Qatrani Formation by a stratigraphic gap . It consists of dense, extremely fine-grained and ferrous basalts . Individually included sand horizons show that the basalts form several streams, the thickness and characteristics of which are, however, highly variable. The greatest thickness is 25 m, in places the Widan-el-Faras basalt can be only 2 m thick.
top layers
The Neogene cover layers lie on top of the Paleogene sedimentary sequence . They are assigned to the Kashab Formation in the Fayyum area , which in turn is distinguished from the Widan-el-Faras basalts by a hiatus . In areas where the basalt is not formed, it directly covers the Gebel Qatrani Formation. The Kashab Formation consists mainly of alluvial sediments, the lower 10 to 12 m being indicated by a boulder-bearing conglomerate . Their total thickness is over 100 m. The Kashab Formation forms the northern hinterland of the Fayyum region.
Development of the Fayyum Depression in the Pleistocene and Holocene
The shape of today's Fayyum Basin was formed during the alternating cold and warm phases of the Pleistocene . The extreme periods of drought during the cold periods repeatedly led to the emptying of the boiler through deflation down to the groundwater level. In the wetter interglacial periods, the basin filled with water and formed a swamp area. These swamp areas were still detectable in historical times and also led to the naming of the Fayyum region (from the Egyptian phiom for "swamp"). The most recent formation of the Holocene is referred to as Lake Moeris . Its natural development began as early as the Lower Holocene and ran through several phases, which can be demonstrated above all by the distribution of diatoms , but also by old shorelines. They led to fluctuating lake levels, which is partly related to the formation and rupture of the natural connection to the Nile. The oldest represents the "Paleomoeris", it existed from 8,830 to 8,220 years BP . During this time, the lake level reached a high of 17 m above sea level. It was followed by the "Premoeris" (between 8,200 and 7,500 years BP) and the "Protomoeris" phase (7,500 to 7,140 years BP), during which the lake level at times fluctuated between 28 and 8 m above sea level. The last phase includes the "Moeris" phase, it started around 6,095 BP and lasted until around 5,500 years BP. Here the lake surface was up to 20 m above sea level and formed a lake with an area of 2100 km² and a volume of 53 km³. In the period that followed, however, it fell back several meters due to the loss of the connection to the Nile, and a swampy area developed.
In ancient Egyptian times, new land was made accessible and arable, among other things, through water management construction measures. This happened, for example, in the Middle Kingdom through the construction of the Bahr Yusuf along the old entrance to the Nile and through a drainage system in the Fayyum Basin. In this way, about 450 km² of land could be used. Later came the construction of an 8 km long dam at el-Lahun and the creation of further fresh water reservoirs in the south-east of the Fayyum area. By the Ptolemaic period , the usable area had been expanded to around 1300 km², which corresponds to a large part of the cultivated land today. At that time, Lake Moeris had shrunk to around 415 km² and occupied the area of today's Lake Qarun. Further declines characterized the Roman era, during which the water surface dropped from 7 m below sea level to 17 m. In the first half of the 19th century it was already around 40 m, while the low point of 46 m below sea level was reached in the 1930s. As a result, Lake Qarun developed from a former freshwater reservoir into a highly saline body of water, which is due on the one hand to the lower water inflow compared to evaporation and on the other hand to the discharge of chemically polluted agricultural wastewater. Aside from that, the lake is an important refuge for migratory and waterfowl. It serves as a breeding ground for Shepherd and Kentish Plovers , Red-winged and Little Terns , while Great Crested Grebes and Black- necked Grebes winter here.
fossil sites
The fossils come from several geological formations . Plant fossils occur in greater numbers only in the Gebel-Qatrani Formation. In contrast, invertebrates can be found in almost all rock units, while vertebrates are again more restricted. Marine forms are distributed among the lower units and are common in the Muweilih, Gharaq, Gehannam and Birket Qarun Formations, but they also survive from the Qasr el Sagha Formation. Sometimes the larger marine mammals in particular were preserved in complete skeletons. The Qasr-el-Sagha Formation and the following Gebel-Qatrani Formation represent the main find areas of the terrestrial life forms. The material is mostly fragmented and disarticulated, so that certain rearrangements can be assumed. There are countless sites and outcrops throughout the Fayyum area. They lie in the different stratigraphic units and are usually concentrated north and west of Lake Qarun. More than 100 sites have been documented for terrestrial mammals alone. These are divided between a few isolated ones in the Birket Qarun Formation and a little less than a dozen in the Qasr el Sagha Formation. The much larger part of over 90% belongs to the Gebel Qatrani formation. The site BQ-2 from the Umm Rigl Member of the Qasr-el-Sagha Formation is of great importance (individual, mainly paleontological studies refer the Umm Rigl Member to the Birket-Qarun Formation, which is attributed to insufficient exploration conditions at the site itself is) near the easternmost tip of Lake Qarun. BQ-2 was only discovered in 2000, from where the most diverse vertebrate fauna of the entire Fayyum area originates. Most of the other sites in the Qasr el Sagha Formation are in the Dir Abu Lifa Member , with the exception of one unstratified site in the Temple Member . In general, however, terrestrial mammals are less common here.
Of the sites from the Gebel-Qatrani Formation, eight ( A , B , E , G , I , M , V and L-41 ) are of particular interest, as they contain around 90% of all terrestrial vertebrate material. The most significant finding here is found with L-41 . It was discovered in 1983 by geologist Thomas M. Bown . Bown was involved in the research done here by Elwyn L. Simons since the 1960's, he was made aware of this locality by two mandibles of hyrax . A more detailed description of the site was presented a few years later by D. Tab Rasmussen , among others. Standing 47 m above the base of the rock unit, L-41 marks the oldest occurrence of the formation to date and is believed to be the most productive. Slightly higher in the geological sequence, but still in the Lower Zone, sites A , B and E are located. They were already opened at the beginning of the 20th century and form the oldest stratified fossil deposits in terms of research history. Site G , which is about 16 m above the sandstone bank separating the lower from the upper zone, is also of some importance . The main investigations took place in the 1960s. Another 66 m higher are the two sites I and M , which are among the youngest within the Gebel-Qatrani Formation.
In general, the fossil remains are diagenetically overprinted and permineralized , so that the organic material has been replaced by anhydrite and gypsum . However, the bones and teeth from the Dir Abu Lifa Member of the Qasr el Sagha Formation appear to have undergone and have undergone another diagenetic process. In many cases, the sulfates were replaced by carbonates . As a result, the surface structures of the bones and teeth were less well preserved than the finds from the Gebel-Qatrani Formation.
finds
flora
Macro remains of plants are in the form of woods , leaves , fruits , and seeds . Leaf remains from the Cymodoceaceae group are occasionally found, especially from the older rock units , including the “sea grasses” Thalassodendron and Cymodocea . Otherwise, plant remains are largely confined to the Gebel-Qatrani Formation. The woods come in a wide variety of sizes, ranging from small twigs to tree trunks that are 44 m long and more than 2 m thick. The bark is still present on some trunks. Some of these tree trunks lie together as fossilized forests, which, among other things, consist of up to 200 individual pieces in the lower sections of the Gebel Qatrani Formation. The vegetation comprises around two dozen different families, eight of which were identified under the woods, fruits and seeds, and thirteen under the leaves. Typical are, among other things, the floating ferns such as Salvinia , which are bound to stagnant or slowly flowing water , as well as mangrove ferns such as Acrostichum , which lives in brackish water . Among the monocots , the palm -like ones stand out, of which the extinct genus Palmoxylon , among others, has been described. Other monocots can be found with the ivy from the group of arum plants , which climb trees like lianas. Some of their fragile fruits can be salvaged while they are still intact. However, the most common are dicots , including the lotus family , mallow family , carob family or sapote family . The entire flora shows relationships to the Indo-Malaysia region.
fauna
invertebrates
Invertebrates can be found in all layers of rock, usually the remains of marine fauna. A large part of this is taken up by the foraminifera , single- celled organisms that carry shells and occur in the Fayyum as planktonic as well as benthic forms. Planktonic foraminifera are represented by Truncorotaloides , Turborotalia or Globigerinatheca , benthic by Operculina Discocyclina or Nummulites . Some forms like Frondicularia or Pullenia refer to quite large sea depths. In individual rock sections, the foraminifera are quite numerous, with up to over 30 species. Other invertebrates belong to the sea urchins such as schizaster , a representative of the heart sea urchins , or more rarely lance sea urchins . In addition, mollusks are also documented. Among the mussels , oysters , moon mussels and saddle clams are of great importance, with Carolia forming an important stratigraphic marker for the latter. In turn, moon snails and tower snails can be found among snails . Cnidarians are also found . Occasionally crabs have survived, as shown by individual shell parts of Lobocarcinus from the group of edible crabs in the middle Birket-Qarun Formation. The carapaces are 9 to 14 cm wide, with marked differences between the sexes in the carapace shape.
