Megaladapis
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''Megaladapis'', informally known as the koala lemur, 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 of
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 belonging to the family Megaladapidae, consisting of three species that once inhabited the island of
Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...
. The largest measured between in length.


Description

''Megaladapis'' was quite different from any living lemur. Its body was squat and built like that of the modern
koala The koala (''Phascolarctos cinereus''), sometimes inaccurately called the koala bear, is an arboreal herbivorous marsupial native to Australia. It is the only Extant taxon, extant representative of the Family (biology), family ''Phascolar ...
. Its long arms, fingers, feet, and toes were specialized for grasping trees, and its legs were splayed for vertical climbing. The
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
of its foot suggests ''Megaladapis'' evolved to live in an
arboreal Arboreal locomotion is the 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 (scansorial), but others are exclusively arboreal. The hab ...
environment, having a large
hallux Toes are the digits 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 ''plantigrade''; ...
and lateral abductor musculature that helped it to grasp vertically on trees. These features are shared by other arboreal species. The hands and feet were curved and the ankles and wrists did not have the usual stability needed to travel on the ground that most other lemurids have. Its body weight reached . Other estimates suggest but its still much larger than any extant lemur. It had the largest body size of any lemur, with double the body mass of the next largest extinct lemur.


Cranial anatomy

The shape of its
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
was unique among all known primates, with a
nasal region A nose is a sensory organ and respiratory structure in vertebrates. It consists of a nasal cavity inside the head, and an external nose on the face. The external nose houses the nostrils, or nares, a pair of tubes providing airflow through the ...
which showed similarities to those of
rhinoceros A rhinoceros ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...
, a feature that probably combined with an enlarged upper lip for grasping leaves. Its head was unlike that of any other primate; most strikingly, its eyes were on the sides of its skull, instead of forward on the skull as in all other primates. An
endocast An endocast is the internal cast of a hollow object, often referring to the cranial vault in the study of brain development in humans and other organisms. Endocasts can be artificially made for examining the properties of a hollow, inaccessible ...
of its skull showed that it had a brain capacity of about , about 3 to 4 times the size of a
domestic cat The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small Domestication, domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have sh ...
's, which is small for its size when compared to other lemurs. Compared to the size of the skull, the diameter of the
orbit In celestial mechanics, an orbit (also known as orbital revolution) 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 ...
s protrudes outwards and forwards in a tabular form, suggesting that ''Megaladapis'' was diurnal. Details about the
anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
parts of the
dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiology ...
, such as the canines and incisors, are difficult to determine. The bulle osseve are broken away. The foremost facial portion and base of the skull is also wanting. The total length of the skull of ''M. madagascariensis'' has been calculated to be about , about from three to four times that of a domestic cat. Based on the wear on the teeth, the obliteration of most of the sutures of the very thick bones, and the strongly developed crests, it is believed to have been an elderly individual. There are several well-preserved fragments of the upper and
lower jaw In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
. Its long canine teeth and cow-like jaw formed a tapering
snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, Rostrum (anatomy), rostrum, beak or proboscis. The wet furless surface around the nostrils of the n ...
. Its
jaw muscles The four classical muscles of mastication elevate the mandible (closing the jaw) and move it forward/backward and laterally, facilitating biting and chewing. Other muscles are responsible for opening the jaw, namely the geniohyoid, mylohyoid, and ...
were powerful for chewing the tough native vegetation. Based on the patterns of
dental microwear Dental microwear analysis is a method to infer diet and behavior in extinct animals, especially in fossil specimens. It has been used on a variety of taxa, including hominids, victoriapithecids, amphicyonids, canids, ursids, hyaenids, hyaenodont ...
, ''Megaladapis'' is believed to have been
folivorous In zoology, a folivore is a herbivore that specializes in eating leaves. Mature leaves contain a high proportion of hard-to-digest cellulose, less energy than other types of foods, and often toxic compounds.Jones, S., Martin, R., & Pilbeam, D. (1 ...
, using a leaf-cropping
foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavi ...
method. These patterns found no permanent upper incisors or the presence of an expanded
articular facet A joint or articulation (or articular surface) is the connection made between bones, ossicles, or other hard structures in the body which link an animal's skeletal system into a functional whole.Saladin, Ken. Anatomy & Physiology. 7th ed. McGraw- ...
on the posterior face of the
mandibular condyle The condyloid process or condylar process is the process on the human and other mammalian species' mandibles that ends in a condyle, the mandibular condyle. It is thicker than the coronoid process of the mandible and consists of two portions: the ...
. This diet and similar
phenotypic trait A phenotypic trait, simply trait, or character state is a distinct variant of a phenotypic characteristic of an organism; it may be either inherited or determined environmentally, but typically occurs as a combination of the two.Lawrence, Eleano ...
s of the teeth are the basis for concluding a shared ancestry with the ''
Lepilemur 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. ...
''. The upper molars of ''Lepilemur'' are very close in shape to those of ''Megaladapis''. The main difference between the two is that the outer crown-surface of ''Lepilemur'' molars forms a nearly straight line, almost parallel with the long axis of the skull, and the outer side is slightly concave inwards. The antero-internal cingulum is missing in the molars of ''Lepilemur''.


