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''Morganucodon'' ("
Glamorgan Glamorgan (), or sometimes Glamorganshire ( or ), was Historic counties of Wales, one of the thirteen counties of Wales that existed from 1536 until their abolishment in 1974. It is located in the South Wales, south of Wales. Originally an ea ...
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 of synapsid tetrapods that includes the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts during the Late Triassic. It is defined a ...
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 ...
that lived from the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
to the
Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 161.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relativel ...
. 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 Glamorgan (), or sometimes Glamorganshire ( or ), was Historic counties of Wales, one of the thirteen counties of Wales that existed from 1536 until their abolishment in 1974. It is located in the South Wales, south of Wales. Originally an ea ...
in
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
(''Morganucodon watsoni''), but fossils have also been found in
Yunnan Yunnan; is an inland Provinces of China, province in Southwestern China. The province spans approximately and has a population of 47.2 million (as of 2020). The capital of the province is Kunming. The province borders the Chinese provinces ...
Province in
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
(''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. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
. The name comes from a Latinization of ''Morganuc'', the name for South
Glamorgan Glamorgan (), or sometimes Glamorganshire ( or ), was Historic counties of Wales, one of the thirteen counties of Wales that existed from 1536 until their abolishment in 1974. It is located in the South Wales, south of Wales. Originally an ea ...
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 at the behest of William the Conqueror. The manuscript was originally known by ...
, the county of
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
where it was discovered by Walter Georg Kühne,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 Glamorgan (), or sometimes Glamorganshire ( or ), was Historic counties of Wales, one of the thirteen counties of Wales that existed from 1536 until their abolishment in 1974. It is located in the South Wales, south of Wales. Originally an ea ...
in southern Wales. Grey conglomerate that formed fissure fill deposits within
karst Karst () is a topography formed from the dissolution of soluble carbonate rocks such as limestone and Dolomite (rock), dolomite. It is characterized by features like poljes above and drainage systems with sinkholes and caves underground. Ther ...
ic voids in Carboniferous limestone was extracted. In 1949, Walter Georg Kühne 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 Glamorgan (), or sometimes Glamorganshire ( or ), was Historic counties of Wales, one of the thirteen counties of Wales that existed from 1536 until their abolishment in 1974. It is located in the South Wales, south of Wales. Originally an ea ...
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 at the behest of William the Conqueror. The manuscript was originally known by ...
, 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 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 an inland Provinces of China, province in Southwestern China. The province spans approximately and has a population of 47.2 million (as of 2020). The capital of the province is Kunming. The province borders the Chinese provinces ...
, China yielded a 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 University College London (Trade name, branded as UCL) is a Public university, public research university in London, England. It is a Member institutions of the University of London, member institution of the Federal university, federal Uni ...
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 teeth from South West England South West England, or the South West of England, ...
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) deposits near Hallau, Switzerland, with the species being named after paleontologist Bernhard Peyer. In 1981, Kermack, Mussett and Rigney published an extensive monograph on the skull of ''Morganucodon''. In 2016 Percy Butler and Denise Sigogneau-Russell named the species ''Morganucodon tardus'' from an upper right molar (M34984) collected from the Watton Cliff locality near Eype in
Dorset Dorset ( ; Archaism, archaically: Dorsetshire , ) is a Ceremonial counties of England, ceremonial county in South West England. It is bordered by Somerset to the north-west, Wiltshire to the north and the north-east, Hampshire to the east, t ...
, England, dating to the late
Bathonian In the geologic timescale the Bathonian is an age (geology), age and stage (stratigraphy), stage of the Middle Jurassic. It lasted from approximately 168.2 ±1.2 annum, Ma to around 165.3 ±1.1 Ma (million years ago). The Bathonian Age succeeds ...
stage of the
Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 161.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relativel ...
. 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 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 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 Cynodontia () is a clade of eutheriodont therapsids that first appeared in the Late Permian (approximately 260 Megaannum, mya), and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extin ...
s and in primitive mammals. The logics 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,Fehrenbach, MJ and Popowics, T. (2026). ''Illustrated Dental Embryology, Histology, and Anatomy'', 6th edition, Elsevier, page 287–296. are ...
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 process ...
; 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 inst ...
ic metabolism seen in current mammals.


Species


Classification

''Morganucodon'' is the
type genus In biological taxonomy, the type genus (''genus typica'') 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-bearin ...
for the order Morganucodonta, a group of generally similar mammaliaforms known from the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
to
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
epochs, with one possible member ('' Purbeckodon'') dating to the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
. 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 of synapsid tetrapods that includes the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts during the Late Triassic. It is defined a ...
. Some researchers limit the term "mammal" to the crown group 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 and quadrate bones 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 (geology), Pennsylvanian sub-period of the late Carboniferous period. By the mid-Triassic, there were many synaps ...


References

{{Taxonbar, from=Q131813 Morganucodonta 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