Numidotherium Temporal range: | |
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Drawing of skull from Numidotherium with restored silhouette. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Proboscidea |
Family: | † Numidotheriidae |
Genus: | † Numidotherium Jaeger, 1986 |
Species: | †N. koholense |
Binomial name | |
†Numidotherium koholense Jaeger, 1986 | |
Numidotherium ("Numidia beast") is an extinct genus of early proboscideans, discovered in 1984, that lived during the middle Eocene of North Africa some 46 million years ago. It was about 90-100 cm tall at the shoulder and weighed about 250-300 kg. [1]
The type species, N. koholense, is known from an almost complete skeleton from the site of El Kohol, southern Algeria, dating from the early/middle Eocene period. The animal had the size and the appearance of a modern tapir. In appearance, it was more slender and more plantigrade than an elephant, its closest modern relative. [2]
Numidotherium savagei, named by Court (1995) for material from late Eocene deposits at Dor el Talha, Libya, has been reassigned to its own genus, Arcanotherium . [3] [4]
Numidotherium is one of the earliest representatives of the Proboscidea, and is known from several tooth and skull finds, and also from several postcranial fossils. [2] It reached a shoulder height of 90 to 100 cm [1] and was thus significantly larger than its predecessors Phosphatherium and Eritherium . Furthermore, it was characterized by a short trunk and quite long legs. The skull had a length of 37 cm and a raised cranial roof and had widely spreading zygomatic arches. [2] As one of the earliest proboscideans, Numidotherium had air-filled bones in the roof of the skull, which, due to the increase in size and weight of the animal, offered a larger attachment point for the neck muscles. The rostrum was short and sloped steeply down from the forehead to the nose. The nasal bone was characterized by an elongated outer area of the nasal space (nares), while the intermaxillary bone was rather short and high. The shape of the rostrum created a high, but rather narrow, nasal interior. On the side of the upper jaw, a deep furrow ran from the intermaxillary bone to the anterior approach of the zygomatic arch. In some individuals the temporal ridges on the parietal bone formed a distinctive crest, in others they remained clearly separated. The occiput was also quite short in shape, with a robust occipital bulge. The joint surfaces here for articulating with the cervical spine were comparatively high.
The lower jaw, up to 39 cm in length, was low but broad. [2] The symphysis extended to the second premolar. The mental foramen lay below the third premolar tooth. The ascending branch loomed up and delimited a prominent masseteric fossa. The crown process was oriented slightly inwards. The number of teeth in the set of teeth was slightly reduced compared to older proboscideans. The second incisor in the upper jaw (I2) was significantly enlarged with a length of 4.5 cm and was conical in shape. Furthermore, it possessed only a thin layer of enamel. In the lower jaw, the first incisor, protruding obliquely forward (procumbent), was enlarged by 3.5 cm - the first examinations here looked at the second incisor as the enlarged one. These enlargements represent the preliminary stages of tusk formation in later proboscideans. There was a large diastema to each of the rear teeth, which could be up to 9.5 cm long. The premolars were characterized by a simple construction with a single raised enamel cusp. The molars, on the other hand, had two transversely positioned enamel ridges (bilophodont) and were reminiscent of those of tapirs. The entire posterior dentition can be described as a low crown (brachydont). [2] [4]
The previously known postcranial skeleton comprises individual vertebrae, especially the first and second cervical vertebrae (atlas and axis), which differ only slightly from those of other proboscideans. The humerus reached a length of 30 cm and was robustly designed. The bones of the forearm, ulna and radius reached about the same length and were fused at the lower end. The upper joint end of the ulna (olecranon) was relatively short and oriented backwards. The thigh bone measured 40.5 cm. The latter was clearly narrowed anteriorly and posteriorly, but had a very broad upper joint end and a third trochanter. The shin was significantly shorter at 26 cm. The forefoot and hindfoot had the "taxeopodous" arrangement of the root bones typical of proboscideans. That is, the individual bones of the palm and tarsal lay in rows one behind the other and not reciprocally to one another. As a result, there was no contact between the lunate bone and the hamate on the forefoot. The special arrangement of the carpal bones suggests that an os centrale was formed between the lunate, the hamate and the trapezoid bone, which should be understood as a novel feature of the Proboscidea. [2] [5] [6] [7]
