Djebelemur
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''Djebelemur'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of early
strepsirrhine 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. Colle ...
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 ...
from the late early or early middle
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
period from the Chambi locality in
Tunisia ) , image_map = Tunisia location (orthographic projection).svg , map_caption = Location of Tunisia in northern Africa , image_map2 = , capital = Tunis , largest_city = capital , ...
. Although they probably lacked a
toothcomb A toothcomb (also tooth comb or dental comb) 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 ( ...
, a specialized dental structure found in living lemuriforms (
lemur Lemurs ( ) (from Latin ''lemures'' – ghosts or spirits) are wet-nosed primates of the superfamily Lemuroidea (), divided into 8 families and consisting of 15 genera and around 100 existing species. They are endemic to the island of Madagas ...
s and lorisoids), they are thought to be a related
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. ...
. The one recognized species, ''Djebelemur martinezi'', was very small, approximately . ''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 The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
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 Arabic (, ' ; , ' or ) is a Semitic language spoken primarily across the Arab world.Semitic languages: an international handbook / edited by Stefan Weninger; in collaboration with Geoffrey Khan, Michael P. Streck, Janet C. E.Watson; Walter ...
''djebel'' means "mountain". The species was named in honor of
geologist A geologist is a scientist who studies the solid, liquid, and gaseous matter that constitutes Earth and other terrestrial planets, as well as the processes that shape them. Geologists usually study geology, earth science, or geophysics, althoug ...
C. Martinez, who was the guide and host for the field party that made the discovery.


Evolutionary history

The fragmentary remains of ''Djebelemur martinezi'' are morphologically similar to those of '' 'Anchomomys' milleri'', a fossil primate originally described as a type of cercamoniine (early adapiforms found in the northern continents). Despite significant differences in age, with ''D. martinezi'' dating to the late early or early middle
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
(~52–46 million years ago) and ''A.' milleri'' dating to the late Eocene (36 million years ago), they form a clade and are both grouped under the family Djebelemuridae. 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. Both are considered stem lemuriforms because despite sharing a number of traits, it is suspected that they lacked a
toothcomb A toothcomb (also tooth comb or dental comb) 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 ( ...
, a dental feature unique to lemuriforms. The confirmed presence of djebelemurids in Eocene Africa is important because it proves that lemuriforms evolved their toothcomb in Africa and differentiated there. This Afro-Arabian clade of stem lemuriforms are thought to have arrived in Africa too early to have descended from the
Holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical reg ...
adapiforms. All dental similarities between the Afro-Arabian clade and European anchomomyins may be due to
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
because the traits appear in the early Eocene (
Ypresian In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The 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 The Lutetian is, in the geologic timescale, a stage or age in the Eocene. It spans the time between . The Lutetian is preceded by the Ypresian and is followed by the Bartonian. Together with the Bartonian it is sometimes referred to as the ...
. This ancient stem lineage of lemuriform primates in Africa possibly descended from an early Asian branch of adapiforms such as a primitive branch of cercamoniines predating '' Donrussellia'' (one of the oldest European adapiforms).


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 The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
found in
Tunisia ) , image_map = Tunisia location (orthographic projection).svg , map_caption = Location of Tunisia in northern Africa , image_map2 = , capital = Tunis , largest_city = capital , ...
at the Chambi locality, which date to the late early or early middle Eocene. It was described as an adapiform, possibly related to the cercamoniine branch, with the informal suggestion that it might merit its own subfamily, "Djebelemurinae". This interpretation was based on their support of the hypothesis favored by paleoanthropologist
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 ...
and others that simians (monkeys, apes, and humans) were descended from African adapids, which in turn were descended from the adapids of Europe. 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. No definitive upper teeth for ''Djebelemur'' are known, but could yield surprises if found. In 1994, paleoanthropologist
Marc Godinot Marc or MARC may refer to: People * Marc (given name), people with the first name * Marc (surname), people with the family name Acronyms * MARC standards, a data format used for library cataloging, * MARC Train, a regional commuter rail system of ...
described ''Djebelemur'' as an early simian, along with ''
Algeripithecus ''Algeripithecus'' is an extinct genus of early fossil primate, weighing approximately . Fossils have been found in Algeria dating from 50 to 46 million years ago. It was once commonly thought to be one of the oldest simian primates (a group t ...
'', once considered a basal simian, but now considered to be a distant stem lemuriform (
lemur Lemurs ( ) (from Latin ''lemures'' – ghosts or spirits) are wet-nosed primates of the superfamily Lemuroidea (), divided into 8 families and consisting of 15 genera and around 100 existing species. They are endemic to the island of Madagas ...
s and lorisoids). Godinot saw similarities between ''Djebelemur'' and early simians, as well as cercamoniines, but also noted issues of premolar and molar compaction that set it apart from European adapiforms. In 1997, Hartenberger continued to favor adapiform affinities, while in 1998, Godinot considered affinities with lemuriforms ("
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 ...
strepsirrhines") while still favoring simian affinities, particularly with oligopithecids. By 2006, Godinot accepted ''Djebelemur'' as a stem lemuriform, admitting that he was misled by the lack of a
toothcomb A toothcomb (also tooth comb or dental comb) 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 ( ...
—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. As of 2010, ''Djebelemur'' is still considered a stem lemuriform in the family Djebelemuridae, although its
infraorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
is unnamed.


Anatomy and physiology

''Djebelemur martinezi'' was a tiny primate, weighing approximately . It was thought to lack a toothcomb since its
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 ...
were only moderately reduced.


Notes


References


Literature cited

* * * * * * * * ** {{Taxonbar, from=Q3032855 Prehistoric strepsirrhines Eocene primates Fossil taxa described in 1992 Eocene mammals of Africa Prehistoric primate genera