Phosphatherium

Last updated

Phosphatherium
Temporal range: [Ypresian]
Phosphatherium escuilliei.JPG
Holotype of the species Phosphatherium escuilliei
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Proboscidea
Family: Numidotheriidae
Genus: Phosphatherium
Gheerbrant, Sudre & Cappetta, 1996 [1]
Species:
P. escuillei
Binomial name
Phosphatherium escuillei
Gheerbrant, Sudre & Cappetta, 1996 [1]

Phosphatherium escuillei Gheerbrant, Sudre et Cappetta 1996 is a basal proboscidean that lived in Africa during in the early Eocene, by 56-55 Ma. [2] [3] It is one of the earliest known proboscidean, together with Eritherium azzouzorum from the Selandian (about 60 Ma). [4] [5] It was found in phosphorites beds from the base of the Ypresian stage of the Ouled Abdoun Basin, which is best known for its exceptionally rich marine vertebrate fauna. [6] [7]

Contents

Description

Restoration of the head of P. escuilliei by Dominique Visset Phosphatherium - head restoration.tif
Restoration of the head of P. escuilliei by Dominique Visset

The species P. escuillei is known from relatively well-preserved fossils, including several sub-complete skulls that have enabled the reconstruction of the animal's head. [8]


The skull of Phosphatherium is very primitive compared to advanced proboscideans (Elephantiformes), for example the nasal openings are anterior, indicating that it lacked a trunk. Other plesiomorphic traits are the elongated face, the narrow rostrum, the strong sagittal and nuchal crests, the presence of an alisphenoid canal, the auditory meatus that remain open ventrally, the tooth rows that does not extend back more than half of the total skull length and the presence of a canine and anterior premolars. However, Phosphatherium shares several advanced characteristics (synapomorphies) with the proboscidean order, such as the development of the maxillary bone which forms the lower rim of the orbit and part of the robust zygomatic process, the petrosal that has large pars mastoidea, the frontal that is in contact with the squamosal, the molars with sharp transverse crests (true bilophodonty),  the lower molar with posterior cusp (hypoconulid) shifted in labial position (first step in the development of a third  loph found in later proboscideans such as elephantiforms) and the enlarged lower central incisor. The first lower incisor (I1) showed clear enlargements, but it was not vergrowing as in the tusks of modern elephants.The dental formula (number of teeth ) is nearly complete as in ancestral placentals, it only lack two anterior teeth (an incisor and the canine) at the lower jaw. Adult animals had indeed the following dental formula:

Dentition
 ?.1.4.3
2.0.4.3

The molars are low (brachyodont) and lophodont, with sharp transverse crests, which indicates that Phosphatherium had a herbivorous diet specialized in eating leaves (follivory). The skull length is about 17 cm.

Skull Phosphatherium, discovered in 1996 Phosphatherium skull 1996.jpg
Skull Phosphatherium, discovered in 1996

Some features of the teeth and jaw structures of P. escuilliei show significant individual variation, including the development of the molar crests (lophodonty). Part of this individual variation is related to sexual dimorphism. This includes the development of the submaxillary fossa housing muscles for mastication.

Discovery

Map showing where fossils have been found Phosphatherium escuilliei distribution.svg
Map showing where fossils have been found

The first finds were probably made by a fossil dealer in the early 1990s from an unknown location. The two fragments of the upper jaw were coated with phosphate, and the preparation of the finds revealed the tooth of a fossil shark, which dates to the late Paleocene (Thanetian) and thus confirms the ancient age of the finds. The holotype (copy number MNHN PM2) consists of a right upper part of the jaw with the last two premolars and the first two molars (P3 to M2). It is now in the Muséum National d'Histoire Naturelle in Paris. The first scientific description took place in 1996 by Emmanuel Gheerbrant and colleagues. The name Phosphatherium is made up of the Greek words φωςφορος (phosphoros "light-bearing") and θηρίον ( thērion "animal") and refers to the fact that it is stored in phosphate-containing sediments. The only known species is Phosphatherium escuilliei. The species name escuilliei honors the person who found the type fossils, François Escuillié. The first description was largely limited to the tooth features, a more extensive template of the then known find material was only published two years later. [1]

