Morganucodon

{{Short description|Early mammaliaform genus of the Triassic and Jurassic periods}}

{{Automatic taxobox

| fossil_range = Late Triassic-Middle Jurassic {{Fossilrange|Rhaetian|Bathonian}}

| image = Restored skull of Morganucodon oehleri.jpg

| image_caption = Scan and reconstruction of the M. oehleri holotype skull

| image2 = Morganucodon watsoni life restoration.jpg

| image2_caption = Life restoration of Morganucodon watsoni

| taxon = Morganucodon

| authority = Kühne, 1949

| type_species = {{Extinct}}Morganucodon watsoni

| type_species_authority = Kühne, 1949

| subdivision_ranks = Species

| subdivision = * {{Extinct}}M. watsoni {{small|(Kühne, 1949)}}

{{Extinct}}M. oehleri {{small|(Rigney, 1963)}}
{{Extinct}}M. heikuopengensis {{small|(Young, 1978)}}
{{Extinct}}M. peyeri {{small|(Clemens, 1980)}}
{{Extinct}}M. tardus {{small|(Butler and Sigogneau-Russell, 2016)}}

}}

Morganucodon ("Glamorgan tooth") is an early mammaliaform genus that lived from the Late Triassic to the Middle Jurassic. 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 in Wales (Morganucodon watsoni), but fossils have also been found in Yunnan Province in China (Morganucodon oehleri) and various parts of Europe and North America. Some closely related animals (Megazostrodon) are known from exquisite fossils from South Africa.Pages 21–33, 174 in Zofia Kielan-Jaworowska, Richard L. Cifelli, and Zhe-Xi Luo, Mammals from the Age of Dinosaurs: Origins, Evolution, and Structure, Columbia University Press, New York, 2004 {{ISBN|0-231-11918-6}}

The name comes from a Latinization of Morganuc, the name for South Glamorgan in the Domesday Book, the county of Wales 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

File:Morganucodon watsoni.JPG

In the summer of 1947, fieldwork was done at Duchy Quarry in Glamorgan in southern Wales. Grey conglomerate that formed fissure fill deposits within karstic 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 in the Domesday Book, with the species name being in honour of D. M. S. Watson. Additional remains of M. watsoni were described by Kühne in 1958.{{Cite journal |last=Kühne |first=Walter Georg |author-link=Walter Georg Kühne |date=1958-08-01 |title=Rhaetische Triconodonten aus Glamorgan, ihre Stellung zwischen den Klassen Reptilia und Mammalia und ihre Bedeutung für die REICHART'sche Theorie |trans-title=Rhaetian triconodonts from Glamorgan, their position between the classes Reptilia and Mammalia and their significance for REICHART's theory. |url=https://doi.org/10.1007/BF02989032 |journal=Paläontologische Zeitschrift |language=de |volume=32 |issue=3 |pages=197–235 |doi=10.1007/BF02989032 |issn=0031-0220 |s2cid=128828761|url-access=subscription }} 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.{{Cite journal|last=Clemens|first=William A.|date=May 1979|title=A problem in morganucodontid taxonomy (Mammalia)|url=https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.1979.tb01898.x|journal=Zoological Journal of the Linnean Society|language=en|volume=66|issue=1|pages=1–14|doi=10.1111/j.1096-3642.1979.tb01898.x|url-access=subscription}} In August 1948, an expedition to Lufeng in Yunnan, China yielded a {{Convert|1|in|cm|-long|adj=mid}} 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 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.{{Cite journal|last=Rigney|first=Harold W.|date=March 1963|title=A Specimen of Morganucodon from Yunnan|url=http://www.nature.com/articles/1971122a0|journal=Nature|language=en|volume=197|issue=4872|pages=1122–1123|doi=10.1038/1971122a0|bibcode=1963Natur.197.1122R|s2cid=4204736|issn=0028-0836|url-access=subscription}} In 1978 C. C. Young described Eozostrodon 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.C.-C. Young. 1978. New materials of Eozostrodon. Vertebrata PalAsiatica 16:1-3 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.W. A. Clemens. 1980. [https://www.zobodat.at/pdf/Zitteliana_5_0051-0092.pdf Rhaeto-Liassic mammals from Switzerland and West Germany]. Zitteliana 5:51-92 In 1981, Kermack, Mussett and Rigney published an extensive monograph on the skull of Morganucodon.{{Cite journal|last1=Kermack|first1=K. A.|last2=Mussett|first2=Frances|last3=Rigney|first3=H. W.|date=January 1981|title=The skull of Morganucodon|url=https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.1981.tb01127.x|journal=Zoological Journal of the Linnean Society|language=en|volume=71|issue=1|pages=1–158|doi=10.1111/j.1096-3642.1981.tb01127.x|url-access=subscription}} 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, England, dating to the late Bathonian stage of the Middle Jurassic. The species being named after the Latin tardus, late, in reference to it being the youngest member of the genus.Butler, P.M. and Sigogneau-Russell, D. 2016. Diversity of triconodonts in the Middle Jurassic of Great Britain. Palaeontologia Polonica 67, 35–65. LSID urn:lsid:zoobank.org: pub: C4D90BB6-A001-4DDB-890E-2061B4793992