fish and amphibians
Fish also show a wide stratigraphic distribution in the Fayyum area. Most of the find material consists of teeth and a few isolated bones, occasionally there are also articulated skeletal remains. Marine fish occur in all geological sections of the Fayyum region, but are more prevalent in the Mokattam Group and the lower reaches of the Maadi Group. The fauna is mainly characterized by sharks and rays . Larger sharks are found with the tiger shark , which is very common in the Midawara Formation, for example. Other forms are sharpnose sharks , hammerhead sharks and mackerel sharks , the latter including a representative of the genus Otodus , which can be up to 5 m long . However, the most extensive finds can be assigned to the lemon sharks , which occur in a larger and smaller form and have their distribution center in the Maadi group. In addition, a large number of medium-sized and smaller sharks were recorded, such as nurse sharks , thresher sharks , sand sharks , sixgill sharks and smooth sharks . The genus Moerigaleus , which is typical for weasel sharks , has a set of teeth with a wide variety of designs, of which over 60 are known to date. With the fossil shark , which also belongs to this group, a form occurs whose present-day relatives mostly hunt cephalopods in the shelf area . Also noteworthy are some requiem sharks such as Carcharhinus and Misrichthys , which form some freshwater component and occur in comparatively larger numbers in the upper reaches of the Maadi Group. An increased fossil evidence in these geological layers is also recorded for the related Abdounia . Large rays mostly belong to the group of stingrays , including cownose rays, which are quite common in the Midawara and Qasr el Sagha formations, but almost disappear in between. In contrast, other forms such as Leidybatis or Lophobatis are generally rare. Propristis and Anoxypristis , representing the sawfish , have also been recorded . On the other hand, Coupatezia and Hypolophodon are related to the stingrays , they indicate a stronger freshwater influence in the upper sections of the Maadi group. A very rare faunal element can be found with the torpedo rays .
In general, marine elements fade into the background in the upper sections of the Maadi Group from the Qasr el Sagha Formation and freshwater or brackish water fish are more prevalent. Here the bony fish dominate with varied forms. From the Gehannam Formation there are remains of Pycnodus from the extinct group of Pycnodontiformes , which apparently fed on hard-shelled mussels. From the more recent deposit series, for example, the bichirs could be covered with Polypterus . Also of note are the Great Nile pike and catfish such as Chrysichthys and Auchenoglanis . An almost complete, 23 cm long individual is available from Chasmoclupea from the herring group . Cross catfish , on the other hand, indicate that there was still a marine influence, which was documented, among other things, by the genus Qarmoutus with a largely complete skeleton. This is also supported by the evidence of individual parts of the skull of Xiphiorhynchus , a relative of the swordfish , the longest piece measures about 33 cm. The genus Lates is in turn handed down with several skulls, they represent the oldest evidence of this representative of the giant perch in Africa. The same statement applies to Parachanna , a representative of the snakehead fish , from which individual skull parts are also handed down from the Fayyum. Cichlids , African tetras and lungfish also occur .
Amphibians have hardly been studied to date. Remains of frogs have been reported from the Gebel-Qatrani Formation .
reptiles and birds
The reptiles , on the other hand, are very diverse and are represented by all the orders that still exist today. As a rule, the remains are in the Qasr el-Sgha and in the Gebel-Qatrani Formation, only rarely in older rock strata. The extensive material of the turtles consists mainly of the shell remains, but there are also skulls and elements of the body skeleton. There are representatives from both of today's orders. Gigantochersina , a tortoise of the size of the Galapagos tortoises , can be referred to the Halsberger tortoise group . Several complete carapaces of up to 88 cm in length have survived from this genus. Initially, the finds were attributed to the Palaearctic tortoises . Among the turner -necked turtles , the genera Cordichelys , Dacquemys , Albertwoodemys , Shetwemys and Stereogenys should be highlighted. All belong to the Podocnemididae family , which contains primarily freshwater forms. It is mostly small to medium-sized forms, as a large form Stereogenys has a carapace length of up to 46 cm.
There are around 100 vertebral fragments from the scaled reptiles. They include the oldest representatives of the monitor lizards . There is also a somewhat more primitive, hitherto undetermined form. The finds suggest that the entire group originated in Africa. The snakes are also predominantly documented by vertebrae, with most of the material being assigned to the Qasr el-Sagha formation. Among them, Gigantophis , a giant snake , is particularly impressive, with an average length of 6.9 m, large specimens may have reached 9 to 11 m in length. The animals probably lived underground. So far, no skull material is known, so that nothing can be said about the dimensions and elasticity of the mouth and thus about the size of the prey; other representatives of the Madtsoiidae , however, did not have the adaptation to an extreme mouth stretching as in today's boa constrictors. Pterosphernus , on the other hand, was adapted to life in water, as indicated by the laterally strongly compressed vertebrae. It also reached great proportions. In addition to these two most common representatives, there are also some smaller, as yet unidentified boas , as well as earth boas . With Renenutet , a form of the adder and viper -like also occurs, but so far only three vertebrae have survived.
Within the crocodiles , two groups of forms can be distinguished in the Fayyum area. One consists of long-snouted representatives who are phylogenetically close to today's Gaviale . As the oldest form, Paratomistoma has been described using a skull from the Gehannam Formation of Wadi el-Hitan. Their location in marine sediments suggests an adaptation to marine life. All other finds are stratigraphically younger. Of importance here is Eogavialis , whose find material was originally attributed to the recent Sunda Gavial . However, Eogavialis turned out to be very basic in the development of the Gavial, possibly belonging to a lineage before the splitting into today's Southeast Asian and the extinct South American forms. The second group is represented by animals with wide snouts, which probably show a closer relationship to real crocodiles . First assigned to today's genus Crocodylus , but it is very likely that they are phylogenetically more original members. In addition to these forms, a lower jaw fragment from a previously unnamed member of the Sebecosuchia provides one of the rare examples of this group from Africa. Together with some finds from South America, the piece from the Fayyum is also one of the most recent examples of a representative of this primitive and more distant crocodile family.
Bird fossils are sparse with isolated leg and foot bones, sometimes with skull fragments. The earliest evidence of birds in the region is a tibiotarsus from the Birket-Qarun Formation of Wadi el-Hitan. The genus Eopelecanus was established for it, and it is also one of the oldest examples of pelicans . The other finds come almost without exception from the Gebel-Qatrani Formation and are distributed over around half a dozen orders. Only a few objects can be assigned exactly. These include the genera Nupharanassa and Janipes , which are closely related to jacanas within the plover family . The wading birds , on the other hand, are represented by Xenerodiops , which was slightly smaller than today's storks and is documented by a skull and a humerus. Another skull is attributed to Palaeoephippiorhynchus , also a relative of the stork. Some other finds are in turn related to today's genus Nycticorax and thus to the herons within the copepods . Also belonging to the same family group are the shoebills , which form a rather rare fossil element. They are represented by Goliathia in the Fayyum. In addition, birds of prey (including hawks and ospreys ) and cranes (with rails ) were present, as were turacos and flamingos , but the material here is too fragmented for more precise statements. In addition to the well-known groups of birds, some traditional forms cannot be classified exactly. Mention should be made here of Eremopezus , a probably flightless bird with the dimensions of today 's rheas . This bird, equipped with long and slender legs, the tarsometatarsus alone measures a good 34 cm, did not resemble any of the known groups of large ratites. A similar form from the Fayyum area, originally named Stromeria , was even initially considered to be the basal form of the Malagasy elephant birds . Today Stromeria is understood as synonymous with Eremopezus . It is probably an independent development within the bird world of Africa. Another arguably flightless bird, indicated by a severely fragmented leg bone, may belong to the extinct Ameghinornithidae ; these have so far only been described from Europe.
mammals
The mammals are very extensive and covered by more than a dozen orders, some of which are now extinct. They are found in all areas of the Maadi group. A distinction must be made between marine and terrestrial forms. The former are predominantly found in the lower (Gehannam and Birket-Qarun Formation), the latter in the upper (Qasr-el-Sagha and Gebel-Qatrani Formation) sections. The most original group form the marsupials , of which at least two representatives exist. These include Peratherium , which represents the first marsupial ever found in Africa and is closely related to the opossum . Ghamidtherium seems to have a similar relationship . Both forms are only known from several lower jaws and individual remains of the upper jaw . However, the assignment of Ghamidtherium to the marsupials is partially questioned, other authors tend to group the remains with the insectivores . With the Ptolemaiida , an extinct order of small, probably insectivorous mammals was introduced, whose systematic position was initially unclear. Characteristic are the oversized central molars from the rear premolars to the foremost molar , while the other molars remain relatively small. At least three genera have been described from the Fayyum. The best known is the type form Ptolemaia , whose first records in the form of remains of teeth date back to the beginning of the 20th century. Later, skull finds came to light. Other forms from the Fayyum include Qarunavus and Cleopatrodon . What is remarkable about the Ptolemaiida finds in the Fayyum is that almost all of them came to light within a radius of around 1 km, so that the animals may have been ecologically very limited. Today the Ptolemaiidae are associated with the Afrotheria .