The first fossil finds were discovered in the north-eastern part of the Ouled Abdoun Basin in a phosphate-containing layer. However, they only comprised two fragmented upper jaws. However, the exact location of this find was not known. Further and much more extensive finds were discovered in the early 2000s in the same basin in the Grand Daoui area. This consists of numerous skull fragments, mandibles and some limb elements. [9] [10] Other very early proboscideans are known from the same area, the related Eritherium and Daouitherium . [11]

Taxonomy

Phosphatherium is known primarily from two maxilla fragments dated to the latest Paleocene deposits of the Ouled Abdoun Basin, Morocco, which date from the Thanetian epoch. It is one of the oldest [12] and smallest members of the Proboscidea, with an estimated shoulder height of about 30 centimetres (12 in) and body mass of 17 kilograms (37 lb). [13] Like its later relative, Moeritherium , the animal was probably an amphibious browser that fed on aquatic plants, akin to a very small tapir. Both animals are included in the family Numidotheriidae, together with Numidotherium .

Below is a phylogenetic tree based on Tabuce et al. 2019. [14]

  Proboscidea  

  Eritherium

 Phosphatherium

  Daouitherium

  Numidotherium

  Arcanotherium

  Saloumia

  Moeritherium

 Derived Proboscidea (Elephantiformes)

  Deinotheriidae

Palaeobiology

Jaw fragments Phosphatherium escuilliei 65.JPG
Jaw fragments

Phosphatherium is thought to have had a broad diet. The dental microwear patterns observed on their teeth show lengthy scratches on the molars of juveniles. Correspondingly, similar patterns are found on adult individuals. Through study of the wear and specifically scratches on the teeth of Phosphatherium, the food items it ingested include shrubs and bushes, indicating a mixed feeding preference. Adult molars are found to have a much higher density of scratches, indicating abrasive food sources and possibly insects and small animals. Overall, Phosphatherium is thought to be an omnivorous browser mainly determined by its preferences, as well as the availability of resources. [15]

Considering its highly adapted folivorous jaw and tooth structure, Phosphatherium provides evidence of the high age of African endemism. The dental structure of Phosphatherium suggests its diet consisted mainly of leaves, which indicates it may have fulfilled a niche role in its environment, although diet can only be inferred. The discovery of this animal has ultimately helped reinforce the African origin of proboscideans, and provide insight into the radiation of modern orders of placental mammals. [10]

Related Research Articles

<i>Anancus</i> Genus of proboscideans

Anancus is an extinct genus of "tetralophodont gomphothere" native to Afro-Eurasia, that lived from the Tortonian stage of the late Miocene until its extinction during the Early Pleistocene, roughly from 8.5–2 million years ago.

Numidotheriidae is an extinct family of primitive proboscideans that lived from the late Paleocene to the early Oligocene periods of North Africa.

<i>Daouitherium</i> Extinct genus of mammals

Daouitherium is an extinct genus of early proboscideans that lived during the early Eocene some 55 million years ago in North Africa.

<i>Arambourgisuchus</i> Extinct genus of reptiles

Arambourgisuchus is an extinct genus of dyrosaurid crocodylomorph from the late Palaeocene of Morocco, found in the region of Sidi Chenane in 2000, following collaboration by French and Moroccan institutions, and described in 2005 by a team led by palaeontologist Stéphane Jouve. Arambourgisuchus was a large animal with an elongated skull 1 meter in length.

<i>Dyrosaurus</i> Extinct genus of reptiles

Dyrosaurus is a genus of extinct crocodylomorph that lived during the early Eocene. The name Dyrosaurus comes from sauros (σαῦρος) the Greek for lizard or reptile, and Dyr for Djebel Dyr (mountain) close to where the type species was discovered. It was a large reptile with an estimated body length of 6.5 metres (21 ft).

