Djebelemur

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Djebelemur
Temporal range: Late Early or Early Middle Eocene
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Strepsirrhini
Family: Djebelemuridae
Genus: Djebelemur
Hartenberger & Marandat, 1992
Species:
D. martinezi
Binomial name
Djebelemur martinezi
Hartenberger & Marandat, 1992

Djebelemur is an extinct genus of early strepsirrhine primate from the late early or early middle Eocene period from the Chambi locality in Tunisia. Although they probably lacked a toothcomb, a specialized dental structure found in living lemuriforms (lemurs and lorisoids), they are thought to be a related stem group. The one recognized species, Djebelemur martinezi, was very small, approximately 100 g (3.5 oz).

Contents

Djebelemur is one of five genera of djebelemurids, others including 'Anchomomys' milleri , a fossil primate formerly thought to be related to other members of the genus Anchomomys , a group of Eocene European adapiform primates. Following its discovery and description in 1992, Djebelemur was thought to be either a relative of European adapiforms or an early simian, mostly due to the fragmentary nature of the lower jaw and two isolated upper molars that may not belong with the jaw. By 2006, it was viewed as a stem lemuriform.

Etymology

The name "Djebelemur" derives from the mountainous outcrops in which the first specimens were found: Arabic djebel means "mountain". The species was named in honor of geologist C. Martinez, who was the guide and host for the field party that made the discovery. [1]

Evolutionary history

The fragmentary remains of Djebelemur martinezi are morphologically similar to those of 'Anchomomys' milleri , [lower-alpha 1] a fossil primate originally described as a type of cercamoniine (early adapiforms found in the northern continents). [4] Despite significant differences in age, with D. martinezi dating to the late early or early middle Eocene (~52–46 million years ago) [5] [6] and 'A.' milleri dating to the late Eocene (36 million years ago), [7] they form a clade [8] and are both grouped under the family Djebelemuridae. [9]

Djebelemurids appear to be a primitive sister group to lemuriforms and the more closely related Plesiopithecus , all of which form an Afro-Arabian clade that excludes the adapids or notharctids from the northern continents. [4] [8] Both are considered stem lemuriforms because despite sharing a number of traits, it is suspected that they lacked a toothcomb, a dental feature unique to lemuriforms. [2]

General phylogeny of strepsirrhines [10]
Primates  
  Haplorhines  

Simians (monkeys, apes, & humans)

Tarsiers

Omomyiforms

  Strepsirrhines  

Adapiforms

Azibiids

Djebelemurids

Plesiopithecus

Lemuriforms (lemurs, lorises, and galagos)

Holarctic clade
Afro-Arabian
clade
An Afro-Arabian clade of early strepsirrhines is thought to have given rise modern lemurs, lorises, and galagos. Azibiid taxonomy is still disputed.

The confirmed presence of djebelemurids in Eocene Africa is important because it proves that lemuriforms evolved their toothcomb in Africa and differentiated there. [4] This Afro-Arabian clade of stem lemuriforms are thought to have arrived in Africa too early to have descended from the Holarctic adapiforms. All dental similarities between the Afro-Arabian clade and European anchomomyins may be due to convergent evolution because the traits appear in the early Eocene (Ypresian stage) in the relatively poor fossil record of Africa long before they appear in the fossil-rich deposits of Europe during the mid-Lutetian. [8] This ancient stem lineage of lemuriform primates in Africa possibly descended from an early Asian branch of adapiforms [11] such as a primitive branch of cercamoniines predating Donrussellia (one of the oldest European adapiforms). [12]

Taxonomy

Djebelemur martinezi was first described in 1992 by Hartenberger and Marandat. The description was based on a lower jaw fragment and two isolated upper molars found in Tunisia at the Chambi locality, which date to the late early or early middle Eocene. [5] [6] It was described as an adapiform, possibly related to the cercamoniine branch, [4] with the informal suggestion that it might merit its own subfamily, "Djebelemurinae". [2] This interpretation was based on their support of the hypothesis favored by paleoanthropologist Philip D. Gingerich and others that simians (monkeys, apes, and humans) were descended from African adapids, which in turn were descended from the adapids of Europe. [4] [6] They based their assumptions of simian relations on the two isolated upper molars, which are now seen as being incompatible with the lower dentition on the jaw. The upper molars were highly bunodont (having cusps that are separate and rounded)—a trait seen in simians—whereas the lower molars were crested. [6] [13] No definitive upper teeth for Djebelemur are known, but could yield surprises if found. [13]