Numidotherium finds come exclusively from North Africa. The first specimens were discovered in El Kohol on the southern edge of the Atlas Mountains in Algeria. They come from the third layer of the El-Kohol Formation and date to the Middle Eocene. [2] The rather extensive finds include a skull, a lower jaw and remains of the musculoskeletal system. A much older find from the Lower Eocene is known with a lower jaw fragment from a phosphate mine near Tamaguèlelt in northern Mali. However, this belongs to a much smaller representative and may have to be assigned to a different taxon. Further dental finds were found in connection with a rich fish fauna from Ad-Dakhla in Western Sahara and can be assigned to the transition from the Middle to the Upper Eocene. [2]
Below is a phylogenetic tree of early Proboscidea, based on the work of Hautier et al. (2021). [8]
The construction of the nasal bone with the elongated outer nasal cavities suggests the presence of a short proboscis in Numidotherium. [6] The early proboscideans are commonly associated with a semi-aquatic way of life, as proven in the closely related Barytherium and in Moeritherium . The fused bones of the forearm support this view, but the plantigrade structure of the forefoot shows adaptations to terrestrial locomotion, just as the low position of the eyes speaks against a water-dwelling way of life. The low-crowned molars with their typical, clearly formed lophodontic chewing surfaces indicate specialization in relatively soft vegetation, which should have predominantly included leaves, fruits and twigs. The fibrous food was probably looked for on bushes at head height or just above it, characteristic grinding marks on the anterior premolars suggest that the animals wedged them in the mouth and tore them off with lateral movements of the head. [3] Isotope analysis of the teeth confirm this assumed terrestrial way of life. In the case of Numidotherium compared to Barytherium or Moeritherium, these resulted in more fluctuating ratios of the oxygen isotope O18, which in the latter two are significantly more balanced due to their semi-aquatic way of life. It is possible that an adaptation to life in a semiaquatic environment occurred several times within the Proboscidea, whereby the question remains unanswered as to whether the proboscis, including today's elephants, actually descended from some water-dwelling ancestors. [9]
The traditional postcranial skeleton of Numidotherium allows conclusions to be drawn concerning its locomotion. Due to the arrangement of individual joint surfaces in the shoulder and pelvic area and the fused bones of the forearm, slightly bowed limbs can be assumed. The bones of the front and rear feet are also more horizontally oriented, which is suggested by the structure of the ankle bone with a forward-facing joint head. The joint facets of the foot bones, which are sometimes clearly positioned to the side, indicate that the individual bones (metapodia and phalanges) were rather spread apart. All in all, it can be assumed that Numidotherium has a sole passage. This is in clear contrast to today's elephants, which have column-like limbs arranged under the body, which can carry the heavy weight. The front and rear feet, with their more vertical structure, show a semi-plantigrade posture (which, however, is again supported by the foot pad). [5]
The cochlea of Numidotherium, which is built somewhat differently than that of today's elephants, turned out to be a further specialty. In Numidotherium, for example, this has one and a half turns, whereas in elephants it has two. In the basal area of the winding, there are deviations in detail, for example in the clear formation of the lamina spiralis secundaria, which is responsible, among other things, for the perception of certain frequencies. Researchers therefore suspect that Numidotherium was more likely to perceive sounds in the higher frequency range, in contrast to today's elephants, which can also communicate in infrasound. The lower number of coils of the cochlea in Numidotherium also suggests a more limited auditory perception. [10]
Proboscidea is a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Three species of elephant are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant.
Catopsbaatar is a genus of multituberculate, an extinct order of rodent-like mammals. It lived in what is now Mongolia during the late Campanian age of the Late Cretaceous epoch, about 72 million years ago. The first fossils were collected in the early 1970s, and the animal was named as a new species of the genus Djadochtatherium in 1974, D. catopsaloides. The specific name refers to the animal's similarity to the genus Catopsalis. The species was moved to the genus Catopsalis in 1979, and received its own genus in 1994. Five skulls, one molar, and one skeleton with a skull are known; the last is the genus' most complete specimen. Catopsbaatar was a member of the family Djadochtatheriidae.
Moeritherium is an extinct genus of basal proboscideans from the Eocene of North and West Africa. It was related to elephants and, more distantly, to sea cows and hyraxes.
Deinotheriidae is a family of prehistoric elephant-like proboscideans that lived during the Cenozoic era, first appearing in Africa, then spreading across southern Asia (Indo-Pakistan) and Europe. During that time, they changed very little, apart from growing much larger in size; by the late Miocene, they had become the largest land animals of their time. Their most distinctive features were their lack of upper tusks and downward-curving tusks on the lower jaw.