<i>Argochampsa</i> Extinct genus of reptiles

Argochampsa is an extinct genus of eusuchian crocodylomorph, usually regarded as a gavialoid crocodilian, related to modern gharials. It lived in the Paleocene of Morocco. Described by Hua and Jouve in 2004, the type species is A. krebsi, with the species named for Bernard Krebs. Argochampsa had a long narrow snout, and appears to have been marine in habits.

<i>Eritherium</i> Extinct genus of mammals

Eritherium is an extinct genus of early Proboscidea found in the Ouled Abdoun basin, Morocco. It lived about 60 million years ago. It was first named by Emmanuel Gheerbrant in 2009 and the type species is Eritherium azzouzorum. Eritherium is the oldest, smallest and most primitive known elephant relative.

<i>Zarafasaura</i> Extinct genus of reptiles

Zarafasaura is an extinct genus of elasmosaurid known from the Ouled Abdoun Basin of Morocco. As a relatively small elasmosaur, it would have measured around 3–4 metres (9.8–13.1 ft) long and weighed about 100 kilograms (220 lb).

<i>Ocepeia</i> Extinct Afrotherian mammal

Ocepeia is an extinct genus of afrotherian mammal that lived in present-day Morocco during the middle Paleocene epoch, approximately 60 million years ago. First named and described in 2001, the type species is O. daouiensis from the Selandian stage of Morocco's Ouled Abdoun Basin. A second, larger species, O. grandis, is known from the Thanetian, a slightly younger stage in the same area. In life, the two species are estimated to have weighed about 3.5 kg (7.7 lb) and 10 kg (22 lb), respectively, and are believed to have been specialized leaf-eaters. The fossil skulls of Ocepeia are the oldest known afrotherian skulls, and the best-known of any Paleocene mammal in Africa.

<span class="mw-page-title-main">Ouled Abdoun Basin</span> Phosphate basin in Morocco

The Oulad Abdoun Basin is a phosphate sedimentary basin located in Morocco, near the city of Khouribga. It is the largest in Morocco, comprising 44% of Morocco's phosphate reserves, and at least 26.8 billion tons of phosphate. It is also known as an important site for vertebrate fossils, with deposits ranging from the Late Cretaceous (Cenomanian-Turonian) to the Eocene epoch (Ypresian), a period of about 25 million years.

<i>Abdounodus</i> Extinct genus of mammal

Abdounodus is an extinct genus of mammal known from the middle Paleocene of Northern Africa. The sole species, A. hamdii, is known from teeth and jaw bones discovered in the Ouled Abdoun Basin of present-day Morocco in 2001.

Lahimia ("carnivore") is an extinct genus of placental mammals from extinct family Hyaenodontidae within extinct order Hyaenodonta, known from the middle Paleocene of Morocco. Lahimia selloumi is the one of the oldest known members of order Hyaenodonta.

Saloumia is an extinct genus of the order Proboscidea. It is one of the oldest members of the order and lived in the middle Eocene of Senegal. It is known only from a single molar, whose pronounced bumpy chewing surface indicates it is probably closely related to Moeritherium.

<i>Ajnabia</i> Extinct genus of reptiles

Ajnabia is a genus of lambeosaurine hadrosaur from the Late Cretaceous (Maastrichtian) of Morocco. It is the first definitive hadrosaur from Africa, and is thought to be related to European dinosaurs like Arenysaurus. The discovery of Ajnabia came as a surprise to the paleontologists who found it, because Africa was isolated by water from the rest of the world during the Cretaceous, such that hadrosaurs were assumed to have been unable to reach the continent. Ajnabia is relatively small and similar in size to its contemporary relative Minqaria, which is estimated to have reached 3.5 metres (11 ft) in total body length. Assuming that the holotype represents an adult, Ajnabia would be one of the smallest if not the smallest known hadrosaurids.