The actual Afrotheria form one of the main find groups in the fossil community of Fayyum. From the small, insect-like forms mostly dentition fragments have survived. This applies to Eochrysochloris as a primordial member of the golden mole -rats as well as to some other, more distant relatives of the tenrecs . Dilambdogale , Widanelfarasia or Qatranilestes should be mentioned here . Both the present-day golden mole-rats and the tenrec-like ones are characterized by three prominent humps and a V-shaped claw on each molar ( zalambdodontes tooth pattern), but the latter three fossil genera have a more primitive W-shaped claw on the front molars ( dilambdodontes tooth pattern). Herodotius , on the other hand, can be regarded as a very early representative of the elephant shrews . The genus is based on several skull and lower jaw fragments. Another lower jaw and a tooth are known from Metoldobotes , a much more modern and larger form.
The fossil record of hyraxes in the Fayyum is outstanding; in the Gebel-Qatrani Formation alone they account for almost a third of all mammalian finds. The hyrax are among the most important biostratigraphic elements of Palaeogene Africa. Skull and body elements survive from Dimaitherium . With its still elongated skull, it represents an archaic hyrax, the structure of its foot indicates a possible climbing locomotion with rapid but not sustained movements. Other early forms are Geniohyus , Bunohyrax or Pachyhyrax , they often differ only in the modifications of the molars from a humped ( bunodont ) chewing surface pattern to one with a crescent-shaped scissors ( selenodont ). Saghatherium and Thyrohyrax appear much more modern , which is indicated by the shortened skull. The former possibly represented a fast runner. Both representatives have chamber-like cavities on the inside of the lower jaw, which only occur in male animals, but their function is unclear. With Antilohyrax , the hind foot configuration suggests a jumping animal. Most of the hyraxes mentioned here were relatively small and only slightly exceeded today's species. In contrast, Megalohyrax reached the size of a donkey, while Titanohyrax was estimated to weigh over 800 kg and thus had the proportions of a small rhinoceros. From both mainly remains of skulls and teeth are preserved.
Arsinoitherium is one of the best-known and probably also most characteristic fossil forms of the entire Fayyum region , a huge animal with a body length of up to 3.4 m. Almost the entire skeleton of the animal is known from several dozen individuals. Its main features can be found in the two pairs of bony horns on the skull, of which the larger front one was formed from the nasal bone. Equally striking is the dentition with 44 high-crowned teeth, with which the probably soft plant food was crushed. Externally, Arsinoitherium resembled the rhinos, with which it is not closely related to the skeletal structure, it is referred to its own extinct group of Embrithopoda . The Fayyum region gained exceptional importance through the fossils of proboscidea . They all belong to a very early phase of development, the common feature of which is the vertical change of teeth. Moeritherium probably formed a side branch within this. It was outwardly pig-like and did not yet have a trunk. In contrast, the much larger Barytherium looked more like a classic trunked animal, but it had a total of eight small tusks. Both forms were pronounced swamp dwellers. Two other genera attested are Palaeomastodon and Phiomia . Both appear somewhat more modern due to their front molars with three transverse ridges compared to two in the aforementioned representatives, and the tusks are longer. It is partly believed that the former is closer to the mammothid lineage , the latter to gomphotheres and elephants . The manatees are stratigraphically broad. Protosiren appears as early as the Gehannam Formation with some well-preserved skeletons. Its very early branch within the sea cows can still be recognized by the clearly developed front and rear extremities and the strong spinous processes of the thoracic vertebrae, which probably allowed a semi-aquatic life. The other genera, Eosiren and Eotheroides , are both phylogenetically and stratigraphically younger and closer to present-day dugongs , having a much reduced limb skeleton and more swollen bones. The up to 2.5 m long Eosiren may not have lived purely in the sea, but was also found upstream. The various manatees of the Fayyum fed on sea grasses, probably occupying different ecological niches.
In addition to the Afrotheria with the manatees, sea-dwelling mammals also emerged within the Laurasiatheria with the whales . They are very richly documented in the Fayyum, with some complete skeletons. Various remnants already appear in the Mokattam group, whales are found very frequently in the Gehannam and Birket-Qarun Formation, but in the Qasr-el-Sagha Formation their proportion decreases again. A good 500 skeletons are known from Wadi el-Hitan alone, which allow the size and individual development of the animals to be studied. Other finds came to light in the outcrops north and south of Lake Qarun. The forms described so far from the Fayyum area belong to an original cetacean group called Archaeoceti . These differ from modern whales in having a full dentition, teeth with numerous small bumps, less specialized flippers , and hind legs that still had feet and toes. The evidence of the fully developed hind limbs could be provided for the first time using the Fayyum fossils. In line with today's whales, the lumbar spine was already extremely elongated. The oldest finds come from the Midawara Formation in Wadi el-Rayyan . One example of this is the hind limbs of Rayanistes , a member of the Remingtonocetidae . These very primitive whales have so far been largely documented from the Indian subcontinent . The rock unit also contained a partial skeleton of Phiomicetus , which in turn can be assigned to the Protocetidae , another very original family group of whales. It also includes Aegicetus , which is available with two individuals from the Gehannam Formation of Wadi el-Hitan. The animals, which weighed just under 900 kg, had hands and feet that were relatively the same size. As a result, they moved less undulating in the water, but used their limbs even more for propulsion. In addition, there are weaker flattened caudal vertebrae, so that possibly no fluke was formed. Likewise, Aegicetus possessed a still fully developed sternum, deviating from later whales. The other whales from the Fayyum area belong to the Basilosauridae , the best known and most common representatives here are Basilosaurus , a 18 m long giant, and the up to 5 m long Dorudon . Basilosaurus stomach remains from Wadi el-Hitan showed that the giant toothed whale hunted its smaller relative Dorudon and preyed on its offspring. The extinct bony fish Pycnodus was found to be another prey animal . Thus, Basilosaurus probably took the position of an apex predator in the seas of the late Eocene . Dorudon , on the other hand, probably fed mainly on fish and used the then shallow sea waters to give birth to their offspring, as individual finds of young animals suggest. It had a weight of around 2.2 t, a small brain weighing only 980 g and, according to the structure of the skull, was not yet capable of echolocation . Ancalecetus reached about the same dimensions as Dorudon , but it showed a differently configured front leg. Saghacetus is quite small with a body length of 3 m and is mainly found in the Qasr el Sagha Formation. Stromerius and Masracetus appear rather rarely , from the latter individual teeth and parts of the body skeleton were found in the Gehannam Formation north of Lake Qarun. The closest relatives of the whales can be found in the even-toed ungulates , of these only representatives of the Anthracotheriidae are documented in the Fayyum , which are in the evolutionary lineage to the hippos . The externally pig-like animals were probably also adapted to a semi-aquatic life due to their physique. While only a few finds are available from Qatraniodon and Nabotherium , the fossil material from Bothriogenys consists of more than 2000 tooth and bone elements from animals of all ages.
The Hyaenodonta replaced the predators in Africa in the Paleogene and were the exclusive terrestrial predators there . Some of them are placed in the more comprehensive but not self-contained group of " Creodonta ". In general, the Afro-Arabian Hyaenodonta are little studied, previous work mostly compared them to forms such as Pterodon or Apterodon that were already known from other continents. The find material from the Fayyum consists largely of skulls and mandibles, postcranial body parts are scarce. Among other things, Masrasector from the Fayyum was described, which occupied the niche of the small, ground-dwelling predators, its body weight was around 1 kg. In contrast, Brychotherium became six times as heavy, both surpassing Akhnatenavus , who weighed about 20 kg and more. The latter was characterized by an extremely developed crushing scissors . Another proven representative is Metapterodon . Two indented terminal phalanges of the fore and hind feet indicate pangolins , a more precise determination has not yet taken place. With a length of 2.7 or 1.1 cm, they are in the size range of today's species. Bats are a rather rare element among Fayyum fossils. Their evidence is based on a little more than three dozen individual finds, which mainly include dentition remains and isolated teeth. The genera Philisis , Witwatia , and possibly also Vampyravus belong to an extinct branch of the order. In contrast to the other bats, Vampyravus is only known for a 4.9 cm long humerus. Presented in the early 20th century, this bone was the first reference to extinct bats in the Fayyum (and in Africa). In addition, Vampyravus weighs around 120 g, surpassing most other fossil bats. Only Aegyptonycteris had a similar size, this is a large, predatory animal. Other bats can be associated with lineages existing today. Dhofarella is related to the smooth-nosed free- tailed bats , Qarunycteris to the mouse -tailed bats and Saharaderma to the large- leaf bats . The latter and also Khonsunycteris , a smooth -nosed bat , can be considered the smallest bats from the Fayyum with a body weight of about 30 g. Phasmatonycteris is also noteworthy , since the closest relatives of the genus can only be found in Madagascar today with the Madagascan adhesive disc bats .
