''Danuvius guggenmosi'' is an extinct species of
great ape
The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the east ...
that lived 11.6 million years ago during the
Middle–
Late
Late may refer to:
* LATE, an acronym which could stand for:
** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia
** Local-authority trading enterprise, a New Zealand business law
** Local average treatment effect, ...
Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
in southern Germany. It is the
sole member of the genus ''Danuvius''. The area at this time was probably a
woodland
A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the ''plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (see ...
with a
seasonal
A season is a division of the year based on changes in weather, ecology, and the number of daylight hours in a given region. On Earth, seasons are the result of the axial parallelism of Earth's tilted orbit around the Sun. In temperate and po ...
climate. A male specimen was estimated to have weighed about , and two females . Both genus and species were
described in November 2019.
It is the first-discovered Late Miocene great ape with preserved
long bone
The long bones are those that are longer than they are wide. They are one of five types of bones: long, Short bone, short, Flat bone, flat, Irregular bone, irregular and Sesamoid bone, sesamoid. Long bones, especially the femur and tibia, are subj ...
s which could be used to reconstruct the limb anatomy and thus the locomotion of contemporary apes. Its discoverer, paleoanthropologist
Madelaine Böhme
Madelaine Böhme (born 1967) is a German palaeontologist and professor of palaeoclimatology at the University of Tübingen.
Böhme was born in 1967 in Plovdiv, Bulgaria. She studied at the Freiberg University of Mining and Technology and Leipzi ...
, says ''Danuvius'' had adaptations for both hanging in trees (
suspensory behavior Suspensory behaviour is a form of arboreal locomotion or a feeding behavior that involves hanging or suspension of the body below or among tree branches. This behavior enables faster travel while reducing path lengths to cover more ground when trave ...
) and walking on two legs (
biped
Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' a ...
alism)—whereas, among present-day great apes, humans are better adapted for the latter and the others the former.
''Danuvius'' thus had a method of locomotion unlike any previously known ape called "extended limb clambering," she says, walking directly along tree branches as well as using arms for suspending itself. The last common ancestor between humans and other apes possibly had a similar method of locomotion. However, paleoanthropologist Scott Williams and others say the fragmentary remains do not differ enough from other fossil apes to provide such a clue to the origins of bipedalism.
Taxonomy
![Tongrube Hammerschmiede 02](https://upload.wikimedia.org/wikipedia/commons/6/6c/Tongrube_Hammerschmiede_02.jpg)
The
genus name ''Danuvius'' is a reference to the
Celtic
Celtic, Celtics or Keltic may refer to:
Language and ethnicity
*pertaining to Celts, a collection of Indo-European peoples in Europe and Anatolia
**Celts (modern)
*Celtic languages
**Proto-Celtic language
* Celtic music
*Celtic nations
Sports Fo ...
–
Roman
Roman or Romans most often refers to:
*Rome, the capital city of Italy
*Ancient Rome, Roman civilization from 8th century BC to 5th century AD
*Roman people, the people of ancient Rome
*''Epistle to the Romans'', shortened to ''Romans'', a letter ...
river-god
Danuvius, a Roman name for the river
Danube
The Danube ( ; ) is a river that was once a long-standing frontier of the Roman Empire and today connects 10 European countries, running through their territories or being a border. Originating in Germany, the Danube flows southeast for , pa ...
, which flows through the region where the remains were found. The
specific name Specific name may refer to:
* in Database management systems, a system-assigned name that is unique within a particular database
In taxonomy, either of these two meanings, each with its own set of rules:
* Specific name (botany), the two-part (bino ...
''guggenmosi'' honours the amateur archaeologist
Sigulf Guggenmos (1941–2018), who discovered the clay pit in which ''Danuvius'' was found.
The remains of ''Danuvius'' were discovered in the
Hammerschmiede clay pit
The Hammerschmiede clay pit (German: Tongrube Hammerschmiede) is a fossil bearing locality in Pforzen, Bavaria, Germany most well known for the discovery of '' Danuvius guggenmosi'', the potentially earliest known bipedal ape. With an age of 11.6 ...
near the town of
Pforzen
Pforzen is a municipality in Ostallgäu, Bavaria.