Dagbatitherium is an extinct genus of proboscideans. So far a single molar from the phosphate basins of Togo in West Africa has been found. The fossil dates to the Middle Eocene, around 47 million years ago. A striking feature of the tooth are the three pairs of cusps oriented transversely to the longitudinal axis of the tooth. This feature is found in more derived proboscideans, which are grouped in the Elephantiformes. For its age, Dagbatitherium is the earliest member of Elephantiformes to date. Furthermore, it is characterized by a low tooth crown and a humped occlusal pattern. The genus was described in 2021.

<i>Thalassotitan</i> Large bodied African mosasaur

Thalassotitan is an extinct genus of large mosasaurs that lived during the late Maastrichtian of the Cretaceous period in what is now Morocco, around 67 to 66 million years ago. The only known species is T. atrox, described in 2022 from fossils discovered in the Ouled Abdoun Basin, initially identified as coming from other genera such as Mosasaurus or Prognathodon. Hypothetical Thalassotitan specimens may have been found in other corners of the world, although researchers also note the possibility that they come from distinct, related taxa. It is considered to be close to the genera Prognathodon and Gnathomortis, together forming the tribe Prognathodontini. The prognathodontines are separated from other mosasaurs based on their massive jaws and robust teeth.

Hadrogeneios is an extinct genus of afrotherian mammal recovered from the Khouribga Phosphates of the Ouled Abdoun Basin dating from the Selandian to the Thanetian. Only the type species H. phosphaticus is known for this genus, with the material being of dental and jaw specimens. The placement of Hadrogeneios recovered it to be the most basal stem-paenungulate, the clade that includes elephants, sea cows, and hyraxes. Despite this Hadrogeneios was a contemporary of more derived members of the group such as Abdounodus, Ocepeia, and the early proboscideans Eritherium, Phosphatherium, and Daouitherium.

<i>Minqaria</i> Extinct genus of hadrosaurid dinosaurs

Minqaria is a genus of arenysaurinin lambeosaurine hadrosaur from the Late Cretaceous (Maastrichtian) Ouled Abdoun Basin of Morocco. The genus contains a single species, M. bata, known from a partial skull.

Grand Daoui is a quarry in the Ouled Abdoun Basin of Morocco known for its fossils. It is the discovery place of Phosphatherium escuilliei, the earliest known proboscidean. It was the location for several field parties between 1997 and 2001, which allowed survey of the geological and paleontological context of Phosphatherium localities. It is also rich in marine vertebrae fossils. All Phosphatherium remains to date have been found in the quarry. The discovery of fossils there has aided in the study of early African placental fauna. The marine snake Palaeophis maghrebianus is also known from the locality, as well as the primitive gavialoid Argochampsa krebsi.

Boualitomus is an extinct genus of hyaenodont that lived in what is now Morocco during the earliest Eocene. It contains a single species, B. marocanensis.