The superorder of the Euarchontoglires is extremely diverse and accounts for more than 50% of the finds in the Gebel-Qatrani Formation alone. The greater part falls on the rodents . Here again the porcupine relatives dominate , characterized by the hystricognathic lower jaw. The rodent group combines an extensive skull and dentition material. Phiomys was already defined in a lower jaw at the beginning of the 20th century and has been documented many times. More closely related are Protophiomys , Acritophiomys and Talahphiomys , but also Qatranimys , Waslamys or Gaudeamus . Within the porcupine relatives, all forms mentioned form mostly extinct lines. However, Birkamys and Mubhammys could belong in the cane rat lineage , as could Monamys . Another special group is found in the spiny -tailed squirrel relatives , which so far only occur in the lower reaches of the Qasr-el-Sagha Formation. You can only show a little skull and tooth material, detected genera are Shazurus and Kabirmys . The former is relatively small, but resembles the real thorntail squirrel in terms of tooth structure . The latter represents the largest known form of the Paleogene thorntail squirrel relatives.
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The primates , which form one of the ancestral mammal groups of the African continent alongside the Afrotheria , also achieve a high level of diversity . Lori -like forms occur, among others, with Karanisia and Saharagalago , as well as with the dwarfish Wadilemur , of which parts of the body skeleton are known in addition to teeth remains. The latter two could belong to the root group of the Galagos , while the former belonged to the Loris . Plesiopithecus , defined on the basis of a lower jaw and later supplemented by additional finds such as a partial skull, again joins the continental ancestor forms of the aye- aye from Madagascar. Another rather original and extinct lineage of primates was established with the Adapiformes . Their position within the wet-nosed or within the dry-nosed primates is sometimes disputed. Afradapis has been described from the Fayyum , whose molars with high cusps and long shearing edges indicate a leaf-eating diet. Masradapis , which weighed around 900 g, may in turn have consumed a higher proportion of seeds and fruit due to its larger rear molars and more powerful lower jaw. About the same size was the related Aframonius , of whom not only parts of the teeth but also parts of the skull have been handed down. The small size of its eye socket speaks for a diurnal primate. Other forms represent the apes and thus the higher primates. The oldest representative is Biretia , it already occurs in the lower section of the Qasr-el-Sagha Formation. Based on the few teeth and remains of a skull that have been found so far, an animal weighing a little more than 200 g can be reconstructed, which, unlike most of today's apes, lived a nocturnal lifestyle. In contrast, all other monkeys have so far only been found in the Gebel-Qatrani Formation. Various lines of development can be distinguished. For example, the genera Apidium , Parapithecus , Proteopithecus or Qatrania belong to the original branch lines of the monkeys, with Apidium in particular being very common. The animals fed mainly on fruit and, judging by their body skeleton, moved through the trees in an apparently jumping manner, although some studies estimate their agility to be less great. Other forms probably belong to the ancestral group of Old World monkeys , such as Catopithecus and Oligopithecus as well as Propliopithecus and Aegyptopithecus . The former two are included in the kinship of the Oligopithecidae , the latter two in that of the Propliopithecidae . The Oligopithecidae occur mainly in stratigraphically older, the Propliopithecidae in younger sections of the Gebel-Qatrani Formation and thus largely exclude each other. However, a single lower jaw of a probably dwarfed, but not yet exactly identified oligopithecid was found in a much higher position (site M ) and possibly represents one of the last records of this primate group. The find appeared here together with various representatives of the Propliopithecid. Of particular note, however , is Aegyptopithecus , which has some good skull finds. This has revealed considerable sexual dimorphism , which is among the earliest evidence in higher primates. The brain volume of the monkey form was about 20 cc. Two other primates, Nosmips and Afrotarsius , are unclear in their phylogenetic position. Only a few teeth and dentition fragments are present from both, showing mixed characteristics of more primordial and higher primates.
trace fossils
The Fayyum region is characterized by excellently preserved trace fossils , which can occur in all stratigraphic units, but have been handed down in greater abundance in the soil formations of the continental Gebel-Qatrani Formation. More than two dozen different forms are known. Among them, the life traces of invertebrates dominate. These are usually tunnels and tubes or spherical formations that are interpreted as feeding or burrowing passages for insects , worms , molluscs or crabs . Among other things, however, there are also complex structures with a diameter of up to 65 cm, which often consist of a wide variety of corridors, chambers and galleries. They represent fossilized nests of termites , which are assigned to the trace genus Termitichnus . Others include multi-tiered tunnels that are of Masrichnus origin and may be due to bees . Significantly larger structures also occur, some of which are spiral or U-shaped passages of up to 1.5 m in length and 20 cm in diameter. Their origin is likely to be due to vertebrates, with mammals that burrow sometimes being the cause of the complex tunnel systems. A third large group of trace fossils can be referred to as rhizoliths (root ducts of plants). They come in all sizes and reflect the diverse vegetation at that time. Among other things, traces of the buttress roots of trees are documented, which probably suggest mangroves . Some of these root structures are 10 feet (3.2 m) in diameter, so trees of this size can be assumed.
age position
Investigations into the age of the outcrops in the Fayyum and their fossils are based on three different approaches. As early as the late 19th century , a classification into the Upper Eocene and the Lower Oligocene was assumed for the fossils of the Fayyum from biostratigraphical considerations, for example in the composition of the shark fauna . The mollusc community also provided similar results , so that at the turn of the 19th and 20th centuries there was little doubt that the age was classified as “early Tertiary” ( Palaeogene ). The biostratigraphy, with its relative chronological view, was subsequently refined and a faunal succession from the lower to the upper sections was worked out. Invertebrates such as mollusks and foraminifera proved to be good index fossils , especially for the marine deposits . For the latter, an age classification in the Middle Eocene, more precisely in the Lutetian , could be made for the lower sections of the Mokattam group. The upper depositional sequences of the group beginning with the Gharaq Formation belong to the Bartonian 41 to 38 million years ago. The transition to the Priabonian and thus to the Upper Eocene 38 to 34 million years ago is found in the Upper Gehannam Formation and is indicated by the change from Truncorotaloides to Globigerinatheca and the appearance of Turborotalia . However, there is a certain spread here, so that no exact stratigraphic position can be given. An age estimation for the upper terrestrial sequence has been made several times with the help of mammals. However, certain difficulties arise here, since the mammals appear to be highly endemic in large parts . They are composed of originally African forms ( Afrotheria ) and primarily Eurasian representatives ( Laurasiatheria and Euarchontoglires ). The position of the Gebel-Qatrani Formation proved to be particularly problematic. Depending on the study, it either belonged entirely to the Upper Eocene or the Lower Oligocene or dated to the transition between the two periods. However, a relative age could be derived from individual groups of fossils. Thus, the primates of the Fayyum often seem more original than the phylogenetically younger forms of the Lower Miocene in East Africa. Because of the peculiarity of the terrestrial mammalian fauna in the Fayyum region, the "Phiomian" was proposed in 1991 by John A. Van Couvering and Judith A. Harris as a designation for a regional stratigraphic stage.
The new dating methods developed over the course of the 20th century allowed, among other things, absolute dating approaches . Already in the 1960s, the first data for the overlying Widan-el-Faras basalt on the Eocene and Oligocene sedimentary sequence were published, which were based on the potassium-argon method . At that time they gave an age of 24.7 to 27 million years, which corresponds to the Upper Oligocene. The values could thus be regarded as the minimum age for the overlying sediment layers. Further radiometric analyzes from the 1980s assigned a slightly older age of 31 million years to the lowest section of the Widan-el-Faras basalt. However, the result later turned out to be inaccurate. Measurements of the same areas taken in the 1990s, combined with verification of previously obtained age estimates, confirmed the Upper Oligocene position for the basalt, the new values being around 23.6 million years.
