Mammaliaformes

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Mammaliaforms
Temporal range: Late Triassic (Earliest Norian) – Present, 227–0  Ma
Clade Mammaliaformes.png
Top: Megazostrodon rudnerae , Morganucodon watsoni (Morganucodonta);

Middle: Castorocauda lutrasimilis (Docodonta), Shenshou lui (Haramiyida);
Bottom: Tachyglossus aculeatus (Monotremata), Ursus arctos (Theria).

Contents

Morganucodon skull.svg
Skull of Morganucodon
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Clade: Therapsida
Clade: Cynodontia
Clade: Prozostrodontia
Clade: Mammaliamorpha
Clade: Mammaliaformes
Rowe, 1988
Subgroups

Mammaliaformes ("mammalian forms") is a clade that contains the crown group mammals and their closest extinct relatives; the group radiated from earlier probainognathian cynodonts. [1] It is defined as the clade originating from the most recent common ancestor of Morganucodonta and the crown group mammals; the latter is the clade originating with the most recent common ancestor of extant Monotremata, Marsupialia, and Placentalia. [2] Besides Morganucodonta and the crown group mammals, Mammaliaformes includes Docodonta and Hadrocodium as well as the Triassic Tikitherium , the earliest known member of the group. [3] [4]

Mammaliaformes is a term of phylogenetic nomenclature. In contrast, the assignment of organisms to Mammalia has traditionally been founded on traits and, on this basis, Mammalia is slightly more inclusive than Mammaliaformes. In particular, trait-based taxonomy generally includes Adelobasileus and Sinoconodon in Mammalia, though they fall outside the Mammaliaformes definition. These genera are included in the broader clade Mammaliamorpha, defined phylogenetically as the clade originating with the last common ancestor of Tritylodontidae and the crown group mammals. [2] This wider group includes some families that trait-based taxonomy does not include in Mammalia, in particular Tritylodontidae and Brasilodontidae.

Animals in the Mammaliaformes clade are often called mammaliaforms, without the e. Sometimes, the spelling mammaliforms is used. The origin of crown-group mammals extends back to the Jurassic, with extensive findings in the Late Jurassic outcrops of Portugal and China. The earliest confirmed specimens of fur are found in them, demonstrating that the ancestors of mammals had already developed fur.

Mammaliaformes in life

Early mammaliaforms were generally shrew-like in appearance and size, and most of their distinguishing characteristics were internal. In particular, the structure of the mammaliaform (and mammal) jaw and the arrangement of teeth are nearly unique. Instead of having many teeth that are frequently replaced, mammals have one set of baby teeth and later one set of adult teeth that fit together precisely. This is thought to aid in the grinding of food to make it quicker to digest. [5] Endothermic animals require more calories than those that are ectothermic, so speeding up the pace of digestion is a necessity. The drawback to the fixed dentition is that worn teeth cannot be replaced, as was possible for the reptiliomorph ancestors of mammaliaforms. To compensate, mammals developed prismatic enamel, characterized by crystallite discontinuities that helped spread out the force of the bite. [6]

Lactation, along with other characteristically mammalian features, is also thought to characterize the Mammaliaformes, but these traits are difficult to study in the fossil record. Evidence of lactation is present in morganucodontans, via tooth replacement patterns. [7] Combined with the more basal tritylodontids that also display evidence of lactation, [8] this seems to imply that milk is an ancestral characteristic in this group. However, the fairly derived Sinoconodon appears to have uniquely discarded milk altogether[ citation needed ]. Prior to hatching, the milk glands would provide moisture to the leathery eggs, a situation still found in monotremes. [9]

The early mammaliaforms did have a harderian gland. In modern mammals, this is used for cleaning the fur, indicating that they, contrary to their Cynodont ancestors, had a furry covering. An insulative covering is necessary to keep a homeothermic animal warm if it is very small, less than 5 cm (1.97 in) long; [10] the 3.2 cm (1.35 in) Hadrocodium must have had fur, therefore, but the 10 cm (3.94 in) Morganucodon might not have needed it. The docodont Castorocauda , further removed from crown group mammals than Hadrocodium, had two layers of fur, guard hairs and underfur, as do mammals today. [11]

It is possible that early mammaliaforms had vibrissae ; Tritheledontidae, a group of Cynodonts, probably had whiskers. [12] A common ancestor of all therian mammals did so. [13] Indeed, some humans even still develop vestigial vibrissal muscles in the upper lip. [14] Thus, it is possible that the development of the whisker sensory system played an important role in mammalian development, more generally. [13]

Like monotremes today, the legs of early mammaliaforms were somewhat sprawling, giving a rather "reptilian" type of gait. However, there was a general tendency to have more erect forelimbs, forms like eutriconodonts even having a fundamentally modern forelimb anatomy while the hindlimbs remained "primitive"; [15] this tendency is in some effect still seen in modern therian mammals, which often have more sprawling hindlimbs. [16] In some forms, the hind feet likely bore a spur similar to those found in the platypus and echidnas. Such a spur would have been connected to a venom gland for protection or mating competition. [17]

Hadrocodium lacks the multiple bones in its lower jaw seen in reptiles. These are still retained, however, in earlier mammaliaforms. [18]

With the possible exception of Megazostrodon and Erythrotherium (as well as placental mammals), [19] all mammaliforms possess epipubic bones, a possibly synapomorphy with tritylodontids, which also have them. [20] These pelvic bones strengthen the torso and support abdominal and hindlimb musculature. They, however, prevent the expansion of the abdomen, and so force species that possess them to either give birth to larval young (as in modern marsupials), or produce minuscule eggs that hatch into larval young (as in modern monotremes). [21] Most mammaliforms, therefore, probably had the same constraints, and some species could have borne pouches.

