Azibius
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''Azibius'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of fossil primate from the late early or early middle
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
from the Glib Zegdou Formation in the Gour Lazib area of
Algeria ) , image_map = Algeria (centered orthographic projection).svg , map_caption = , image_map2 = , capital = Algiers , coordinates = , largest_city = capital , relig ...
. They are thought to be related to the living
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 ( ...
ed primates, the
lemur Lemurs ( ) (from Latin ''lemures'' – ghosts or spirits) 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 exist ...
s and lorisoids (known as
strepsirrhines 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. Collec ...
), although
paleoanthropologists Paleoanthropology or paleo-anthropology is a branch of paleontology and anthropology which seeks to understand the early development of anatomically modern humans, a process known as hominization, through the reconstruction of evolutionary kinship ...
such as
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 o ...
have argued that they may be 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 monkeys (Platyrrhini) and Catarrhi ...
s (monkeys and apes). Originally described as a type of
plesiadapiform Plesiadapiformes (" Adapid-like" or "near Adapiformes") is a group of Primates, a sister of the Dermoptera. While none of the groups normally directly assigned to this group survived, the group appears actually not to be literally extinct (in ...
(an extinct group of arboreal
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s considered to be a
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 t ...
to the
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 huma ...
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
), its fragmentary remains have been interpreted as a hyopsodontid (a type of extinct
condylarth Condylarthra is an informal group – previously considered an order – of extinct placental mammals, known primarily from the Paleocene and Eocene epochs. They are considered early, primitive ungulates. It is now largely considered to be a wast ...
), an adapid (an extinct type of
adapiform Adapiformes is a group of early primates. Adapiforms radiated throughout much of the northern continental mass (now Europe, Asia and North America), reaching as far south as northern Africa and tropical Asia. They existed from the Eocene to the M ...
primate from Europe), and a macroscelidid (
elephant shrew Elephant shrews, also called jumping shrews or sengis, are small insectivorous mammals native to Africa, belonging to the family Macroscelididae, in the order Macroscelidea. Their traditional common English name "elephant shrew" comes from a pe ...
s). Less fragmentary remains discovered between 2003 and 2009 demonstrated a close relationship between ''Azibius'' and ''
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 th ...
'', a fossil primate once thought to be the oldest known simian. Descriptions of the talus (ankle bone) in 2011 have helped to strengthen support for the strepsirrhine status of ''Azibius'' and ''Algeripithecus'', which would indicate that the
evolutionary history of lemurs 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 ...
and their kin is rooted in Africa. ''Azibius trerki'' is the only named species, although a few teeth and a talus (ankle bone) of a larger, unnamed species (
cf. The abbreviation ''cf.'' (short for the la, confer/conferatur, both meaning "compare") is used in writing to refer the reader to other material to make a comparison with the topic being discussed. Style guides recommend that ''cf.'' be used onl ...
 ''Azibius'' sp.) have also been found. ''A. trerki'' is estimated to have weighed , while cf. ''Azibius'' sp. was larger, weighing approximately . Based on the fragmentary fossils, both are thought to have been
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
and agile
arboreal Arboreal locomotion is the Animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. Th ...
quadrupeds Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
.