Prodeinotherium is an extinct representative of the family Deinotheriidae that lived in Africa, Europe, and Asia in the early and middle Miocene. Prodeinotherium, meaning "before terrible beast", was first named in 1930, but soon after, the only species in it, P. hungaricum, was reassigned to Deinotherium. During the 1970s, however, the two genera were once again separated, with Prodeinotherium diagnosed to include Deinotherium bavaricum, Deinotherium hobleyi, and Deinotherium pentapotamiae, which were separated based on geographic location. The three species are from Europe, Africa, and Asia, respectively. However, because of usage of few characters to separate them, only one species, P. bavaricum, or many more species, including P. cuvieri, P. orlovii, and P. sinense may be possible.
Paraceratherium is an extinct genus of hornless rhinocerotoids belonging to the family Paraceratheriidae. It is one of the largest terrestrial mammals that has ever existed and lived from the early to late Oligocene epoch. The first fossils were discovered in what is now Pakistan, and remains have been found across Eurasia between China and the Balkans. Paraceratherium means "near the hornless beast", in reference to Aceratherium, the genus in which the type species P. bugtiense was originally placed.
Pyrotherium is an extinct genus of South American ungulate, of the order Pyrotheria, that lived in what is now Argentina and Bolivia, during the Late Oligocene. It was named Pyrotherium because the first specimens were excavated from an ancient volcanic ash deposit. Fossils of the genus have been found in the Deseado and Sarmiento Formations of Argentina and the Salla Formation of Bolivia.
Numidotheriidae is an extinct family of primitive proboscideans that lived from the late Paleocene to the early Oligocene periods of North Africa.
Daouitherium is an extinct genus of early proboscideans that lived during the early Eocene some 55 million years ago in North Africa.
Phosphatherium escuillei is a basal proboscidean that lived from the Late Paleocene to the early stages of the Ypresian age. Research has suggested that Phosphatherium existed during the Eocene period.
Palaeomastodon is an extinct genus within the elephant order Proboscidea. Its fossils have been extracted from Oligocene strata conventionally dated to 33.9-23.03 million years old. Usually considered an ancestor or near-ancestor of elephants or mastodons as a member of Elephantiformes it lived in marshes and fluvial-deltaic environments of what is now Egypt, Ethiopia, Libya, and Saudi Arabia.
Notiomastodon is an extinct genus of gomphothere proboscidean, endemic to South America from the Pleistocene to the beginning of the Holocene. Notiomastodon specimens reached a size similar to that of the modern Asian elephant, with a body mass of 3-4 tonnes. Like other brevirostrine gomphotheres such as Cuvieronius and Stegomastodon, Notiomastodon had a shortened lower jaw and lacked lower tusks, unlike more primitive gomphotheres like Gomphotherium.
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.
Karenites is an extinct genus of therocephalian therapsids from the Late Permian of Russia. The only species is Karenites ornamentatus, named in 1995. Several fossil specimens are known from the town of Kotelnich in Kirov Oblast.
Homogalax is an extinct genus of tapir-like odd-toed ungulate. It was described on the basis of several fossil finds from the northwest of the United States, whereby the majority of the remains come from the state of Wyoming. The finds date to the Lower Eocene between 56 and 48 million years ago. In general, Homogalax was very small, only reaching the weight of today's peccaries, with a maximum of 15 kg. Phylogenetic analysis suggests the genus to be a basal member of the clade that includes today's rhinoceros and tapirs. In contrast to these, Homogalax was adapted to fast locomotion.
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.P
Elephantiformes is a suborder within the order Proboscidea. Members of this group are primitively characterised by the possession of upper tusks, an elongated mandibular symphysis and lower tusks, and the retraction of the facial region of the skull indicative of the development of a trunk. The earliest known member of the group, Dagbatitherium is known from the Eocene (Lutetian) of Togo, which is only known from isolated teeth, while other primitive elephantiforms like Phiomia and Palaeomastodon are known from the Early Oligocene onwards. Phiomia and Palaeomastodon are often collectively referred to as "palaeomastodonts" and assigned to the family Palaeomastodontidae. Most diversity of the group is placed in the subclade Elephantimorpha, which includes mastodons, as well as modern elephants and gomphotheres (Elephantida). It is disputed as to whether Phiomia is closely related to both Mammutidae and Elephantida with Palaeomastodon being more basal, or if Palaeomastodon is closely related to Mammutidae and Phiomia more closely related to Elephantida.
Arcanotherium is an extinct genus of early proboscidean belonging to the family Numidotheriidae that lived in north Africa during the late Eocene/early Oligocene interval.
Kerberos ("Cerberus") is an extinct genus of hyainailourid hyaenodonts in the subfamily Hyainailourinae, that lived in Europe. It contains the single species Kerberos langebadreae.
Periptychus is an extinct genus of mammal belonging to the family Periptychidae. It lived from the Early to Late Paleocene and its fossil remains have been found in North America.