In 1994, paleoanthropologist Marc Godinot described Djebelemur as an early simian, along with Algeripithecus , [14] once considered a basal simian, but now considered to be a distant stem lemuriform (lemurs and lorisoids). [15] Godinot saw similarities between Djebelemur and early simians, as well as cercamoniines, [16] but also noted issues of premolar and molar compaction that set it apart from European adapiforms. [4] In 1997, Hartenberger continued to favor adapiform affinities, [16] while in 1998, Godinot considered affinities with lemuriforms ("crown strepsirrhines") while still favoring simian affinities, [8] particularly with oligopithecids. [6] By 2006, Godinot accepted Djebelemur as a stem lemuriform, admitting that he was misled by the lack of a toothcomb—a distinguishing dental feature of living lemuriform primates—despite other dental similarities. He also noted that the lower molars of lemuriforms and simians can be difficult to distinguish. [6] As of 2010, Djebelemur is still considered a stem lemuriform in the family Djebelemuridae, although its infraorder is unnamed. [9]

Anatomy and physiology

Djebelemur martinezi was a tiny primate, weighing approximately 100 g (3.5 oz). [16] It was thought to lack a toothcomb since its canine teeth were only moderately reduced. [17]

Notes

  1. In 2006, Godinot noted that 'Anchomomys' milleri requires a new generic designation because it does not belong in the genus Anchomomys , which contains cercamoniine adapiforms found in Europe. [2] Both djebelemurids share more characteristics—and therefore share a close phylogenetic relationship—with African lemuriforms than Eocene European cercamoniines. [3]

Related Research Articles

Strepsirrhini Suborder of primates which includes lemurs, galagos, pottos and lorises

Strepsirrhini or Strepsirhini is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. Collectively they are referred to as strepsirrhines. Also belonging to the suborder are the extinct adapiform primates which thrived during the Eocene in Europe, North America, and Asia, but disappeared from most of the Northern Hemisphere as the climate cooled. Adapiforms are sometimes referred to as being "lemur-like", although the diversity of both lemurs and adapiforms does not support this comparison.

Prosimian Obsolete primate taxon

Prosimians are a group of primates that includes all living and extinct strepsirrhines, as well as the haplorhine tarsiers and their extinct relatives, the omomyiforms, i.e. all primates excluding the simians. They are considered to have characteristics that are more "primitive" than those of simians.

Lemuriformes Infraorder of primates

Lemuriformes is an infraorder of primate that falls under the suborder Strepsirrhini. It includes the lemurs of Madagascar, as well as the galagos and lorisids of Africa and Asia, although a popular alternative taxonomy places the lorisoids in their own infraorder, Lorisiformes.

Notharctinae Extinct subfamily of mammals

Notharctinae is an extinct subfamily of primates that were common in North America during the early and middle Eocene. The six genera that make up the group contain species that are among the most primitive of the adapiform group, which is one of the most primitive groups of primates. The evolutionary history of this subfamily has been comparatively well documented and has been used to argue for evolutionary gradualism. Though it is generally accepted that adapiforms gave rise to modern day lemurs and lorises, it is not currently known which branch of Adapiformes these living species are most closely related to. Notharctines became extinct in the middle Eocene, most likely because of a combination of factors including climatic change and competition with other North American primates.

<i>Pelycodus</i> Extinct genus of primates

Pelycodus is an extinct genus of adapiform primate that lived during the early Eocene (Wasatchian) period in Europe and North America, particularly Wyoming and New Mexico. It is very closely related to Cantius and may even be its subgenus. It may also have given rise to the Middle Eocene Uintan primate Hesperolemur, although this is controversial. From mass estimates based on the first molar, the two species, P. jarrovii and P. danielsae, weighed 4.5 kg and 6.3 kg respectively and were frugivores with an arboreal, quadrupedal locomotion.