Cemetery
Near Pforzen is an extended Alemannic cemetery that was in use from the 5th up to the 8th century. A total of 442 graves were excavated in two campaigns in 1991/2 and 1996. Two items bea ...
in southern Germany,
magnetostratigraphically dated to 11.62
million years ago
The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds.
Usage
Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
(mya) at the
Serravallian
The Serravallian is, in the geologic timescale, an List of time periods, age or a stage (stratigraphy), stage in the middle Miocene epoch (geology), Epoch/series (stratigraphy), Series, which spans the time between 13.82 annum, Ma and 11.63 Ma (m ...
-
Tortonian boundary (the
Astaracian–
Vallesian
The Vallesian age is a period of geologic time (11.6–9.0 Ma) within the Miocene used more specifically with European Land Mammal Ages. It precedes the Turolian age and follows the Astaracian age. The so-called Vallesian Crisis resulted in the e ...
boundary in
ELMA),
[ and were unearthed between 2015 and 2018.][ The ]holotype
A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
GPIT/MA/10000 comprises a partial skeleton with elements of the mouth, vertebrae, and long bone
The long bones are those that are longer than they are wide. They are one of five types of bones: long, Short bone, short, Flat bone, flat, Irregular bone, irregular and Sesamoid bone, sesamoid. Long bones, especially the femur and tibia, are subj ...
s. There are also three paratypes: an adult left femur
The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
(GPIT/MA/10001); an adult left femur, big toe, and teeth (GPIT/MA/10003); and juvenile teeth and a middle finger bone (GPIT/MA/10002). There are 37 specimens in total.[
Its tooth anatomy is most similar to that of other ]dryopithecine
Dryopithecini is an extinct tribe of Eurasian and African great apes that are believed to be close to the ancestry of gorillas, chimpanzees and humans. Members of this tribe are known as dryopithecines.
Taxonomy
* Tribe Dryopithecini †
** ''K ...
great ape
The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the east ...
s. Having both adaptations for hanging in trees ( suspensory locomotion) and standing on two legs (biped
Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' a ...
alism), ''Danuvius'' may have been very similar in locomotory methods to the last common ancestor between humans and other apes, which adds weight to the hypothesis that ape suspensory activity and human bipedalism both originated from a form capable of both.[ However, it is too early to draw more definitive conclusions because it is unclear how ''Danuvius'' is related to modern great apes, including humans.] Its discovery could also influence reconstructions of contemporary great ape limb anatomy and locomotion, which were previously by-and-large unknown.[
]
Description
''Danuvius'' was small and probably weighed on average . The holotype specimen, an adult male, was calculated, based on the sizes of the hip
In vertebrate anatomy, hip (or "coxa"Latin ''coxa'' was used by Celsus in the sense "hip", but by Pliny the Elder in the sense "hip bone" (Diab, p 77) in medical terminology) refers to either an anatomical region or a joint.
The hip region is ...
and knee
In humans and other primates, the knee joins the thigh with the leg and consists of two joints: one between the femur and tibia (tibiofemoral joint), and one between the femur and patella (patellofemoral joint). It is the largest joint in the hu ...
joints, to have weighed with a point estimate
In statistics, point estimation involves the use of sample data to calculate a single value (known as a point estimate since it identifies a point in some parameter space) which is to serve as a "best guess" or "best estimate" of an unknown popu ...
of . The adult female specimen GPIT/MA/10003 was calculated to be with a point estimate of , and the adult female GPIT/MA/10001 with a point estimate of . This is bigger than siamang
The siamang (, ; ''Symphalangus syndactylus'') is an arboreal, black-furred gibbon native to the forests of Indonesia, Malaysia, and Thailand. The largest of the gibbons, the siamang can be twice the size of other gibbons, reaching in height, a ...
s but much smaller than contemporary great apes;[ for example, male ]bonobo
The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee and less often the dwarf chimpanzee or gracile chimpanzee, is an endangered great ape and one of the two species making up the genus '' Pan,'' the other being the comm ...
s weigh and females .
The sex of the individuals was determined by the size of the canines
Canine may refer to:
Zoology and anatomy
* a dog-like Canid animal in the subfamily Caninae
** ''Canis'', a genus including dogs, wolves, coyotes, and jackals
** Dog, the domestic dog
* Canine tooth, in mammalian oral anatomy
People with the surn ...