Direct dating of the marine and fluviatile-lacustrine fossil-bearing sedimentary rocks of the Fayyum has not yet been possible. However, the radiometric surveys of the 1990s were associated with measurements of paleomagnetism . The sequences analyzed concerned the upper section of the Qasr el Sagha Formation ( Dir Abu Lifa Memeber ) and the entire Gebel Qatrani Formation. Within the sequence of deposits, a multiple change in the polarity of the earth's magnetic field could be demonstrated, which the authors associated with the magnetostratigraphic sections Chron 16 to Chron 12 . The sections corresponded to a (then) age classification of around 37 to 33 million years. As a result of the work, it could be assumed that the Fayyum deposit series belonged almost entirely to the Eocene, only the uppermost section of the Gebel-Qatrani Formation fell into the Lower Oligocene transition. Subsequent analyzes recorded a much more extensive sedimentary sequence beginning in the basal areas of the Qasr el Sagha Formation and also carried out a comparison with other localities in the near and far vicinity. They corrected and specified the results of the first measurements. From the point of view of magnetostratigraphy, the depositional sequence ranges from Chron 17 (Qasr-el-Sagha Formation) to Chron 10 (Gebel-Qatrani Formation), the corresponding age dates are 38 to 28.5 million years. The Eocene-Oligocene transition occurs in Chron 13 , which occupies the lower part of the Gebel-Qatrani Formation. Accordingly, the far greater part of the rock unit is of Oligocene origin, only the lowest 48 m were already formed in the Upper Eocene. The important site L-41 is located exactly in this transition area, its age dates to around 34 million years. The stratigraphically younger sites I and M are embedded in Chron 11 and are therefore about 29.5 million years old. Significantly older is the classification of site BQ-2 , which contains some of the earliest terrestrial fossil finds and is located in the lower section of the Qasr el Sagha Formation. It belongs to Chron 17 with an absolute age value of about 37 million years.
landscape reconstruction
Today the Fayyum region lies in the middle of the Sahara and is characterized by a desert climate with an average annual temperature of around 22 °C with absolute maximum values of around 49 °C and minimum values of −1 °C. The annual precipitation is only 10 mm and is largely limited to the winter months, while the evaporation rate can reach more than 200 times.
In the Paleogene , Africa formed a continental island together with the Arabian Peninsula . This Afro-Arabian landmass was separated from the Eurasian continent by the Tethys Ocean , a land bridge formed only in the transition to the Miocene around 24 million years ago with the gradual closure of the Tethys. The largely limestone formations of the Mokattam Group are marine and shallow marine in origin and consequently deposited within the Tethys during the Middle Eocene. The same applies to parts of the Maadi group, but the influence of the proximity to the coast on the southern edge of the Tethys is already noticeable here. The Birket-Qarun Formation and the lower sections of the Qasr-el-Sagha Formation ( Umm Rigl Member to Temple Member ) indicate the further retreat of the seawater to the north, resulting in a bay -like , later lagoon-like landscape in the course of the Upper Eocene formed. On the other hand, the coarser clastic deposits of the Dir Abu Lifa member , which are layered on the shallow marine sediments, were formed under the conditions of a river delta or an estuary , which increasingly indicates terrestrial conditions.
The Gebel-Qatrani Formation caps the underlying strata. It goes back to a coastal swamp or flooded area that existed in the transition from the Eocene to the Oligocene in the form of a mangrove or terra firma landscape . The area was traversed by several rivers that were relatively wide and powerful enough to move larger rock components. Forests grew on the river banks, which also reached into the hinterland. The relative proximity to the sea had a certain influence, so that brackish water conditions prevailed in some areas. This landscape supported a rich and varied fauna, composed of terrestrial and aquatic animals, the latter consisting of freshwater, saltwater and brackish water forms. According to the further paleontological and geological data, a tropical to subtropical climate prevailed during the formation of the Gebel-Qatrani Formation in the Lower Oligocene . The formation of the paleosoils shows an alternation of wet and dry periods, which may be due to the influence of a monsoon .
Comparison with regionally and nationally important sites
The Fayyum region represents one of the most important Late Palaeogene fossil sites in Africa and the rest of the world. It is also one of the few sites on the African continent that cover the Late Eocene and Lower Oligocene periods. In contrast, older sites in northern Africa are comparatively more common. One of the earliest Cenozoic terrestrial fossil assemblages was discovered in the Ouled-Abdoun Basin in Morocco and dates to the Paleocene and the Early Eocene. Older layers containing finds reach back as far as the Maastrichtian , so that a find period from 72 million years ago to about 48 million years ago is recorded. The basin is known, among other things, for its phosphate richness. The upper part of the sequence contains very primordial forms of mammals, such as Ocepeia and Abdounodus , two forms close to present-day Afrotheria. With Eritherium and Phosphatherium , two of the earliest known proboscis animals also appear, and Stylolophus , an original member of the Embrithopoda, was also discovered. El-Kohol in northern Algeria can be considered comparable to the uppermost section of the Ouled-Abdoun Basin . Marsupials, hyaenodonts, hyraxes and proboscidea were found here, among other things. The mammalian fauna is much closer to that of Fayyum, but still appears comparatively archaic with the hyrax Seggeurius and the proboscidea Numidotherium . The Lower to Middle Eocene site complex Gour Lazib , western Algeria, assumes a stronger intermediary role, since here forms already occur with Megalohyrax and Titanohyrax that are also known from Fayyum. The very early primate Azibius and Helioseus , a mammal whose relationship is not exactly known, are documented as special features. The material is mostly very fragmented, which limits the possible statements. Chambi in Tunisia also shows connections to Fayyum, in addition to hyraxes and primates, some of the oldest elephant shrews have also been documented here. Also of importance is the collection of finds from the Ouarzazate Basin in Morocco , which largely consists only of teeth . The numerous sites of the basin scatter chronologically from the late Paleocene to the Middle Eocene. Older localities such as Adrar Mgorn and N'Tagourt yielded ancestral insectivorous mammals such as Afrodon or Todralestes , which may correspond to an African endemic fauna. In addition, there is a primate, Altiatlasius , who is probably on the threshold of the development of the apes, and Tinerhodon , a representative of the Hyaenodonta. The collection at the most recent site, Aznag, is made up of shrew-like insectivores, bats, elephant shrews and primitive ungulates. Aznag is the only clearly Middle Eocene locality with land mammals from northern Africa.
The Nementcha site (also Bir el-Ater) in northern Algeria dates from the Upper Eocene, and is therefore contemporaneous with the Qasr el-Sagha Formation. This is indicated by Moeritherium and Bunohyrax , for example, and the first anthracotheres could also be detected as Eurasian immigrants in Africa. Other small mammals include rodents of the Phiomyidae family and elephant shrews. The finds from Ad-Dakhla in Western Sahara are of a comparable age . The fossils stored in the Guerrani Member of the Samlat Formation are composed of numerous fish remains, turtles, crocodiles, isolated birds and marine and land-dwelling mammals. Among the marine mammals, remains of the whales Saghacetus , Basilosaurus and Dorudon and the manatee Eosiren stand out. Among other things, the tooth of a proboscidea was discovered as a terrestrial mammal, which may correspond to Numidotherium .
The sites Dor el-Talha and Jebel al-Hasawnah , both in Libya, form a chronological equivalent to the Gebel-Qatrani Formation in the Fayyum. A comparable composition of the faunal community unites all three find areas. Among the large mammals of Dor el-Talha, for example , Barytherium , Moeritherium and Palaeomastodon can be found, as well as Arsinoitherium , and among the reptiles, for example, gavial-like forms stand out. Also of note are early primates such as Karanisia and early monkeys such as Biretia or Talahpithecus , as well as early elephant-shrews such as Eotmantsoius . In Jebel al-Hasawnah, among other things, Saghatherium and Titanohyrax could be proven. The site is also outstanding for taphonomic reasons, as it is the only fossil site in Palaeogene Africa to date that has produced skeletons of land-living mammals in an anatomical context. The site complex of Zallah in the Sirte Basin of central Libya is in a similar chronological context . Fossils have been recovered here since the 1960s, but are largely limited to teeth. The rodent fauna is very numerous with remains of various porcupine relatives such as Gaudeamus , Metaphiomys or Neophiomys as well as thorntail squirrel relatives such as Kabirmys . In addition, hyraxes with Thyrohyrax and representatives of the Anthracotheriidae with Bothriogenys have also been detected. Africtis , in turn, represents the earliest known African form from the family environment of the carnivores, which has its origin in Eurasia. The fossil deposit of Minqar Tibaghbagh in the southwest of the Qattara Depression , discovered only in 2007, also corresponds to the Gebel Qatrani Formation with its banded sandstones. The Oligocene deposits contain not only remains of sharks, turtles and crocodiles, but also of terrestrial mammals such as Phiomia , Antilohyrax and Bothriogenys . An older find horizon from the Upper Eocene also contained fossils of whales and manatees. The sites of Thaytiniti and Taqah in Oman can also be mentioned from the Arabian Peninsula , which contain fauna that is about the same time as the upper sections of Fayyum.