Biology

The island's
topography Topography is the study of the forms and features of land surfaces. The topography of an area may refer to the landforms and features themselves, or a description or depiction in maps. Topography is a field of geoscience and planetary sci ...
was always changing, and like other lemurs, ''Megaladapis'' was specialized within its own
niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development and growth *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ec ...
. The general expectations of tree climbers such as ''Megaladapis'' is that with an increase in size, the body's forelimbs will also increase proportionally. The diet of ''Megaladapis'' might be the factor that influences the dental development; species with a larger brain, later initiation of molar crowns, and longer formation of crown are considered to have more of an omnivorous diet. In contrast, ''Megaladapis'' lived on a folivorous diet, despite having a smaller brain, early initiation of molar crowns, and fast crown formation. Based on when molar crown initiation occurred, ''Megaladapis''
gestation period In mammals, pregnancy is the period of reproduction during which a female carries one or more live offspring from implantation in the uterus through gestation. It begins when a fertilized zygote implants in the female's uterus, and ends once i ...
is thought to have been at least 198 days, but was likely longer. ''Megaladapis'' were diurnal, or active during the day. Lemurs in general had small group sizes and were highly
seasonal breeder Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and ch ...
s (they breed for about one to two weeks a year), which is assumed to be the case for ''Megaladapis''.Scott, Rob. "The Lost Lemurs: Extinction in Madagascar." Rutgers University. Hickman Hall, New Brunswick, NJ. ''n.d.'' Lecture.