Biology

File:Morganucodon.jpg of M. oehleri]]

File:Morganucodon skull.svg

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 [4 inches]. In general appearance, it would have looked like a shrew or mouse".Kemp T.S. 2005. The origin and evolution of mammals, Oxford University Press, page 143. {{ISBN|0-19-850760-7}}. There is evidence that it had specialized glands used for grooming, which may indicate that, like present day mammals, it had fur.{{cite journal| last1=Ruben|first1= J.A. |last2= Jones| first2= T.D. |year=2000|title= Selective Factors Associated with the Origin of Fur and Feathers |journal=American Zoologist|volume= 40|pages= 585–596 | doi = 10.1093/icb/40.4.585| issue=4 |doi-access= free}}

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.{{cite journal|last1=Hall|first1=M. I.|last2=Kamilar|first2=J. M.|last3=Kirk|first3=E. C.|title=Eye shape and the nocturnal bottleneck of mammals|journal=Proceedings of the Royal Society B: Biological Sciences|date=24 October 2012|volume=279|issue=1749|pages=4962–4968|doi=10.1098/rspb.2012.2258|pmid=23097513|pmc=3497252}}{{cite journal|last1=Muchlinski|first1=Magdalena N.|title=A comparative analysis of vibrissa count and infraorbital foramen area in primates and other mammals|journal=Journal of Human Evolution|date=June 2010|volume=58|issue=6|pages=447–473|doi=10.1016/j.jhevol.2010.01.012|pmid=20434193}} Likewise, burrowing was widespread both in non-mammalian cynodonts and in primitive mammals.{{cite journal|last1=Damiani|first1=R.|last2=Modesto|first2=S.|last3=Yates|first3=A.|last4=Neveling|first4=J.|title=Earliest evidence of cynodont burrowing|journal=Proceedings of the Royal Society B: Biological Sciences|date=22 August 2003|volume=270|issue=1525|pages=1747–1751|doi=10.1098/rspb.2003.2427|pmid=12965004|pmc=1691433}}{{cite journal |last1=Kielan-Jaworowska |first1=Zofia |author-link=Zofia Kielan-Jaworowska |last2=Gambaryan |first2=Petr P. |name-list-style=and |date=December 1994 |title=Postcranial anatomy and habits of Asian multituberculate mammals |journal=Lethaia |volume=27 |issue=4 |pages=300 |doi=10.1111/j.1502-3931.1994.tb01578.x |s2cid=85021289}} The logics of phylogenetic bracketing would make Morganucodon nocturnal and burrowing too. Plant material from the conifer Hirmeriella 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.{{cite journal |last1=Gill |first1=Pamela G. |last2=Purnell |first2=Mark A. |last3=Crumpton |first3=Nick |author-link4=Kate Robson Brown|last4=Robson-Brown |first4=Kate |last5=Gostling |first5=Neil J. |last6=Stampanoni |first6=M. |last7=Rayfield |first7=Emily J. |date=21 August 2014 |title=Dietary specializations and diversity in feeding ecology of the earliest stem mammals |journal=Nature |volume=512 |issue=7514 |pages=303–305 |doi=10.1038/nature13622 |pmid=25143112 |bibcode=2014Natur.512..303G |hdl=2381/29192 |s2cid=4469841 |hdl-access=free }} Like most modern mammal insectivores, it grew fairly quickly to adult size.{{cite journal |last=Chinsamy |first=A. |last2=Hurum |first2=J.H. |year=2006 |title=Bone microstructure and growth patterns of early mammals |url=http://www.app.pan.pl/archive/published/app51/app51-325.pdf |journal=Acta Palaeontologica Polonica |volume=51 |issue=2 |pages=325–338 |access-date=30 August 2013}} Its eggs were probably small and leathery, a condition still found in monotremes.{{cite journal |last=Parente |first=Raphael Câmara Medeiros |last2=Bergqvist |first2=Lílian Paglarelli |last3=Soares |first3=Marina Bento |last4=Filho |first4=Olimpio Barbosa Moraes |year=2011 |title=The history of vaginal birth |journal=Archives of Gynecology and Obstetrics |volume=284 |issue=1 |pages=1–11 |doi=10.1007/s00404-011-1918-6 |pmid=21547459 |s2cid=22997887}}

