Saadanius Hijazensis 003
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''Saadanius'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of fossil
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
dating to the
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but the ...
that is closely related to the
common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
of the
Old World monkey Old World monkey is the common English name for a family of primates known taxonomically as the Cercopithecidae (). Twenty-four genera and 138 species are recognized, making it the largest primate family. Old World monkey genera include baboons ...
s and
ape Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its siste ...
s, collectively known as
catarrhines The parvorder Catarrhini , catarrhine monkeys, Old World anthropoids, or Old World monkeys, consisting of the Cercopithecoidea and apes (Hominoidea). In 1812, Geoffroy grouped those two groups together and established the name Catarrhini, "Old W ...
. It is represented by a single species, ''Saadanius hijazensis'', which is known only from a single partial skull tentatively dated between 29 and 28 million years ago. It was discovered in 2009 in western
Saudi Arabia Saudi Arabia, officially the Kingdom of Saudi Arabia (KSA), is a country in Western Asia. It covers the bulk of the Arabian Peninsula, and has a land area of about , making it the fifth-largest country in Asia, the second-largest in the A ...
near
Mecca Mecca (; officially Makkah al-Mukarramah, commonly shortened to Makkah ()) is a city and administrative center of the Mecca Province of Saudi Arabia, and the Holiest sites in Islam, holiest city in Islam. It is inland from Jeddah on the Red ...
and was first described in 2010 after a comparison with both living and fossil catarrhines. ''Saadanius'' had a longer face than living catarrhines and lacked the advanced
frontal sinus The frontal sinuses are one of the four pairs of paranasal sinuses that are situated behind the brow ridges. Sinuses are mucosa-lined airspaces within the bones of the face and skull. Each opens into the anterior part of the corresponding middle na ...
found in living catarrhines. However, it had a bony
ectotympanic The ectotympanic, or tympanicum, is a bony structure found in all mammals, located on the tympanic part of the temporal bone, which holds the tympanic membrane (eardrum) in place. In catarrhine primates (including humans), it takes a tube-shape. I ...
and teeth comparable to those of living catarrhines. The discovery of ''Saadanius'' may help answer questions about the evolution and appearance of the last common ancestors of Old World monkeys and apes.


Discovery

''Saadanius'' is known from a single specimen, the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, named "SGS-UM 2009-002", stored in
Jeddah Jeddah ( ), also spelled Jedda, Jiddah or Jidda ( ; ar, , Jidda, ), is a city in the Hejaz region of the Kingdom of Saudi Arabia (KSA) and the country's commercial center. Established in the 6th century BC as a fishing village, Jeddah's pro ...
, Saudi Arabia, at the Paleontology Unit of the
Saudi Geological Survey The Saudi Geological Survey (SGS; ar, هيئة المساحة الجيولوجية السعودية) is the national geological survey of the Kingdom of Saudi Arabia. History and profile The SGS was established as an independent entity attached ...
(SGS). Its type locality was on top of an oolitic
ironstone Ironstone is a sedimentary rock, either deposited directly as a ferruginous sediment or created by chemical replacement, that contains a substantial proportion of an iron ore compound from which iron (Fe) can be smelted commercially. Not to be con ...
fossil bed of the middle Shumaysi Formation located in the southwest corner of Harrat Al Ujayfa, in western Saudi Arabia, close to Mecca. The specimen was discovered in February 2009 by paleontologist Iyad Zalmout, who had traveled to the region to search for ancient whale and
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
fossils. While looking for dinosaur fossils in an area that, according to the maps he was working from, contained rock layers that dated to the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
, Zalmout found the jawbone of an anthracotheriid, which dated to the
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
or Oligocene. This indicated that the rock layers were much younger than what was dated on the maps. The following day, he noticed fossil teeth, which he immediately recognized as those of a primate. Zalmout emailed a photo of the teeth to paleontologist
Philip D. Gingerich Philip Dean Gingerich (born March 23, 1946) is a paleontologist and educator. He is Professor Emeritus of Geology, Biology, and Anthropology at the University of Michigan, Ann Arbor. He directed the Museums at the University of Michigan#Museum ...
, with whom he was working as a postdoctoral fellow. Gingerich, an expert on ancient primates and whales, confirmed that it was indeed a primate. Due to a tight schedule, Zalmout had to leave the exposed fossil embedded in the rock for the next few days because collecting it would require days of work. The fossil was soon recovered by a joint expedition involving the SGS and the
University of Michigan , mottoeng = "Arts, Knowledge, Truth" , former_names = Catholepistemiad, or University of Michigania (1817–1821) , budget = $10.3 billion (2021) , endowment = $17 billion (2021)As o ...
. The fossil was not formally described until 2010 when its discovery was announced in the journal ''
Nature Nature, in the broadest sense, is the physics, physical world or universe. "Nature" can refer to the phenomenon, phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. ...
''. The fossil was dated to about 28 or 29 million years ago based on comparisons of fossils of anthracotheriids and paenungulates found nearby with fossils from similarly aged African deposits. However, the date has yet to be confirmed by other dating techniques. The
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
name, ''Saadanius'', comes from the Arabic word, ''saadan'' ( ar, سَعدان), which is the collective term for apes and monkeys. The species name, ''hijazensis'', is a reference to the al Hijaz region, in which it was discovered.


