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''Morganucodon'' ("
Glamorgan , HQ = Cardiff , Government = Glamorgan County Council (1889–1974) , Origin= , Code = GLA , CodeName = Chapman code , Replace = * West Glamorgan * Mid Glamorgan * South Glamorgan , Motto ...
tooth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tea ...
") is an early
mammaliaform Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recent comm ...
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 nomencla ...
that lived from the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch. Th ...
to the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations c ...
. It first appeared about 205 million years ago. Unlike many other early mammaliaforms, ''Morganucodon'' is well represented by abundant and well preserved (though in the vast majority of cases disarticulated) material. Most of this comes from
Glamorgan , HQ = Cardiff , Government = Glamorgan County Council (1889–1974) , Origin= , Code = GLA , CodeName = Chapman code , Replace = * West Glamorgan * Mid Glamorgan * South Glamorgan , Motto ...
in
Wales Wales ( cy, Cymru ) is a country that is part of the United Kingdom. It is bordered by England to the east, the Irish Sea to the north and west, the Celtic Sea to the south west and the Bristol Channel to the south. It had a population in 2 ...
(''Morganucodon watsoni''), but fossils have also been found in
Yunnan Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the ...
Province in China (''Morganucodon oehleri'') and various parts of Europe and North America. Some closely related animals (''
Megazostrodon ''Megazostrodon'' is an extinct genus of basal mammaliaforms belonging to the order Morganucodonta. It is approximately 200 million years old.
'') are known from exquisite fossils from
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. It is bounded to the south by of coastline that stretch along the Atlantic Ocean, South Atlantic and Indian Oceans; to the ...
. The name comes from a Latinization of ''Morganuc'', the name for South
Glamorgan , HQ = Cardiff , Government = Glamorgan County Council (1889–1974) , Origin= , Code = GLA , CodeName = Chapman code , Replace = * West Glamorgan * Mid Glamorgan * South Glamorgan , Motto ...
in the
Domesday Book Domesday Book () – the Middle English spelling of "Doomsday Book" – is a manuscript record of the "Great Survey" of much of England and parts of Wales completed in 1086 by order of King William I, known as William the Conqueror. The manusc ...
, the county of
Wales Wales ( cy, Cymru ) is a country that is part of the United Kingdom. It is bordered by England to the east, the Irish Sea to the north and west, the Celtic Sea to the south west and the Bristol Channel to the south. It had a population in 2 ...
where it was discovered by
Walter Georg Kühne Walter Georg Kühne (February 26, 1911 in Berlin – March 16, 1991 ibid.) was a German paleontologist, known as a "legendary explorer of Mesozoic mammals". He graduated in March 1930 from reform boarding school '' Schule am Meer'' on the island ...
,Walter G. Kühne, "On a Triconodont tooth of a new pattern from a Fissure-filling in South Glamorgan", ''Proceedings of the Zoological Society of London'', volume 119 (1949–1950) pages 345–350 giving the meaning "Glamorgan tooth".