Evolutionary history and taxonomy

For years following its discovery, ''Azibius'' was difficult to classify. Originally described by Jean Sudre in 1975 as a possible 'paromomyiform' (a type of
plesiadapiform Plesiadapiformes (" Adapid-like" or "near Adapiformes") is a group of Primates, a sister of the Dermoptera. While none of the groups normally directly assigned to this group survived, the group appears actually not to be literally extinct (in ...
), ''A. trerki'' was also interpreted as a hyopsodontid by paleoanthropologist Frederick S. Szalay that same year. The following year, 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 ...
reclassified it as an adapid. Throughout the 1990s and 2000s, debates over its classification continued, with some researchers suggesting it might be related to macroscelidids (
elephant shrew Elephant shrews, also called jumping shrews or sengis, are small insectivorous mammals native to Africa, belonging to the family Macroscelididae, in the order Macroscelidea. Their traditional common English name "elephant shrew" comes from a pe ...
s), while others supported initial interpretations as adapids or plesiadapiforms (particularly carpolestids). In 2006, 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 o ...
favored a relationship between ''Azibius'' and simians, but tentatively suggested ''Azibius'' may be more closely related to
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 ( ...
ed primates, which include all extant strepsirrhines. This latter view has gained increasing support with the reclassification of ''
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 th ...
'' (once considered a basal
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 monkeys (Platyrrhini) and Catarrhi ...
) as a closely related azibiid. The
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
of ''Algeripithecus'' indicates it had an inclined
canine tooth 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 however ...
, similar to that found in toothcombed primates. Although the anterior dentition of azibiids is unknown, they may have possessed a toothcomb, indicating an ancient
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
lineage of lemuriform primates in Africa, possibly descended from an early Asian branch of adapiforms such as a primitive branch of cercamoniines predating ''
Donrussellia ''Donrussellia'' is a genus of adapiform primate that lived in Europe during the early 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 Per ...
'' (one of the oldest European adapiforms). ''Tabelia hammadae'', which was also considered to be one of the oldest known simians along with ''Algeripithecus'', was shown to be a
synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
of ''Azibius'' when more complete fossils were discovered at Gour Lazib between 2003 and 2009. Likewise, the second upper molar (M2) of ''Dralestes hammadaensis'' have been reinterpreted as being the upper fourth premolar (P4) of ''Azibius'' and has been considered a synonym. However, in 2010, Godinot cautiously suggested that ''Dralestes'' may be a synonym of ''Algeripithecus'' based on a blade-like premolar. He also reasserted his view that ''Algeripithecus'' was a simian based on its upper molar morphology and hypothesized that this applied to all azibiids, favoring his earlier view that they may be early simians instead of stem lemuriforms. In 2011, Marivaux ''et al.'' published an interpretation of recently discovered
talus bone The talus (; Latin for ankle or ankle bone), talus bone, astragalus (), or ankle bone is one of the group of foot bones known as the tarsus. The tarsus forms the lower part of the ankle joint. It transmits the entire weight of the body from the ...
s found at Gour Lazib, which they claimed were more similar to those of living strepsirrhines and extinct adapiforms, not simians, thus reinforcing the strepsirrhine status favored by Tabuce ''et al.'' two years earlier. The tali morphology also differed radically from those of plesiadapiforms, confirming that azibiids are true primates. Remains of a second, unnamed species of ''Azibius'', cf. ''Azibius'' sp., have been discovered in the HGL-50 layer at Gour Lazib. It is known for a few upper and lower teeth. These teeth are three times larger than those of ''A. trerki''. A larger right talus has also been found, and is assumed to belong to this new species.


Anatomy and physiology

All known azibiids were small-bodied primates. ''A. trerki'' is estimated to have weighed between , nearly twice the weight of the only other named azibiid, ''Algeripithecus'', and was comparable in size to a
gray mouse lemur The gray mouse lemur (''Microcebus murinus''), grey mouse lemur or lesser mouse lemur, is a small lemur, a type of strepsirrhine primate, found only on the island of Madagascar. Weighing , it is the largest of the mouse lemurs (genus ''Microceb ...
(''Microcebus murinus''). The unnamed species, cf. ''Azibius'' sp., is estimated to have weighed and been comparable in size to a
sportive lemur The sportive lemurs are the medium-sized primates that make up the family Lepilemuridae. The family consists of only one extant genus, ''Lepilemur''. They are closely related to the other lemurs and exclusively live on the island of Madagascar. ...
(''Lepilemur''). ''Azibius'' also had a short
rostrum Rostrum may refer to: * Any kind of a platform for a speaker: **dais **pulpit * Rostrum (anatomy), a beak, or anatomical structure resembling a beak, as in the mouthparts of many sucking insects * Rostrum (ship), a form of bow on naval ships * Ros ...
and very large
orbits In celestial mechanics, an orbit is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an object or position in space such as a p ...
.


Distribution

''Azibius'' has only been found in the Glib Zegdou Formation of the Gour Lazib area in southwest Algeria. The Glib Zegdou Formation dates to the late early or early middle Eocene, particularly the late
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 ...
or early
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 Midd ...
.


Behavior

Judging from the properties of its talus, ''Azibius'' was probably an arboreal
quadruped Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
, capable of leaping and climbing in trees, very similar to living cheirogaleid lemurs. Based on what can be inferred from the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
ry remains, ''Azibius'' is thought to have had large eyes and extra
vibrissae Vibrissae (; singular: vibrissa; ), more generally called Whiskers, are a type of stiff, functional hair used by mammals to sense their environment. These hairs are finely specialised for this purpose, whereas other types of hair are coarser ...
, which suggests it was
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
.


Notes


References


Literature cited

* * * * ** {{Taxonbar, from=Q4832562 Prehistoric strepsirrhines Eocene primates Eocene mammals of Africa Prehistoric primate genera Fossil taxa described in 1975