Comparatively few sites in Africa are occupied from the Upper Oligocene. One of the most significant is Chilga in Ethiopia, which is about 27 million years old. The large mammals include proboscidea as well as hyrax and Embrithopoda. They show a mixture of both older forms, such as those found in the Fayyum, such as Arsinoitherium , Megalohyrax or Palaeomastodon , but also more modern elements. For example, Chilgatherium and Gomphotherium are added to the trunk animals, both of which belong to phylogenetically younger lines such as Deinotherien and Gomphotherien. Several Upper Oligocene fossiliferous ranges are found along the Great Rift Valley, extending from northwestern Kenya to southwestern Tanzania. Only the Eragaleit Beds west of Lake Turkana should be mentioned here as an example . Here, too, there is a faunal mixture of older and younger elements. In addition to some forms that have been known since the Fayyum ( Arsinoitherium , Thyrohyrax ), Losodokodon has been proven here as a basal member of the mammothid lineage and Kamoyapithecus , a close relative of Proconsul and thus of the Miocene humanoids predators before.
research history
The beginnings up to the 19th century
It can be assumed that the Fayyum depression was already known as a fossil-bearing site in ancient Egyptian times. Quarries from the upper areas of the Qasr-el-Sagha Formation are documented, in which the numerous gypsum found was mined and processed into vessels as early as the Old Kingdom . The quarries are located in close proximity to fossilized tree trunk sites. At the same time, the Widan-el-Faras basalt served as a raw material store for the lining of mortuary temples. The region was also considered the origin of life for the people of that time. The goddess Isis is said to have buried the bones of her deceased husband Osiris here.
The first fossil finds of modern times go back to Arthur Bedford Orlebar (1810-1866), who found some sintered tree stumps up to 20 m long in sandstone layers in 1845 . Additional finds are due to the German geologist Georg Schweinfurth (1836–1925), who was an experienced explorer of Africa. In 1879 he discovered teeth and bones of sharks and bones of whales on the island of Geziret el-Qorn in Lake Qarun, in addition to numerous molluscs . In general, the finds are now assigned to the Birket-Qarun Formation . The vertebrate remains were processed in 1883 by Wilhelm B. Dames , who recognized, among other things, two species of Zeuglodon (today mostly Basilosaurus ). In the same year, the Swiss geologist Karl Mayer-Eymar took on the mollusks. Schweinfurth returned to Fayyum in the mid-1880s and surveyed the areas north of Lake Qarun. He discovered not only the temple ruins of Qasr el-Sagha (also called "Schweinfurth's Temple"), but other vertebrate remains. These were in turn scientifically evaluated by Dames, among other things he described the new whale species, Zeuglodon osiris ( Saghacetus ). Schweinfurth published his own report about his trip in 1886. In the 1890s, Mayer-Eymar stayed in the Fayyum for short periods of time.
From the 19th to the 20th century - a great period of research
The transition from the 19th to the 20th century coincided with an intensive research phase in the Fayyum, in which a wide variety of groups became active. The British paleontologist Hugh John Llewellyn Beadnell (1874–1944) started the process. From 1898 he mapped the northern and eastern borders of the basin on behalf of the Geological Survey of Egypt. The aim was to create an irrigation system for the agricultural use of the basin. During his work, Beadnell discovered numerous fossils of fish and crocodiles, but also of whales and manatees. He sent these to the Natural History Museum in London, where they aroused the interest of Charles William Andrews (1866–1924). From 1901 Andrews joined Beadnell's field investigations. Together they explored the northern part of the Fayyum Basin, which lasted until 1904. Their investigations concerned both the stratigraphically older and lower-lying (Birket-Qarun and Qasr-el-Sagha Formation) as well as the younger and higher outcrops (Gebel-Qatrani Formation, mainly the lower zone) north of Lake Qarun. Her most important discoveries from the former include the giant snake Gigantophis , as well as Barytherium and Moeritherium , two proboscidea, and the manatee Eosiren . Arsinoitherium and Phiomia emerged in the latter , as did original hyrax, such as Saghatherium . In 1902, Beadnell also discovered Wadi el-Hitan . Both Andrews and Beadnell published the first results of their work in short essays, usually combined with the first scientific descriptions of new species and genera. These appeared frequently in Geological magazine , but also elsewhere. In addition, Beadnell gave a brief overview of his geological work, which was followed in 1905 by a detailed monograph. Here he first named the Wadi el-Hitan with the Zeuglodon valley . A year later, Andrews published a comprehensive catalog of the vertebrate finds under the title A descriptive catalog of the Tertiary Vertebrata of the Fayum, Egypt , which is still one of the standard works on the Fayyum fossils today. He made his last trip to the Fayyum Basin in the spring of the same year.
Research activities by German scientists also took place during the same period. Max Blanckenhorn (1861–1947) presented his geological photographs as early as 1900 after a short stay in the Fayyum. Two years later he visited the fossil sites of the basin together with Ernst Stromer (1871-1952). The material they collected was published relatively early. They were followed by the Austrian fossil collector Richard Markgraf (1869–1916), who had left his homeland for Egypt for health reasons. Markgraf met the German researcher Eberhard Fraas (1862–1915) there in 1897. Fraas himself planned further stays in eastern and southern Africa, but could not realize this for the time being. Instead, he returned to Stuttgart and instructed Margrave in the search for fossils by telex. In return, Margrave sent him his fossils, which he first collected at Gebel Mokattam east of Cairo. In 1903, Markgraf met Stromer and joined his three-month expedition to the Fayyum Basin. Both explored the northern parts of the Depression, which was also the focus of British scientists. After that, Markgraf worked for Fraas again. In turn, he organized a trip to Egypt in 1906 to join forces with Margrave. The expedition finally took place after a series of difficulties, starting March 11 for 10 days. The Qasr el Sagha Formation and the Gebel Qatrani Formation were the focus of their attention. Among other things, they discovered finds of Arsinoitherium and Basilosaurus , but also of crocodiles. In 1911 , Max Schlosser published an extensive publication on the activities of German researchers in the Fayyum Basin . In it he described Propliopithecus and Parapithecus , the first unequivocal primates from the area where they were found. Stromer briefly returned to the Fayyum on his third Egyptian expedition in 1914, but the main focus of the trip was on the Bahariyya oasis . In the following two decades, numerous specialist articles on Stromer's research appeared in the treatises of the Bavarian Academy of Sciences . Margrave remained in Egypt until his death in 1916 and collected regularly in the Fayyum Basin, he offered his finds to various scientific institutes.
Parallel to Markgraf's field work and after the appearance of the Beadnell and Andrews publications, the American Museum of Natural History organized an expedition to the Fayyum Basin under the leadership of Henry Fairfield Osborn (1857–1935). This is the museum's first foreign expedition, which was to be followed by a few more, very successful ones, including to Mongolia . The expedition was supported by then US President Theodore Roosevelt . The Americans arrived there in early February 1907 and stayed until the end of May of that year. Walter W. Granger (1872-1942) and George Olsen served as Osborn's assistants . Osborn himself only stayed in Egypt for the beginning of the expedition, but he made a short detour into the Wadi el-Hitan, which he named, according to Beadnell, the Zeuglodon valley . The remaining members of the expedition worked in various outcrops and met Margrave several times during this time. The Americans hired him to look for fossils, but he worked independently. Much of the exploration was on the Gebel-Qatrani Formation and continued work on the lower zone discoveries previously explored by Andrews and Beadnell. However, the researchers also discovered finds in the upper zone for the first time. Fossils found include remains of Arsinoitherium , several proboscidea forms, hyaenodonts, rodents, and early hyraxes, among others. The entire find material, about 550 individual objects, was shipped to America. In the aftermath of the expedition, Osborn published several articles about the fossil finds, as well as the expedition itself. The full account, which includes Granger's journal, was not published until 2002.