Type specimen

''Saadanius'' had a longer face than living catarrhines, more closely resembling
New World monkey New World monkeys are the five families of primates that are found in the tropical regions of Mexico, Central and South America: Callitrichidae, Cebidae, Aotidae, Pitheciidae, and Atelidae. The five families are ranked together as the Ceboidea ( ...
s in appearance, although it was larger—possibly the size of a
gibbon Gibbons () are apes in the family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical rainforest from eastern Bangladesh to Northeast India ...
. Its enlarged, deep-rooted
canine teeth In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...
, the
diastema A diastema (plural diastemata, from Greek διάστημα, space) is a space or gap between two teeth. Many species of mammals have diastemata as a normal feature, most commonly between the incisors and molars. More colloquially, the condition ...
between its canine teeth and second
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...
s, and its
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
suggest that the specimen was a male. These features are shared among male Old World monkeys. ''Saadanius'' lacks the advanced
frontal sinus The frontal sinuses are one of the four pairs of paranasal sinuses that are situated behind the brow ridges. Sinuses are mucosa-lined airspaces within the bones of the face and skull. Each opens into the anterior part of the corresponding middle na ...
found in living catarrhines, but it does possess an
ectotympanic The ectotympanic, or tympanicum, is a bony structure found in all mammals, located on the tympanic part of the temporal bone, which holds the tympanic membrane (eardrum) in place. In catarrhine primates (including humans), it takes a tube-shape. I ...
, found in living catarrhines. Propliopithecoids, the oldest
stem group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
of catarrhines, which date back 35 to 30 mya, lacked a fully developed ectotympanic. The specimen had serious puncture wounds on the front of the skull, indicating that it had been preyed upon by a large
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
. One puncture wound was on the right side of the
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
and may have been the fatal blow. There was also a bite mark on the frontal trigone. Like other catarrhine primates, ''Saadanius'' was probably a tree-dweller. During the time it would have lived, the
Red Sea The Red Sea ( ar, البحر الأحمر - بحر القلزم, translit=Modern: al-Baḥr al-ʾAḥmar, Medieval: Baḥr al-Qulzum; or ; Coptic: ⲫⲓⲟⲙ ⲛ̀ϩⲁϩ ''Phiom Enhah'' or ⲫⲓⲟⲙ ⲛ̀ϣⲁⲣⲓ ''Phiom ǹšari''; T ...
had not yet formed, and new plant and animal species would have been arriving from nearby
Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...
as it converged with the Afro-Arabian landmass.


