Djebelemur
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''Djebelemur'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
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 is an order (biology), order of mammals, which is further divided into the Strepsirrhini, strepsirrhines, which include lemurs, galagos, and Lorisidae, lorisids; and the Haplorhini, haplorhines, which include Tarsiiformes, tarsiers a ...
from the late early or early middle
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
period from the Chambi locality in
Tunisia Tunisia, officially the Republic of Tunisia, is a country in the Maghreb region of North Africa. It is bordered by Algeria to the west and southwest, Libya to the southeast, and the Mediterranean Sea to the north and east. Tunisia also shares m ...
. Although they probably lacked a
toothcomb A toothcomb (also called a tooth comb or dental comb) is a Dentition, 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 Lem ...
, a specialized dental structure found in living lemuriforms (
lemur Lemurs ( ; from Latin ) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea ( ), divided into 8 Family (biology), families and consisting of 15 genera and around 100 existing species. They are Endemism, ...
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 The simians, anthropoids, or higher primates are an infraorder (Simiiformes ) of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders New World monkey, Platyrrhini (New World mon ...
, mostly due to the fragmentary nature of the lower jaw and two isolated upper
molars The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
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 Central Semitic languages, Central Semitic language of the Afroasiatic languages, Afroasiatic language family spoken primarily in the Arab world. The International Organization for Standardization (ISO) assigns lang ...
''djebel'' means "mountain". The species was named in honor of
geologist A geologist is a scientist who studies the structure, composition, and History of Earth, history of Earth. Geologists incorporate techniques from physics, chemistry, biology, mathematics, and geography to perform research in the Field research, ...
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 ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
(~52–46 million years ago) and ''A.' milleri'' dating to the late Eocene (36 million years ago), they form a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
and are both grouped under the family Djebelemuridae. Djebelemurids appear to be a primitive
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
to lemuriforms and the more closely related ''
Plesiopithecus ''Plesiopithecus'' is an extinct genus of early strepsirrhine primate from the late Eocene. Anatomy Originally described from the right mandible (lower jaw), its confusing anatomy resulted in it being classified as an ape—its name translates " ...
'', 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 called a tooth comb or dental comb) is a Dentition, 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 Lem ...
, 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 comm ...
because the traits appear in the early Eocene (
Ypresian In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...
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 (stratigraphy), stage or age (geology), 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 ...
. This ancient
stem Stem or STEM most commonly refers to: * Plant stem, a structural axis of a vascular plant * Stem group * Science, technology, engineering, and mathematics Stem or STEM can also refer to: Language and writing * Word stem, part of a word respon ...
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 tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
found in
Tunisia Tunisia, officially the Republic of Tunisia, is a country in the Maghreb region of North Africa. It is bordered by Algeria to the west and southwest, Libya to the southeast, and the Mediterranean Sea to the north and east. Tunisia also shares m ...
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 and others that
simian The simians, anthropoids, or higher primates are an infraorder (Simiiformes ) of primates containing all animals traditionally called monkeys and apes. More precisely, they consist of the parvorders New World monkey, Platyrrhini (New World mon ...
s (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 described ''Djebelemur'' as an early simian, along with ''
Algeripithecus ''Algeripithecus'' is an extinct genus of early List of fossil primates, 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 sim ...
'', once considered a basal simian, but now considered to be a distant stem lemuriform (
lemur Lemurs ( ; from Latin ) are Strepsirrhini, wet-nosed primates of the Superfamily (biology), superfamily Lemuroidea ( ), divided into 8 Family (biology), families and consisting of 15 genera and around 100 existing species. They are Endemism, ...
s and lorisoids). Godinot saw similarities between ''Djebelemur'' and early simians, as well as cercamoniines, but also noted issues of
premolar The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
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, parti ...
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 called a tooth comb or dental comb) is a Dentition, 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 Lem ...
—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 () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomic rank used in the classific ...
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, dogteeth, eye teeth, vampire teeth, or fangs, are the relatively long, pointed teeth. In the context of the upper jaw, they are also known as '' fangs''. They can appear more fl ...
were only moderately reduced.


Notes


References


Literature cited

* * * * * * * * ** {{Taxonbar, from=Q3032855 Djebelemuridae Eocene primates Fossil taxa described in 1992 Eocene mammals of Africa Monotypic prehistoric primate genera