History of discovery

In the summer of 1947, fieldwork was done at Duchy Quarry in
Glamorgan , HQ = Cardiff , Government = Glamorgan County Council (1889–1974) , Origin= , Code = GLA , CodeName = Chapman code , Replace = * West Glamorgan * Mid Glamorgan * South Glamorgan , Motto ...
in southern Wales. Grey conglomerate that formed fissure fill deposits within
karst Karst is a topography formed from the dissolution of soluble rocks such as limestone, dolomite, and gypsum. It is characterized by underground drainage systems with sinkholes and caves. It has also been documented for more weathering-resistant ...
ic voids in
Carboniferous limestone Carboniferous Limestone is a collective term for the succession of limestones occurring widely throughout Great Britain and Ireland that were deposited during the Dinantian Epoch of the Carboniferous Period. These rocks formed between 363 and 3 ...
was extracted. In 1949,
Walter Georg Kühne Walter Georg Kühne (February 26, 1911 in Berlin – March 16, 1991 ibid.) was a German paleontologist, known as a "legendary explorer of Mesozoic mammals". He graduated in March 1930 from reform boarding school '' Schule am Meer'' on the island ...
noted the lower cheek tooth of a primitive mammal while examining samples of the rock. He named it ''Morganucodon watsoni,'' with the genus name being derived from Morganuc, which Kühne stated was the name of South
Glamorgan , HQ = Cardiff , Government = Glamorgan County Council (1889–1974) , Origin= , Code = GLA , CodeName = Chapman code , Replace = * West Glamorgan * Mid Glamorgan * South Glamorgan , Motto ...
in the
Domesday Book Domesday Book () – the Middle English spelling of "Doomsday Book" – is a manuscript record of the "Great Survey" of much of England and parts of Wales completed in 1086 by order of King William I, known as William the Conqueror. The manusc ...
, with the species name being in honour of D. M. S. Watson. Additional remains of ''M. watsoni'' were described by Kühne in 1958. Also in 1958,
Kenneth Kermack Kenneth A. Kermack (1919 – 2000) was a British palaeontologist at University College London most notable for his work on early mammals with his wife, Doris Mary Kermack. Among Kermack's other significant contributions was the observation ...
and Frances Mussett described additional remains from Pant Quarry, about a mile from Duchy Quarry, that had been collected in 1956. In August 1948, an expedition to Lufeng in
Yunnan Yunnan , () is a landlocked province in the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is Kunming. The province borders the ...
, China yielded a 1 in (2.5 cm) long skull. It was shortly sent to Beijing (then Peking) and then eventually sent out of China, and deposited with Kenneth Kermack at
University College London , mottoeng = Let all come who by merit deserve the most reward , established = , type = Public research university , endowment = £143 million (2020) , budget = � ...
in 1960. The specimen was preliminarily described in 1963 by Harold W. Rigney, who noted the similarity to ''Morganucodon'' from Britain, and considered it cogeneric, naming the new species ''Morganucodon oehleri'' in honor of the reverend Edgar T. Oehler, who had originally collected the specimen. In 1978 C. C. Young described ''
Eozostrodon ''Eozostrodon'' is an extinct morganucodont mammaliaform. It lived during the Rhaetian stage of the Late Triassic. ''Eozostrodon'' is known from disarticulated bones from Wales and South West England and estimated to have been less than in head- ...
heikuopengensis'' from the Hei Koa Peng locality near Lufeng, based on an associated skull and dentary, as well as a right maxilla and associated dentary. A revision by William A. Clemens in 1979 assigned this species to ''Morganucodon,'' based on its close similarity to the two previously named species.'''' In 1980 Clemens named the species ''Morganucodon peyeri'', from isolated teeth found in Late Triassic (
Rhaetian The Rhaetian is the latest age of the Triassic Period (in geochronology) or the uppermost stage of the Triassic System (in chronostratigraphy). It was preceded by the Norian and succeeded by the Hettangian (the lowermost stage or earliest age of ...
) deposits near
Hallau Hallau is a municipality in the canton of Schaffhausen in Switzerland. History Though Bronze Age weapons have been found in Hallau, the first traces of a settlement date from the Roman era. A Roman warehouse was found in Hüttenhau as well as ...
, Switzerland, with the species being named after paleontologist
Bernhard Peyer Bernhard Peyer (25 July 1885 – 23 February 1963) was a Swiss paleontologist and anatomist who served as a professor at the University of Zurich. A major contribution was on the evolution of vertebrate teeth. Peyer was born in Schaffhausen, Switze ...
. In 1981, Kermack, Mussett and Rigney published an extensive monograph on the skull of ''Morganucodon''. In 2016 Percy Butler and
Denise Sigogneau-Russell Denise Sigogneau-Russell (born ''c.'' 1941/42) is a French palaeontologist who specialises in mammals from the Mesozoic, particularly from France and the UK. She is currently based at the Muséum National d'Histoire Naturelle. Background Denise S ...
named the species ''Morganucodon tardus'' from an upper right molar (M34984) collected from the Watton Cliff locality near
Eype Eype ( ) is a small village in southwest Dorset, England, situated southwest of Bridport. It lies on the Jurassic Coast World Heritage Site on the English Channel and is within the civil parish of Symondsbury. Eype means "steep place". Many of ...
in
Dorset Dorset ( ; archaically: Dorsetshire , ) is a county in South West England on the English Channel coast. The ceremonial county comprises the unitary authority areas of Bournemouth, Christchurch and Poole and Dorset. Covering an area of , Do ...
, England, dating to the late
Bathonian In the geologic timescale the Bathonian is an age and stage of the Middle Jurassic. It lasted from approximately 168.3 Ma to around 166.1 Ma (million years ago). The Bathonian Age succeeds the Bajocian Age and precedes the Callovian Age. Stra ...
stage of the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations c ...
. The species being named after the Latin ''tardus'', late, in reference to it being the youngest member of the genus.