Phylogeny and significance

Comparative anatomy Comparative anatomy is the study of similarities and differences in the anatomy of different species. It is closely related to evolutionary biology and phylogeny (the evolution of species). The science began in the classical era, continuing in t ...
and
cladistic analysis Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived chara ...
indicate that ''Saadanius'' is more closely related to the last common ancestor of
crown A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, partic ...
catarrhines than any other known fossil catarrhines, placing the common ancestry of Catarrhini in Arabia and Africa. Other stem catarrhines include propliopithecoids, such as ''
Aegyptopithecus ''Aegyptopithecus'' ("Egyptian ape", from Greek ''Αίγυπτος'' "Egypt" and ''πίθηκος'' "ape") is an early fossil catarrhine that predates the divergence between hominoids (apes) and cercopithecids (Old World monkeys). It is known ...
'', and pliopithecoids, such as ''
Pliopithecus ''Pliopithecus'' {meaning "more ape") is a genus of extinct primates of the Miocene. It was discovered in 1837 by Édouard Lartet (1801–1871) in France, with fossils subsequently discovered in Switzerland, Slovakia and Spain. ''Pliopithecus'' ...
''. The closer similarities between ''Saadanius'' and crown catarrhines, particularly its ectotympanic, suggest ''Saadanius'' of all known fossil primates most closely resembled the last common ancestor of living catarrhines. Its discovery has provided new details about
human evolution Human evolution is the evolutionary process within the history of primates that led to the emergence of ''Homo sapiens'' as a distinct species of the hominid family, which includes the great apes. This process involved the gradual development of ...
, particularly the split between the Old World monkeys and the ape lineage that eventually led to humans. The discovery of ''Saadanius'' provides new evidence for competing hypotheses about the facial appearance of the ancestral crown catarrhines, or common ancestor. One reconstruction is based on living catarrhine traits and predicts a short face and a rounded braincase, similar to that of a gibbon. Another reconstruction, based on the
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
of early
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
apes and the basal cercopithecoid ''
Victoriapithecus ''Victoriapithecus macinnesi'' was a primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the s ...
'', predicts that the last common ancestor had a projecting snout and tall face, like that of living
baboon Baboons are primates comprising the genus ''Papio'', one of the 23 genera of Old World monkeys. There are six species of baboon: the hamadryas baboon, the Guinea baboon, the olive baboon, the yellow baboon, the Kinda baboon and the chacma ba ...
s and the oldest fossil apes and Old World monkeys. The conservative features of ''Saadanius'', similar to those of the older stem catarrhines, support the latter hypothesis, according to Zalmout ''et al.'' However, one palaeontologist, Eric Delson, has cautioned that geological pressure may have distorted the shape of the skull. According to Zalmout ''et al.'', ''Saadanius'' may also help resolve the age of the hominoid–cercopithecoid split. Paleoanthropological work has typically placed the divergence between 25 and 23 mya, but genetic-based estimates have placed it in the early Oligocene, approximately 33 mya. Despite the predictions from the genetic tests, little fossil evidence has been found for a last common ancestor between 30 and 23 mya, favoring a later split. Only isolated teeth of ''
Kamoyapithecus ''Kamoyapithecus'' ('Kamoya' + Greek - “ape”) was a primate that lived in Africa during the late Oligocene period, about 24.2-27.5 million years ago. First found in 1948 as part of a University of California, Berkeley expedition, it was at fi ...
'' hinted at the existence of potential basal
hominoids Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its siste ...
in the late Oligocene (between 24 and 27.5 mya), while the oldest fossil Old World monkey, ''Victoriapithecus macinnesi'', dates to 19 mya. With the discovery of ''Saadanius'', Zalmout ''et al.'' suggested a later split than the genetic data, dating between 29–28 and 24 mya. However, Pozzi ''et al.'' later argued that although ''Saadanius'' is a significant discovery, because it is a stem catarrhine, it could not be used to date the divergence of the crown group. The presence of stem
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
in the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
does not indicate that crown groups have evolved, and stem taxa may survive for millions of years after the crown taxa appear. For this reason, the fossil record can only suggest a hard minimum boundary for divergence dates, which corresponds to the first appearance of a crown taxon. Furthermore, Pozzi ''et al.'' pointed out that the supplementary material published by Zalmout ''et al.'' demonstrated that Pliopithecoidea were more closely related to living catarrhines than ''Saadanius''. The fossil find has also been seen by the SGS as an important find for Saudi Arabia, because it enriches the fossil record for the region. As a result of the find, both the SGS and the University of Michigan are considering more collaborative field explorations in the country.


References


External links


A piece in the monkey puzzle
– A short video from Nature explaining the ''Saadanius'' discovery and its significance. {{Taxonbar, from=Q134357, from2=Q18618800, from3=Q18618801, from4=Q18618803 Prehistoric primate genera Oligocene primates Fossil taxa described in 2010 Catarrhini Articles containing video clips Oligocene mammals of Asia