, with males presumed to have had larger canines than females. Male dryopithecines are thought to have had an elongated face with the molars pushed more towards the front of the mouth. Like those of other dryopithecines, the molars of ''Danuvius'' were wide, and there was a broad length between the two cusps; however, the premolar
The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
s had three roots
A root is the part of a plant, generally underground, that anchors the plant body, and absorbs and stores water and nutrients.
Root or roots may also refer to:
Art, entertainment, and media
* ''The Root'' (magazine), an online magazine focusing ...
instead of two, and the canines were more vertically oriented rather than somewhat sticking out.[
''Danuvius'' is thought to have had a broad chest. It is the first recorded Miocene great ape to have had the ]diaphragm
Diaphragm may refer to:
Anatomy
* Thoracic diaphragm, a thin sheet of muscle between the thorax and the abdomen
* Pelvic diaphragm or pelvic floor, a pelvic structure
* Urogenital diaphragm or triangular ligament, a pelvic structure
Other
* Diap ...
located in the lower chest cavity
The thoracic cavity (or chest cavity) is the chamber of the body of vertebrates that is protected by the thoracic wall (rib cage and associated skin, muscle, and fascia). The central compartment of the thoracic cavity is the mediastinum. There ...
, as in ''Homo
''Homo'' () is the genus that emerged in the (otherwise extinct) genus ''Australopithecus'' that encompasses the extant species ''Homo sapiens'' ( modern humans), plus several extinct species classified as either ancestral to or closely relate ...
'', indicating an extended lower back and a greater number of functional lumbar vertebra
The lumbar vertebrae are, in human anatomy, the five vertebrae between the rib cage and the pelvis. They are the largest segments of the vertebral column and are characterized by the absence of the foramen transversarium within the transverse p ...
e. This may have caused lordosis (the normal curvature of the human spine) and moved the center of mass
In physics, the center of mass of a distribution of mass in space (sometimes referred to as the balance point) is the unique point where the weighted relative position of the distributed mass sums to zero. This is the point to which a force may ...
over the hips and legs, which implies some habitual bipedal activity.[
The robust finger and ]hypertrophied
Hypertrophy is the increase in the volume of an organ or tissue due to the enlargement of its component cells. It is distinguished from hyperplasia, in which the cells remain approximately the same size but increase in number.Updated by Linda J. ...
wrist and elbow bones indicate a strong grip and load bearing adaptations for the arms. The legs also show adaptations for load-bearing, especially at the hypertrophied knee joint
In humans and other primates, the knee joins the thigh with the leg and consists of two joints: one between the femur and tibia (tibiofemoral joint), and one between the femur and patella (patellofemoral joint). It is the largest joint in the hu ...
. There was likely limited ankle loading, and the ankle would have had a hinge-like function, being most stable if positioned perpendicularly to the leg as opposed to at an angle in apes. ''Danuvius'' was likely able to achieve a strong grip with its big toe
Toes are the digits (fingers) of the foot of a tetrapod. Animal species such as cats that walk on their toes are described as being ''digitigrade''. Humans, and other animals that walk on the soles of their feet, are described as being ''plant ...
s, unlike modern African great apes, which would have allowed it to grasp onto thinner trees. The limb proportions are most similar to those of bonobos.[
]
Adaptations for load bearing in both the arm and leg joints to this degree is unknown in any other primate. Plantigrade catarrhine monkeys lack the capacity for suspensory locomotion or to focus body weight over the knee joint; knuckle-walking apes lack strong big toes and thumbs, and have more robust finger bones; and both lack an extendable knee. Orangutan
Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s have a clambering motion too, but their knees lack weight-bearing ability.[
]
Paleoecology
The total anatomy of the limbs suggest ''Danuvius'' was capable of a seemingly unique manner of locomotion called "extended limb clambering". ''Danuvius'' likely walked along mildly inclined tree branches with its foot directly laid onto the branch, using its strong big toes for grasping. The strong knee joint would have provided balance while walking by counteracting torque
In physics and mechanics, torque is the rotational equivalent of linear force. It is also referred to as the moment of force (also abbreviated to moment). It represents the capability of a force to produce change in the rotational motion of th ...
s, and the strong hands would have carried out a similar function during suspension or palm-walking. Extended limb clambering emphasizes knee extension and lordosis, as well as the suspensory mechanisms seen in apes, and may be a precursor to obligate bipedalism seen in human ancestors.[
Hammerschmiede has also yielded the remains of several small creatures such as ]mollusc
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is esti ...
s, fish, and cold blooded vertebrates. Small mammals are the dominant fauna,[ such as ]rodent
Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...
s, rabbits
Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit specie ...