Biology

''Morganucodon'' was a small,
plantigrade 151px, Portion of a human skeleton, showing plantigrade habit In terrestrial animals, plantigrade locomotion means walking with the toes and metatarsals flat on the ground. It is one of three forms of locomotion adopted by terrestrial mammals. T ...
animal. The tail was moderately long. According to Kemp (2005), "the skull was 2–3 cm in length and a presacral body length of about 10 cm inches In general appearance, it would have looked like a shrew or mouse". There is evidence that it had specialized glands used for grooming, which may indicate that, like present day mammals, it had fur. Like present day mammals of similar size and presumed habit, ''Morganucodon'' was likely nocturnal and spent the day in a burrow. There is no direct fossil evidence, but several lines of evidence point to a
nocturnal bottleneck The nocturnal bottleneck hypothesis is a hypothesis to explain several mammalian traits. In 1942, Gordon Lynn Walls described this concept which states that placental mammals were mainly or even exclusively nocturnal through most of their evolu ...
in the evolution of the mammal class, and almost all modern mammals of similar size to ''Morganucodon'' are still nocturnal. Likewise, burrowing was widespread both in non-mammalian
cynodont The cynodonts () ( clade Cynodontia) are a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 mya), and extensively diversified after the Permian–Triassic extinction event. Cynodonts had a wide variet ...
s and in primitive mammals. The logics of
phylogenetic bracketing Phylogenetic bracketing is a method of inference used in biological sciences. It is used to infer the likelihood of unknown traits in organisms based on their position in a phylogenetic tree. One of the main applications of phylogenetic bracketing ...
would make ''Morganucodon'' nocturnal and burrowing too. Plant material from the conifer ''
Hirmeriella ''Hirmeriella'' is a genus of fossil tree, a conifer that was widespread in Late Triassic and Early Jurassic of Germany, the UK, and Poland.BARBACKA M., ZIAJA J., WCISŁO-LURANIEC E. 200Hirmeriella muensteri (Schenk) Jung from Odrowąż (Poland) ...
'' was also found in the fissure fills, indicating ''Morganucudon'' lived in, or near, a forested area. The diet appears to have been insects and other small animals, with a preference for hard prey such as beetles. Like most modern mammal insectivores, it grew fairly quickly to adult size. Its eggs were probably small and leathery, a condition still found in monotremes. The teeth grew in mammalian fashion, with
deciduous teeth Deciduous teeth or primary teeth, also informally known as baby teeth, milk teeth, or temporary teeth,Illustrated Dental Embryology, Histology, and Anatomy, Bath-Balogh and Fehrenbach, Elsevier, 2011, page 255 are the first set of teeth in the ...
being replaced by permanent teeth that were retained throughout the rest of the animal's life.Alexander F. H. van Nievelt and Kathleen K. Smith, "To replace or not to replace: the significance of reduced functional tooth replacement in marsupial and placental mammals", ''Paleobiology'', Volume 31, Issue 2 (June 2005) pages 324–346 The combination of rapid growth in juveniles and a toothless stage at infancy strongly suggests that ''Morganucodon'' raised its young by
lactation Lactation describes the secretion of milk from the mammary glands and the period of time that a mother lactates to feed her young. The process naturally occurs with all sexually mature female mammals, although it may predate mammals. The proce ...
; indeed, it may have been among the first animals to do so. The molars in the adult had a series of raised humps and edges that fit into each other, allowing for efficient chewing. However, unlike the situation in most later mammals, the upper and lower molars did not occlude properly when they first met; as they wore against each other, however, their shapes were modified by wear to produce a precise fit. A 2020 study suggests that the metabolism of ''Morganucodon'' was significantly slower than that of comparably sized modern mammals, and that it had a life-span more similar to that of reptiles, with the oldest specimen having a lifespan of 14 years. Thus it likely did not possess the fully
endotherm An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inste ...
ic metabolism seen in current mammals.