Toothcomb Dental structure found in some mammals

A toothcomb is a dental structure found in some mammals, comprising a group of front teeth arranged in a manner that facilitates grooming, similar to a hair comb. The toothcomb occurs in lemuriform primates, treeshrews, colugos, hyraxes, and some African antelopes. The structures evolved independently in different types of mammals through convergent evolution and varies both in dental composition and structure. In most mammals the comb is formed by a group of teeth with fine spaces between them. The toothcombs in most mammals include incisors only, while in lemuriform primates they include incisors and canine teeth that tilt forward at the front of the lower jaw, followed by a canine-shaped first premolar. The toothcombs of colugos and hyraxes take a different form with the individual incisors being serrated, providing multiple tines per tooth.

Altiatlasius is an extinct genus of mammal, which may have been the oldest known primate, dating to the Late Paleocene from Morocco. The only species, Altiatlasius koulchii, was described in 1990.

<i>Europolemur</i> Extinct genus of primates

Europolemur is a genus of adapiform primates that lived in Europe during the middle Eocene.

Bugtilemur is an extinct genus of Strepsirhine primate belonging to the adapiform family Ekgmowechashalidae.It is represented by only one species, B. mathesoni, which was found in the Chitarwata Formation of Pakistan.

Algeripithecus is an extinct genus of early fossil primate, weighing approximately 65 to 85 grams. Fossils have been found in Algeria dating from 50 to 46 million years ago.

Evolution of lemurs History of primate evolution on Madagascar

Lemurs, primates belonging to the suborder Strepsirrhini which branched off from other primates less than 63 million years ago, evolved on the island of Madagascar, for at least 40 million years. They share some traits with the most basal primates, and thus are often confused as being ancestral to modern monkeys, apes, and humans. Instead, they merely resemble ancestral primates.

Azibiidae is an extinct family of fossil primate from the late early or early middle Eocene from the Glib Zegdou Formation in the Gour Lazib area of Algeria. They are thought to be related to the living toothcombed primates, the lemurs and lorisoids, although paleoanthropologists such as Marc Godinot have argued that they may be early simians. It includes the genera Azibius and Algeripithecus, the latter of which was originally considered the oldest known simian, not a strepsirrhine.

Azibius is an extinct genus of fossil primate from the late early or early middle Eocene from the Glib Zegdou Formation in the Gour Lazib area of Algeria. They are thought to be related to the living toothcombed primates, the lemurs and lorisoids, although paleoanthropologists such as Marc Godinot have argued that they may be early simians. Originally described as a type of plesiadapiform, its fragmentary remains have been interpreted as a hyopsodontid, an adapid, and a macroscelidid. Less fragmentary remains discovered between 2003 and 2009 demonstrated a close relationship between Azibius and Algeripithecus, a fossil primate once thought to be the oldest known simian. Descriptions of the talus in 2011 have helped to strengthen support for the strepsirrhine status of Azibius and Algeripithecus, which would indicate that the evolutionary history of lemurs and their kin is rooted in Africa.

Djebelemuridae is an extinct family of early strepsirrhine primates from Africa. It consists of five genera. The organisms in this family were exceptionally small, and were insectivores. This family dates to the early to late Eocene. Although they gave rise to the crown strepsirrhines, which includes today's lemurs and lorisoids, they lacked the toothcomb that identifies that group.

<i>Afrasia djijidae</i> Eocene fossil primate from Myanmar

Afrasia djijidae is a fossil primate that lived in Myanmar approximately 37 million years ago, during the late middle Eocene. The only species in the genus Afrasia, it was a small primate, estimated to weigh around 100 grams (3.5 oz). Despite the significant geographic distance between them, Afrasia is thought to be closely related to Afrotarsius, an enigmatic fossil found in Libya and Egypt that dates to 38–39 million years ago. If this relationship is correct, it suggests that early simians dispersed from Asia to Africa during the middle Eocene and would add further support to the hypothesis that the first simians evolved in Asia, not Africa. Neither Afrasia nor Afrotarsius, which together form the family Afrotarsiidae, is considered ancestral to living simians, but they are part of a side branch or stem group known as eosimiiforms. Because they did not give rise to the stem simians that are known from the same deposits in Africa, early Asian simians are thought to have dispersed from Asia to Africa more than once prior to the late middle Eocene. Such dispersals from Asia to Africa also were seen around the same time in other mammalian groups, including hystricognathous rodents and anthracotheres.

Plesiopithecus is an extinct genus of early strepsirrhine primate from the late Eocene.