, marten
A marten is a weasel-like mammal in the genus ''Martes'' within the subfamily Guloninae, in the family Mustelidae. They have bushy tails and large paws with partially retractile claws. The fur varies from yellowish to dark brown, depending on t ...
s, hamsters, and shrew
Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to different fa ...
s. However, large mammals are also known, such as the extinct mouse-deer
Chevrotains, or mouse-deer, are small even-toed ungulates that make up the family Tragulidae, the only extant members of the infraorder Tragulina. The 10 extant species are placed in three genera, but several species also are known only f ...
''Dorcatherium
''Dorcatherium'' is an extinct genus of tragulid ruminant which existed in Europe, East Africa and the Siwaliks during the Miocene and Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5 ...
'',[ the ]antelope
The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia.
Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mammals ...
''Miotragocerus
''Miotragocerus'', also known as the European eland, is an extinct species of antelope that once lived in Europe in 10 to 8 million years ago. They were most likely browsers
Browse, browser or browsing may refer to:
Programs
* Web browser, a ...
'', and the rhinoceros
A rhinoceros (; ; ), commonly abbreviated to rhino, is a member of any of the five extant species (or numerous extinct species) of odd-toed ungulates in the family Rhinocerotidae. (It can also refer to a member of any of the extinct species o ...
''Aceratherium
''Aceratherium'' (Greek: "without (a) horn" (keratos), "beast" (therion)) was a genus of rhinoceros of the subfamily Aceratheriinae that lived in Eurasia during the Miocene.
Taxonomy
''Aceratherium'' was coined by Kaup (1832) for ''"Rhinoceros ...
''. The site is located in the Upper Freshwater Molasse
Upper may refer to:
* Shoe upper or ''vamp'', the part of a shoe on the top of the foot
* Stimulant, drugs which induce temporary improvements in either mental or physical function or both
* ''Upper'', the original film title for the 2013 found fo ...
of the Molasse basin
The Molasse basin (or North Alpine foreland basin) is a foreland basin north of the Alps which formed during the Oligocene and Miocene epochs. The basin formed as a result of the flexure of the European plate under the weight of the orogenic ...
; by the late Miocene, the Paratethys Sea
The Paratethys sea, Paratethys ocean, Paratethys realm or just Paratethys was a large shallow inland sea that stretched from the region north of the Alps over Central Europe to the Aral Sea in Central Asia.
Paratethys was peculiar due to its pal ...
had dried up and the Alps
The Alps () ; german: Alpen ; it, Alpi ; rm, Alps ; sl, Alpe . are the highest and most extensive mountain range system that lies entirely in Europe, stretching approximately across seven Alpine countries (from west to east): France, Sw ...
had lifted, allowing the expansion of wetland
A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (anoxic) processes prevailing, especially in the soils. The ...
habitats in the basin. The late Miocene may have been the beginning of a drying trend characterized by increased seasonality, causing deciduous
In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, aft ...
forest to turn into a less dense woodland
A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the ''plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (see ...
, and fruit and leaf production to occur cyclically rather than year-round. The late Miocene cooling trend may have led to the replacement of more tropical flora by mid-latitude and alpine
Alpine may refer to any mountainous region. It may also refer to:
Places Europe
* Alps, a European mountain range
** Alpine states, which overlap with the European range
Australia
* Alpine, New South Wales, a Northern Village
* Alpine National Pa ...
varieties, and ultimately the extinction of European great apes.[ ]
See also
References
{{Taxonbar, from=Q73803131
Prehistoric apes
Serravallian life
Tortonian life
Neogene Germany
Miocene primates
Fossils of Germany
Fossil taxa described in 2019