The teeth grew in mammalian fashion, with deciduous teeth 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; indeed, it may have been among the first animals to do so.{{cite book |last1=Kielan-Jaworowska |first1=Zofia |author-link=Zofia Kielan-Jaworowska |title=Mammals from the age of dinosaurs : origins, evolution, and structure |last2=Cifelli |first2=Richard L. |last3=Luo |first3=Zhe-Xi |date=2004 |publisher=Columbia University Press |isbn=978-0231119184 |location=New York |pages=148–153}} 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.{{cite journal |last1=Crompton |first1=A. W. |last2=Jenkins |first2=Farish A. Jr. |author-link2=Farish Jenkins |year=1968 |title=Molar occlusion in late Triassic mammals |journal=Biological Reviews |volume=43 |issue=4 |pages=427–458 |doi=10.1111/j.1469-185x.1968.tb00966.x |pmid=4886687 |s2cid=1044399}}

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 endothermic metabolism seen in current mammals.{{cite journal |last1= Newham |first1=Elis |last2=Gill |first2=Pamela |display-authors=1 |date=2020 |title=Reptile-like physiology in Early Jurassic stem-mammals |journal=Nature Communications |volume=11 |issue=1 |pages=5121 |doi=10.1038/s41467-020-18898-4 |pmid=33046697 |pmc=7550344 |bibcode=2020NatCo..11.5121N |doi-access=free}}

Species

File:Morganucodon.svg

class="wikitable sortable mw-collapsible mw-uncollapsed"

!Species

! width="50" |Author

!Year

!Status

! width="200" |Temporal range

!Location

!Formations

Morganucodon watsoni

|Kuehne

|1949

|Early Jurassic (Hettangian–Sinemurian)

|England, Wales

|Various fissure fill deposits

Morganucodon heikuopengensis

|Young

|1978

|Early Jurassic (Sinemurian)

| rowspan="2" |Yunnan, China

| rowspan="2" |Lufeng

Morganucodon oehleri

|Rigney

|1963

|Early Jurassic (Hettangian)

Morganucodon peyeri

|Clemens

|1980

|Late Triassic (Rhaetian)

|Switzerland, France

|Klettgau, grès infraliasiques (Saint-Nicolas-de-Port)

Morganucodon tardus

|Butler and Sigogneau-Russel

|2016

|Middle Jurassic (Bathonian)

|England

|Forest Marble

Classification

File:Morganucodon watsoni life restoration.jpg

Morganucodon is the type genus for the order Morganucodonta, a group of generally similar mammaliaforms known from the Late Triassic to Late Jurassic epochs,{{cite journal |last1=Martin |first1=T. |last2=Averianov |first2=A. O. |last3=Jäger |first3=K. R. K. |last4=Schwermann |first4=A. H. |last5=Wings |first5=O. |title=A large morganucodontan mammaliaform from the Late Jurassic of Germany |journal=Fossil Imprint |date=2019 |volume=75 |issue=3–4 |pages=504–509 |doi=10.2478/if-2019-0030 |url=http://fi.nm.cz/wp-content/uploads/2019/12/13_Martin.pdf|doi-access=free }}pages 511–512, Malcolm C. McKenna and Susan K. Bell, Classification of Mammals Above the Species Level, Columbia University Press, 1997. {{ISBN|0-231-11012-X}} with one possible member (Purbeckodon) dating to the Early Cretaceous.{{cite journal |last=Butler |first=P. M. |last2=Sigogneau-Russell |first2=D. |last3=Ensom |first3=P. C. |year=2011 |title=Possible persistence of the morganucodontans in the Lower Cretaceous Purbeck Limestone Group (Dorset, England) |journal=Cretaceous Research |volume=33 |pages=135–145 |doi=10.1016/j.cretres.2011.09.007}} 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. 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.{{Cite journal | doi=10.1111/j.1096-3642.1981.tb01127.x|title = The skull of Morganucodon| journal=Zoological Journal of the Linnean Society| volume=71| pages=1–158|year = 1981|last1 = Kermack|first1 = K. A.| last2=Mussett| first2=Frances| last3=Rigney| first3=H. W.}} 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.[https://web.archive.org/web/20040925071034/http://www.geocities.com/trevor_dykes/mammalsbasal.htm Mammals of the Mesozoic: The least mammal-like mammals]

;Phylogeny{{cite journal|last1=Close|first1=Roger A.|last2=Friedman|first2=Matt|last3=Lloyd|first3=Graeme T.|last4=Benson|first4=Roger BJ|year=2015|title=Evidence for a mid-Jurassic adaptive radiation in mammals.|journal=Current Biology|volume=25|issue=16|pages=2137–2142|doi=10.1016/j.cub.2015.06.047|pmid=26190074|doi-access=free}}

{{clade |style=font-size:95%;line-height:80%;

|label1=Mammaliaformes

|1={{clade

|1=Adelobasileus

|2={{clade

|1=Sinoconodon

|2={{clade

|1={{clade

|1=Morganucodon

|2=Megazostrodon

}}

|2={{clade