Extinction

When humans arrived on Madagascar 2,300 years ago, there were at least 17 species of now-extinct "giant" lemur, in addition to the species alive today. Their seasonal breeding and long gestation times, resulting in low breeding rates, coupled with their diurnality and presumed slow movement were disadvantageous in this new ecological context; ''Megaladapis'' (along with the other species of giant lemur) were more susceptible to new predators (humans), forest fires, and
habitat destruction Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...
due to these traits. Over-hunting by humans was deemed a major contributor to the extinction of "giant" lemurs. Minor
drought A drought is a period of drier-than-normal conditions.Douville, H., K. Raghavan, J. Renwick, R.P. Allan, P.A. Arias, M. Barlow, R. Cerezo-Mota, A. Cherchi, T.Y. Gan, J. Gergis, D.  Jiang, A.  Khan, W.  Pokam Mba, D.  Rosenfeld, J. Tierney, ...
s are frequent in Madagascar, but a major drought approximately 1000 years ago significantly lowered water levels in lakes, caused a severe vegetation transition, and created conditions suitable for
wildfire A wildfire, forest fire, or a bushfire is an unplanned and uncontrolled fire in an area of Combustibility and flammability, combustible vegetation. Depending on the type of vegetation present, a wildfire may be more specifically identified as a ...
s in grasslands and
savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
s.
Crop failure Harvesting is the process of collecting plants, animals, or fish (as well as fungi) as food, especially the process of gathering mature crops, and "the harvest" also refers to the collected crops. Reaping is the cutting of grain or pulses fo ...
s due to these conditions would drive inhabitants to hunt for
bushmeat Bushmeat is meat from wildlife species that are Hunting, hunted for human consumption. Bushmeat represents a primary source of animal protein and a cash-earning commodity in poor and rural communities of humid tropical forest regions of the worl ...
to survive, and these giant lemurs were an easy source of said meat. Some exterior scratches and incisions were found on both the
metatarsus The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
and
mandibula In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...
of ''Megaladapis'' fossils. The cuts on the metatarsus are comparable to those found in caves and are thought to have been produced by humans, while those on the mandibula seem to have been produced by some instrument engineered for cutting – indications that the ''Megaladapis'' was at some point in direct contact with the anatomically-modern humans of its time. The
landscape A landscape is the visible features of an area of land, its landforms, and how they integrate with natural or human-made features, often considered in terms of their aesthetic appeal.''New Oxford American Dictionary''. A landscape includes th ...
in which giant lemurs were found were largely
forest A forest is an ecosystem characterized by a dense ecological community, community of trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, ...
ed areas with dense vegetation. Almost directly after human arrival, there was a rapid decline in the spores of the
coprophilous fungus A coprophilous fungus (''dung-loving'' fungus) is a type of saprobic fungus that grows on animal dung. The hardy spores of coprophilous species are unwittingly consumed by herbivores from vegetation, and are excreted along with the plant matter. ...
''
Sporormiella ''Sporormiella'' is a genus of fungi in the phylum Ascomycota whose species can be found worldwide, including the Arctic. It grows primarily on dung but also can be found in soil and plant debris. The exact number of species is debated and can ra ...
'', which indicates a decrease in megafaunal
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
. Charcoal
microparticle Microparticles are particles between 0.1 and 100 μm in size. Commercially available microparticles are available in a wide variety of materials, including ceramics, glass, polymers, and metals. Microparticles encountered in daily life incl ...
s being found in surveys of various areas in Madagascar give evidence to the fact that human habitat modification only occurred after this decline in megafaunal biomass.
Charcoal Charcoal is a lightweight black carbon residue produced by strongly heating wood (or other animal and plant materials) in minimal oxygen to remove all water and volatile constituents. In the traditional version of this pyrolysis process, ca ...
deposits provide evidence to the fact that humans used fire to clear large pieces of land very rapidly. The habitats that ''Megaladapis'' were well-adapted to live in were converted into
grassland A grassland is an area where the vegetation is dominance (ecology), dominated by grasses (Poaceae). However, sedge (Cyperaceae) and rush (Juncaceae) can also be found along with variable proportions of legumes such as clover, and other Herbaceo ...
s, which provided little to no cover from outside forces for these creatures. Thus, the final decline of "giant" lemur populations, including ''Megaladapis'', were hypothesized to be due to
habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological proces ...
through human activities (for example, clearing of land through "
slash-and-burn Slash-and-burn agriculture is a form of shifting cultivation that involves the cutting and burning of plants in a forest or woodland to create a Field (agriculture), field called a swidden. The method begins by cutting down the trees and woody p ...
" techniques), resulting in the extinction of these lemurs between 500 and 600 years ago.


Cultural references

It is often believed that Malagasy legends of the '' tretretretre'' or ''tratratratra'', an extinct animal, refer to ''Megaladapis'', but the details of these tales, notably the "human-like" face of the animal, match the related '' Palaeopropithecus'' much better.


Gallery


References


External links


''Megaladapis edwardsi'': Scientific information


{{Taxonbar, from=Q137557 Subfossil lemurs Lemur genera Prehistoric primate genera Prehistoric animals of Madagascar Holocene extinctions Fossil taxa described in 1894 Species made extinct by human activities Taxa named by Charles Immanuel Forsyth Major