Regardless of the previous investigations, the Muséum national d'histoire naturelle of Paris under the organizational direction of Marcellin Boule and Jean Albert Gaudry carried out a two-week expedition to the Fayyum in March/April 1904, which was financially supported by Edmond de Rothschild . Also involved was René Fourtau , a French engineer who had lived in Egypt since 1888 and was interested in geology. Among other things, Fourtau analyzed fossil invertebrates, mainly sea urchins , for the Egyptian Geological Survey . The work took place in the Gebel-Qatrani formation. As a result, the French brought around 80 vertebrate fossils to Paris, including remains of Titanohyrax , a giant hyrax. In contrast to the other research projects, those of the Muséum national d'histoire naturelle were hardly processed, so that the 1904 expedition was almost forgotten.
intermezzo
In the following, almost five decades after the predominantly British, German and American research on site, hardly any activities took place in the Fayyum. An exception is a pan-African expedition from the University of California in 1947 under Wendell Phillips . Several scientists dug in previously opened outcrops. Due to the limited time, the find material is not very extensive, but consists of proboscidea, hyaenodonts, anthracotheres and embrithopods. It now resides in Berkeley . A skull of a primeval whale was also discovered three years later.
Modern research
In the 1950s, Elwyn L. Simons (1930–2016) described the cranial bone of a Fayyum primate held in the holdings of the American Museum of Natural History. He then initiated an initiative for further field research in the Fayyum, which should, among other things, investigate the question of the origin of the closest human relatives. In cooperation with Yale University and later Duke University with the Geological Survey of Egypt and the Geological Museum of Cairo, the work on site began in 1961 and ended for the time being in 1967. In addition to the early primates, the focus was mainly on small vertebrates, which had previously received less attention had been. The annual fieldwork led, among other things, to the description of Oligopithecus and Aegyptopithecus , two primate forms, and Phiocricetomys , a rodent. In addition, there were also studies on stratigraphy and age. A declared goal was, for example, to document the exact position of the older outcrops. In 1968 Simons published an overview of the geological and paleontological work.
In 1977 a second phase of investigations in the Fayyum started, again under the overall direction of Simons. It lasted until 2005 and thus over a period of 28 years. The second phase was characterized by an increasingly interdisciplinary way of working. Numerous internationally renowned scientists were involved, each with their own research focus. These included Philip D. Gingerich (marine mammals), David Tab Rasmussen (birds, hyraxes) and Erik R. Seiffert (primates). In addition, other fields of work were increasingly in focus, such as the palaeoenvironment, palaeomagnetics or soil science. In addition, new fossil groups such as trace fossils have also been included. Another focus was the training of young scientists. Advances in excavation methods have significantly increased the find yield. In the late 1970s alone, more than 1,400 remains of mammals were recovered. The work on site then continued, the processing of the find material continues to this day, whereby new species and genera from the Fayyum Basin are described almost every year.
lost and found safekeeping
To date, tens of thousands of fossils have been found in the Fayyum, most notably the Egyptian Geological Museum in Cairo, the Peabody Museum of Natural History at Yale University , the Duke Lemur Center at Duke University , the American Museum of Natural History in New York , in the Natural History Museum in London, in the Muséum national d'histoire naturelle in Paris, in the State Museum of Natural History in Stuttgart and in the Bavarian State Collection for Paleontology and Geology in Munich.
literature
- Charles W. Andrews: A descriptive catalog of the Tertiary Vertebrata of the Fayum, Egypt. London 1906, pp. 1-324
- Vincent L. Morgan and Spencer G. Lucas: Notes From Diary–Fayum Trip, 1907 (based on the expedition diary and photographs of Walter Granger). Bulletin of the New Mexico Museum of Natural History and Science 22, 2002, pp. 1-148
- Elwyn L. Simons: Eocene and Oligocene mammals of the Fayum, Egypt. In: First International Conference on the Geology of the Tethys, Cairo University, November, 2005, Volume II. Cairo 2005, pp. 439–450
itemizations
- ↑ a b c d Hugh John Llewellyn Beadnell: The Topography and Geology of the Fayum Province of Egypt. Cairo, 1905, pp. 1-101
- ↑ a b c d e f g h i Thomas M. Bown and Mary J. Kraus: Geology and Paleoenvironment of the Oligocene Jebel Qatrani Formation and Adjacent Rocks, Fayum Depression, Egypt. US Geological Survey Professional Paper 1452, 1988, pp. 1-60
- ↑ a b c d e f g h i j k l m Philip D. Gingerich: Marine mammals (Cetacea and Sirenia) from the Eocene of Gebel Mokattam and Fayum, Egypt: Stratigraphy, age, and paleoenvironments. Papers on Paleontology 30, 1992, pp. 1-84
- ↑ a b c Gamal M. El-Shabrawy and Henri J. Dumont: The Fayum Depression and Its Lakes. In: Henri J. Dumont (ed.): The Nile. Origin, Environments, Limnology and Human Use. Monographiae Biologicae, Springer, 2009, pp. 95–124
- ↑ a b c d e f g h i j k l m Chris King, Charlie Underwood and Etienne Steurbaut: Eocene stratigraphy of the Wadi Al-Hitan World Heritage Site and adjacent areas (Fayum, Egypt). Stratigraphy 11 (3-4), 2014, pp. 185-234
- ↑ a b Ulrich Jux and Thomas Steubner: Lake Karun in Egyptian Fayum. Its sediments as reflected in history. Naturwissenschaften 77, 1990, pp. 262-270
- ↑ a b Rushdi Said, Claude C. Albritton, Fred Wendorf, Romuald Schild and Michał Kobusiewicz: Remarks on the Holocene geology and archeology of Northern Fayum desert. Archaeologia Polona 13, 1972, pp. 7-22
- ↑ a b El-Sayed AAYoussef, MA Abdel Fattah and A. Refaat: Sequence stratigraphy of the Late Eocene/Oligocene alluvial/paralic succession in the Fayum area, Western Desert, Egypt. Bulletin of the Tethys Geological Society 1, 2006, pp. 59-70
- ↑ Timothy M Kusky, Talaat M Ramadan, Mahmoud M Hassaan, and Safwat Gabr: Structural and Tectonic Evolution of El-Faiyum Depression, North Western Desert, Egypt Based on Analysis of Landsat ETM+, and SRTM Data. Journal of Earth Science 22(1), 2011, pp. 75-100
- ↑ a b c d e Iyad S. Zalmout and Philip D. Gingerich: Late Eocene Sea Cows (Mammalia, Sirenia) from Wadi Al Hitan in the Western Desert of Fayum, Egypt. Papers on Paleontology 37, 2012, pp. 1-158
- ↑ a b c d e f M Gameil, M Al Anbaawy, M Abdel Fattah and G Abu El-Kheir: Lithofacies and biofacies characteristics and whales skeletons distribution in the Eocene rock units of Fayoum Area, Egypt. Journal of African Earth Sciences 116, 2016, pp. 42-55
- ↑ a b c d Zaki A Abdel-Fattah, Murray K Gingras, Michael W Caldwell and S George Pemberton: Sedimentary environments and depositional characteristics of the Middle to Upper Eocene whale-bearing succession in the Fayum Depression, Egypt. Sedimentology 57, 2010, pp. 446-476
- ↑ a b c d Thomas M Bown, Mary J Kraus, Scott L Wing, John G Fleagle, Bruce H Tiffney, Elwyn L Simons, and Carl F Vondra: The Fayum Primate Forest Revisited. Journal of Human Evolution 11, 1982, pp. 603-632
- ↑ a b Abdelfattah A. Zalat: Holocene diatom assemblages and their palaeoenvironmental interpretations in Fayoum depression, Western Desert, Egypt. Quaternary International 369, 2015, pp. 86-98
- ↑ a b c d e f g h Vincent L. Morgan and Spencer G. Lucas: Notes From Diary–Fayum Trip, 1907 (based on the expedition diary and photographs of Walter Granger). Bulletin of the New Mexico Museum of Natural History and Science 22, 2002, pp. 1–148 (online)