References

  1. 1 2 3 Gheerbrant, E.; Sudre, J.; Cappetta, H. (1996). "A Palaeocene proboscidean from Morocco". Nature . 383 (6595): 68–71. Bibcode:1996Natur.383...68G. doi:10.1038/383068a0. S2CID   4362199.
  2. http://www3.planetarioroma.it/content/download/4908/62991/file/637_638.pdf. Archived 2016-03-05 at the Wayback Machine
  3. Gheerbrant, Gheerbrant (2003). "Les localités à mammifères des carrières de Grand Daoui, Bassin des Ouled Abdoun, Maroc, Yprésien : premier état des lieux" (PDF). Bulletin de la Société Géologique de France. 174 (3): 279–293. doi:10.2113/174.3.279.
  4. Gheerbrant, Emmanuel (2009-06-30). "Paleocene emergence of elephant relatives and the rapid radiation of African ungulates". Proceedings of the National Academy of Sciences. 106 (26): 10717–10721. doi: 10.1073/pnas.0900251106 . ISSN   0027-8424. PMC   2705600 . PMID   19549873.
  5. Kocsis, Lászlό; Gheerbrant, Emmanuel; Mouflih, Mustapha; Cappetta, Henri; Yans, Johan; Amaghzaz, Mbarek (2014). "Comprehensive stable isotope investigation of marine biogenic apatite from the late Cretaceous–early Eocene phosphate series of Morocco". Palaeogeography, Palaeoclimatology, Palaeoecology. 394: 74–88. doi:10.1016/j.palaeo.2013.11.002.
  6. Arambourg, Camille (1952). Les vertébrés fossiles des gisements de phosphates (Maroc - Algérie Tunisie) (in French). Notes et Mémoires du Service géologique du Maroc (Rabat).
  7. Zouhri, Samir (2017). Paléontologie des vertébrés du Maroc: état des connaissances. Mémoires de la Société géologique de France. Paris: Société géologique de France. ISBN   978-2-85363-099-3.
  8. 1 2 Gheerbrant, Emmanuel; Sudre, Jean; Tassy, Pascal; Amaghzaz, Mbarek; Bouya, Baâdy; Iarochène, Mohamed (2005). "Nouvelles données sur Phosphatherium escuilliei (Mammalia, Proboscidea) de l'Éocène inférieur du Maroc, apports à la phylogénie des Proboscidea et des ongulés lophodontes". Geodiversitas. 27 (2): 239–333.
  9. Gheerbrant, E. ( 1 ), et al. "New Data On Phosphatherium Escuilliei (Mammalia, Proboscidea) From The Early Eocene Of Morocco, And Its Impact On The Phylogeny Of Proboscidea And Lophodont Ungulates." Geodiversitas 27.2 (2005): 239-333.
  10. 1 2 Gheerbrant, Emmanuel (1998). "The Oldest Known Proboscidean and the Role of Africa in the Radiation of Modem Orders of Placentals". Bulletin of the Geological Society of Denmark. 44: 181–85. doi: 10.37570/bgsd-1998-44-12 .
  11. Gheerbrant, E.; Sudre, J.; Cappetta, H.; Iarochène, M.; Amaghzaz, M.; Bouya, B. (2002). "A new large mammal from the Ypresian of Morocco: Evidence of surprising diversity of early proboscideans" (PDF). Acta Palaeontologica Polonica. 47 (3): 493–506.
  12. Gheerbrant, Emmanuel; Sudre, Jean; Cappetta, Henri; Mourer-Chauviré, Cécile; Bourdon, Estelle; Iarochene, Mohamed; Amaghzaz, Mbarek; Bouya, Baâdi (2003). "The mammal localities of Grand Daoui Quarries, Ouled Abdoun Basin, Morocco, Ypresian : A first survey" (PDF). Bulletin de la Société Géologique de France. 174 (3): 279–293. doi:10.2113/174.3.279.
  13. Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans" (PDF). Acta Palaeontologica Polonica. 61. doi: 10.4202/app.00136.2014 .
  14. Rodolphe Tabuce, Raphaël Sarr, Sylvain Adnet, Renaud Lebrun, Fabrice Lihoreau, Jeremy E. Martin, Bernard Sambou, Mustapha Thiam and Lionel Hautier; 2019 "Filling a gap in the proboscidean fossil record: a new genus from the Lutetian of Senegal." Journal of Paleontology. doi:10.1017/jpa.2019.98
  15. Calandra, Ivan; Göhlich, Ursula B.; Merceron, Gildas (2010). "Feeding preferences of Gomphotherium subtapiroideum (Proboscidea, Mammalia) from the Miocene of Sandelzhausen (Northern Alpine Foreland Basin, southern Germany) through life and geological time: evidence from dental microwear analysis". Paläontologische Zeitschrift. 84 (1): 205–215. Bibcode:2010PalZ...84..205C. doi:10.1007/s12542-010-0054-0. S2CID   56260944.

Further reading