Species


Classification

''Morganucodon'' is the
type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal ...
for the order
Morganucodonta Morganucodonta ("Glamorgan teeth") is an extinct order of basal Mammaliaformes, a group including crown-group mammals (Mammalia) and their close relatives. Their remains have been found in Southern Africa, Western Europe, North America, India a ...
, a group of generally similar mammaliaforms known from the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch. Th ...
to
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
epochs, with one possible member (''
Purbeckodon ''Purbeckodon'' is an extinct genus of mammaliaforms, possibly belonging to Morganucodonta, that is known from Early Cretaceous deposits of southeastern Dorset, England. It was collected in the Purbeck Limestone Group of Dorset. It was first n ...
'') dating to the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous ( chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology P ...
. All were small and likely insectivorous. ''Morganucodon'' is the best preserved and best understood member of Morganucodonta. There is currently controversy about whether or not to classify ''Morganucodon'' as a mammal or as a non-mammalian
mammaliaform Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. It is defined as the clade originating from the most recent comm ...
. Some researchers limit the term "mammal" to the
crown 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. ...
mammals, which would not include ''Morganucodon'' and its relatives. Others, however, define "mammals", as a group, by the possession of a special, secondarily evolved jaw joint between the dentary and the squamosal bones, which has replaced the primitive one between the
articular The articular bone is part of the lower jaw of most vertebrates, including most jawed fish, amphibians, birds and various kinds of reptiles, as well as ancestral mammals. Anatomy In most vertebrates, the articular bone is connected to two oth ...
and
quadrate bone The quadrate bone is a skull bone in most tetrapods, including amphibians, sauropsids (reptiles, birds), and early synapsids. In most tetrapods, the quadrate bone connects to the quadratojugal and squamosal bones in the skull, and forms upper ...
s in all modern mammalian groups. Under this definition, ''Morganucodon'' would be a mammal. Nevertheless, its lower jaw retains some of the bones found in its non-mammalian ancestors in a very reduced form rather than being composed solely of the dentary. Furthermore, the primitive reptile-like jaw joint between the articular and quadrate bones, which in modern mammals has moved into the middle ear and become part of the ear ossicles as malleus and incus, is still to be found in ''Morganucodon''. ''Morganucodon'' also suckled (it may have been the earliest animal to do so), had only two sets of teeth and grew rapidly to adult size and stopped growing thereafter, all typical mammalian traits.Mammals of the Mesozoic: The least mammal-like mammals
/ref> ;Phylogeny


See also

*
Evolution of mammals The evolution of mammals has passed through many stages since the first appearance of their synapsid ancestors in the Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synapsid species that looked l ...


References

{{Taxonbar, from=Q131813 Morganucodonts Prehistoric cynodont genera Late Triassic synapsids Triassic synapsids of Europe Jurassic synapsids Transitional fossils Rhaetian first appearances Middle Jurassic extinctions Fossil taxa described in 1949 Taxa named by Walter Georg Kühne