- ↑ Gunther Garbrecht: Historical water storage for irrigation in the Fayum depression (Egypt). Irrigation and Drainage Systems 10, 1996, pp. 47-76
- ↑ a b c d David Tab Rasmussen, Thomas M. Bown and Elwyn L. Simons: The Eocene-Oligocene transition in continental Africa. In: Donald R. Prothero and William A. Berggren (eds.): Eocene-Oligocene Climatic and Biotic Evolution. Princeton University Press, Princeton, 1992, pp. 548-566
- ↑ a b c d e Patricia A Holroyd, Elwyn L Simons, Thomas M Bown, Paul D Polly and Mary J Kraus New records of terrestrial mammals from the Upper Eocene Qasr el Sagha Formation, Fayum depression, Egypt. Palaeovertebrata 25 (2-4), 1996, pp. 175-192
- ↑ a b c d Erik R. Seiffert, Thomas M. Bown, William C. Clyde, and Elwyn Simons: Geology, Paleoenvironment, and Age of Birket Qarun Locality 2 (BQ-2), Fayum Depression, Egypt. In: JG Fleagle and CC Gilbert (eds.): Elwyn Simons: A Search for Origins. Springer, 2008, pp. 71–86
- ^ a b c Elwyn L. Simons: Description of two genera and species of Late Eocene Anthropoidea from Egypt. PNAS 86, 1989, pp. 9956-9960
- ↑ a b David Tab Rasmussen and EL Simons: The oldest hyracoids (Mammalia: Pliohyracidae): new species of Saghatherium and Thyrohyrax from the Fayum. New Yearbook of Geology and Paleontology Papers 182, 1991, pp. 187–209
- ^ a b c d e f Elwyn L. Simons and Albert E. Wood: Early Cenozoic Mammalian Faunas Fayum Province, Egypt. Bulletin of the Peabody Museum of Natural History 28, 1968, pp. 1-105
- ↑ a b Elwyn L. Simons: Egyptian Oligocene Primates: A Review. Yearbook of Physical Anthropology 38, 1995, pp. 199-238
- ↑ Samar Nour-El-Deen, Romain Thomas and Wagieh El-Saadawi: First record of fossil Trachycarpeae in Africa: three new species of Palmoxylon from the Oligocene (Rupelian) Gebel Qatrani Formation, Fayum, Egypt. Journal of Systematic Palaeontology 16 (9), 2018, pp. 741-766, doi:10.1080/14772019.2017.1343258
- ↑ Richard Krausel: Results of the research trips of Prof. E. Stromer in the deserts of Egypt. IV. The fossil floras of Egypt. Papers of the Bavarian Academy of Sciences Mathematical and Natural Sciences Departments, Neue Folge, 47, 1939, pp. 1-140 ( [1] )
- ↑ Wagieh E. El-Saadawi: On the fossil flora of Jebel Qatrani area, Fayum, Egypt. Taeckholmia 26, 2006, pp. 131-140
- ↑ a b Amin Strougo, Mahmoud Faris, Mona AY Haggag, Radwan, A Abdul-Nasr, and Philip D Gingerich: Planktonic foraminifera and calcareous nannofossil biostratigraphy through the middle to late Eocene transition at Wadi Hitan, Fayum Province, Egypt. Contributions from the Museum of Paleontology 32, 2013, pp. 111-138
- ↑ Akmal Mohamed Marzouk, Ahmed Moustafa El Shishtawy and Atef Masoud Kasem: Calcareous nannofossil and planktonic foraminifera biostratigraphy through the Middle to Late Eocene transition of Fayum area, Western Desert, Egypt. Journal of African Earth Sciences 100, 2014, pp. 303-323
- ↑ Jessica L. Anderson and Rodney M. Feldmann: Lobocarcinus lumacopius (Decapoda: Cancridae), a new species of cancrid crab from the Eocene of Fayum, Egypt. Journal of Paleontology 69 (5), 1995, pp. 922-932
- ↑ a b c Manja Voss, Mohammed Sameh M Antar, Iyad S Zalmout and Philip D Gingerich: Stomach contents of the archaeocete Basilosaurus isis: Apex predator in oceans of the late Eocene. PLoS ONE 14(1), 2019, p.e0209021, doi:10.1371/journal. pone.0209021
- ↑ Charlie J. Underwood and David J. Ward: New Hemigaleid Shark from the Late Eocene of Wadi Al-Hitan, Egypt. Journal of Vertebrate Paleontology 31(3), 2011, pp. 707-711
- ↑ Gerard R. Case and Henri Cappetta: The Eocene selachian fauna from the Fayum Depression in Egypt. Palaeontographica Department A 212, 1990, pp. 1-30
- ↑ a b Alison M Murray, Todd D Cook, Yousry S Attia, Prithijit Chatrath and Elwyn L Simons: A Freshwater Ichthyofauna from the Late Eocene Birket Qarun Formation, Fayum, Egypt. Journal of Vertebrate Paleontology 30 (3), 2010, pp. 665-680
- ↑ Charlie J Underwood, David J Ward, Christopher King, Sameh M Antar, Iyad S Zalmout, and Philip D Gingerich: Shark and ray faunas in the Middle and Late Eocene of the Fayum Area, Egypt. Proceedings of the Geologists' Association 122, 2011, pp. 47-66
- ↑ Alison M Murray, Elwyn L Simons and Yousry S Attia: A new clupeid fish (Clupeomorpha) from the Oligocene of Fayum, Egypt, with notes on some other fossil clupeomorphs. Journal of Vertebrate Paleontology 25(2), 2005, pp. 300-308
- ↑ Sanaa E El-Sayed, Mahmoud A Kora, Hesham M Sallam, Kerin M Claeson, Erik R Seiffert and Mohammed S Antar: A new genus and species of marine catfishes (Siluriformes; Ariidae) from the upper Eocene Birket Qarun Formation, Wadi El-Hitan, Egypt. PLoS ONE 12(3), 2017, p.e0172409, doi:10.1371/journal.pone.0172409
- ↑ Harry L. Fierstine and Philip D. Gingerich: A second and more complete rostrum of Xiphiorhynchus aegyptiacus Weiler, 1929 (Perciformes: Xiphioidei, Xiphiidae, Xiphiorhynchinae), from the Birket Qarun Formation, late Eocene, Egypt. Journal of Vertebrate Paleontology 29(2), 2009, pp. 589-593
- ↑ Alison M. Murray and Yousry S. Attia: A new species of Lates (Teleostei: Perciformes) from the Lower Oligocene of Egypt. Journal of Vertebrate Paleontology 24(2), 2004, pp. 299-308
- ↑ Alison M. Murray: A new channid fish (Teleostei: Channiformes) from the Eocene and Oligocene of Egypt. Journal of Paleontology 80 (6), 2006, pp. 1172-1178
- ↑ Alison M. Murray: Lower pharyngeal jaw of a cichlid fish (Actinopterygii; Labroidei) from an early Oligocene site in the Fayum, Egypt. Journal of Vertebrate Paleontology 22(2), 2002, pp. 453-455
- ↑ a b Alison M. Murray: Late Eocene and Early Oligocene teleost and associated ichthyofauna of the Jebel Qatrani Formation, Fayum, Egypt. Palaeontology 47 (3), 2004, pp. 711-724
- ↑ Alison M Murray, Thodoris Argyriou and Todd D Cook: Palaeobiogeographic relationships and palaeoenvironmental implications of an earliest Oligocene Tethyan ichthyofauna from Egypt. Canadian Journal of Earth Sciences 51(10), 2014, pp. 909-918
- ↑ Patricia A. Holroyd and James Ford Parham: The antiquity of African tortoises. Journal of Vertebrate Paleontology 23 (3), 2003, pp. 688-690
- ↑ a b Charles W. Andrews: On some Pleurodiran Chelonians from the Eocene of the Fayum, Egypt. Annals and magazine of natural history 7 (11), 1903, pp. 115–121 ( [2] )
- ↑ Eugene S Gaffney, Donald D DeBlieux, Elwyn L Simons, Marcelo R Sánchez-Villagra, and Peter A Meylan: Redescription of the Skull of Dacquemys Williams, 1954, a Podocnemidid Side-Necked Turtle from the Late Eocene of Egypt . American Museum Novitates 3372, 2002, pp. 1-16
- ↑ Eugene S Gaffney, Peter A Meylan, Roger C Wood, Elwyn Simons, and Diogenes De Almeida Campos: Evolution of the Side-Necked Turtles: The Family Podocnemididae. Bulletin of the American Museum of Natural History 350, 2011, pp. 1-237
- ↑ Adán Pérez-García: New shell information and new generic attributions for the Egyptian podocnemidid turtles "Podocnemis" fajumensis (Oligocene) and "Podocnemis" aegyptiaca (Miocene). Fossil Record 24, 2021, pp. 247-262, doi:10.5194/fr-24-247-2021
- ↑ Krister T Smith, Bhart-Anjan S Bhullar and Patricia A Holroyd: Earliest African Record of the Varanus Stem-Clade (Squamata: Varanidae) from the Early Oligocene of Egypt. Journal of Vertebrate Paleontology 28(3), 2008, pp. 909-913
- ↑ Robert B Holmes, Alison M Murray, Yousry S Attia, Elwyn L Simons and Prithijit Chatrath: Oldest known Varanus (Squamta: Varanidae) from the Upper Eocene and Lower Oligocene of Egypt: Suport for an African origin of the Genus . Palaeontology 53(5), 2010, pp. 1099-1110
- ↑ Jacob A McCartney and Erik R Seiffert: A late Eocene snake fauna from the Fayum depression, Egypt. Journal of Vertebrate Paleontology 36(1), 2016, p.e1029580, doi:10.1080/02724634.2015.1029580
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