<i>Afrotarsius</i> Extinct genus of primates

Afrotarsius is a primate found in the Paleogene of Africa.

The Lignites de Soissonais is a geologic formation in the Var, Marne departments of France. It preserves fossils dating back to the Ypresian stage of the Eocene period.

Sivaladapis is a genus of adapiform primate that lived in Asia during the middle Miocene.

'Anchomomys' milleri is an extinct primate related to lemuriforms that lived in Africa during the early late Eocene. It was originally thought to be a member of the European genus Anchomomys, but was later aligned with the djebelemurids, although a new generic name needs to be assigned.

References

  1. Hartenberger & Marandat 1992, p. 10.
  2. 1 2 3 Godinot 2006, p. 452.
  3. Godinot 2006, p. 453.
  4. 1 2 3 4 5 6 Godinot 2010, p. 325.
  5. 1 2 Godinot 2010, p. 320.
  6. 1 2 3 4 5 6 Godinot 2006, p. 447.
  7. Simons 1997, p. 180.
  8. 1 2 3 4 Seiffert et al. 2005, p. 11400.
  9. 1 2 Godinot 2010, p. 324.
  10. Tabuce et al. 2009, pp. 4091–4092.
  11. Godinot 2006, pp. 461–462.
  12. Godinot 2006, pp. 454–455.
  13. 1 2 Godinot 2010, p. 326.
  14. Godinot & Mahboubi 1994, p. 358.
  15. Tabuce et al. 2009, p. 4087.
  16. 1 2 3 Gebo 2002, p. 35.
  17. Tabuce et al. 2009, p. 4092.

Literature cited

  • Gebo, D.L. (2002). "Chapter 4: Adapiformes: Phylogeny and adaptation". In Hartwig, W.C (ed.). The Primate Fossil Record. Cambridge University Press. pp. 21–43. ISBN   0-521-66315-6.
  • Godinot, M.; Mahboubi, M. (1994). "Les petits primates simiiformes de Glib Zegdou (Eocène inférieur à moyen d'Algérie)" [Small simiiform primates from Glib Zegdou (early to middle Eocene, Algeria)]. Comptes rendus de l'Académie des sciences. Série 2. Sciences de la terre et des planètes. (in French). 319 (3): 357–364. ISSN   1251-8050.
  • Godinot, M. (2006). "Lemuriform origins as viewed from the fossil record". Folia Primatologica. 77 (6): 446–464. doi:10.1159/000095391. PMID   17053330. S2CID   24163044.
  • Godinot, M. (2010). "Chapter 19: Paleogene Prosimians". In Werdelin, L.; Sanders, W.J (eds.). Cenozoic Mammals of Africa. University of California Press. ISBN   978-0-520-25721-4.
  • Hartenberger, J.-L.; Marandat, B. (1992). "A new genus and species of an early Eocene primate from North Africa". Human Evolution. 7: 9–16. doi:10.1007/BF02437474. S2CID   84275103.
  • Seiffert, E.R.; Simons, E.L.; Ryan, T.M.; Attia, Y. (2005). "Additional remains of Wadilemur elegans, a primitive stem galagid from the late Eocene of Egypt" (PDF). Proceedings of the National Academy of Sciences. 102 (32): 11396–11401. doi: 10.1073/pnas.0505310102 . PMC   1183603 . PMID   16087891.
  • Simons, E. L. (1997). "Discovery of the smallest Fayum Egyptian primates (Anchomomyini, Adapidae)". Proceedings of the National Academy of Sciences. 94 (1): 180–184. doi: 10.1073/pnas.94.1.180 . PMC   19275 . PMID   11038538.
  • Tabuce, R.; Marivaux, L.; Lebrun, R.; Adaci, M.; Bensalah, M.; Fabre, P. -H.; Fara, E.; Gomes Rodrigues, H.; Hautier, L.; Jaeger, J. -J.; Lazzari, V.; Mebrouk, F.; Peigne, S.; Sudre, J.; Tafforeau, P.; Valentin, X.; Mahboubi, M. (2009). "Anthropoid versus strepsirhine status of the African Eocene primates Algeripithecus and Azibius: Craniodental evidence". Proceedings of the Royal Society B: Biological Sciences. 276 (1676): 4087–4094. doi:10.1098/rspb.2009.1339. PMC   2821352 . PMID   19740889.