Phylogeny

The cladogram below follows the analysis of Luo and colleagues in 2015. [22]

Mammaliamorpha

Expanded from above

Trechnotheria

Cladogram based on Rougier et al. (1996) [23] with Tikitherium included following Luo and Martin (2007). [3]

  Mammaliamorpha

See also

Related Research Articles

<span class="mw-page-title-main">Synapsida</span> Clade of tetrapods

Synapsida is one of the two major clades of vertebrate animals in the group Amniota, the other being the Sauropsida. The synapsids were the dominant land animals in the late Paleozoic and early Mesozoic, but the only group that survived into the Cenozoic are mammals. Unlike other amniotes, synapsids have a single temporal fenestra, an opening low in the skull roof behind each eye orbit, leaving a bony arch beneath each; this accounts for their name. The distinctive temporal fenestra developed about 318 million years ago during the Late Carboniferous period, when synapsids and sauropsids diverged, but was subsequently merged with the orbit in early mammals.

<span class="mw-page-title-main">Therapsida</span> Clade of tetrapods including mammals

Therapsida is a clade composing of a major group of eupelycosaurian synapsids that includes mammals and their ancestors and close relatives. Many of the traits today seen as unique to mammals had their origin within early therapsids, including limbs that were oriented more underneath the body, as opposed to the sprawling posture of many reptiles and salamanders.

<span class="mw-page-title-main">Cynodontia</span> Clade of therapsids

Cynodontia is a clade of eutheriodont therapsids that first appeared in the Late Permian, and extensively diversified after the Permian–Triassic extinction event. Mammals are cynodonts, as are their extinct ancestors and close relatives (Mammaliaformes), having evolved from advanced probainognathian cynodonts during the Late Triassic.

<i>Morganucodon</i> Early mammaliaform genus of the Triassic and Jurassic periods

Morganucodon 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 material. Most of this comes from Glamorgan in Wales, but fossils have also been found in Yunnan Province in China and various parts of Europe and North America. Some closely related animals (Megazostrodon) are known from exquisite fossils from South Africa.

<span class="mw-page-title-main">Allotheria</span> Extinct subclass of mammals

Allotheria is an extinct clade of mammals known from the Mesozoic and early Cenozoic. Shared characteristics of the group are the presence of lower molariform teeth equipped with longitudinal rows of cusps and enlarged incisors. Typically, the canine teeth are also lost. Allotheria includes Multituberculata, Gondwanatheria, and probably Haramiyida, although some studies have recovered haramiyidans to be basal mammaliaforms unrelated to multituberculates. Allotherians are often placed as crown group mammals, more closely related to living marsupials and placentals (Theria) than to monotremes or eutriconodonts, though some studies place the entirety of Allotheria outside of crown Mammalia.

<i>Megazostrodon</i> Extinct genus of mammaliaforms

Megazostrodon is an extinct genus of basal mammaliaforms belonging to the order Morganucodonta. It is approximately 200 million years old. Two species are known: M. rudnerae from the Early Jurassic of Lesotho and South Africa, and M. chenali from the Late Triassic of France.

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 and estimated to have been less than 10 cm (3.9 in) in head-body length, slightly smaller than the similar-proportioned Megazostrodon.

<span class="mw-page-title-main">Docodonta</span> Extinct order of mammaliaforms

Docodonta is an order of extinct Mesozoic mammaliaforms. They were among the most common mammaliaforms of their time, persisting from the Middle Jurassic to the Early Cretaceous across the continent of Laurasia. They are distinguished from other early mammaliaforms by their relatively complex molar teeth. Docodont teeth have been described as "pseudotribosphenic": a cusp on the inner half of the upper molar grinds into a basin on the front half of the lower molar, like a mortar-and-pestle. This is a case of convergent evolution with the tribosphenic teeth of therian mammals. There is much uncertainty for how docodont teeth developed from their simpler ancestors. Their closest relatives may have been certain Triassic "symmetrodonts", namely Woutersia, Delsatia, and Tikitherium.

Sinoconodon is an extinct genus of mammaliamorphs that appears in the fossil record of the Lufeng Formation of China in the Sinemurian stage of the Early Jurassic period, about 193 million years ago. While sharing many plesiomorphic traits with other non-mammaliaform cynodonts, it possessed a special, secondarily evolved jaw joint between the dentary and the squamosal bones, which in more derived taxa would replace the primitive tetrapod one between the articular and quadrate bones. The presence of a dentary-squamosal joint is a trait historically used to define mammals.

Tritylodontidae is an extinct family of small to medium-sized, highly specialized mammal-like cynodonts, with several mammalian traits including erect limbs, endothermy and details of the skeleton. They were the last-known family of the non-mammaliaform synapsids, persisting into the Early Cretaceous.

<span class="mw-page-title-main">Evolution of mammals</span> Derivation of mammals from a synapsid precursor, and the adaptive radiation of mammal species

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 like mammals. The lineage leading to today's mammals split up in the Jurassic; synapsids from this period include Dryolestes, more closely related to extant placentals and marsupials than to monotremes, as well as Ambondro, more closely related to monotremes. Later on, the eutherian and metatherian lineages separated; the metatherians are the animals more closely related to the marsupials, while the eutherians are those more closely related to the placentals. Since Juramaia, the earliest known eutherian, lived 160 million years ago in the Jurassic, this divergence must have occurred in the same period.

<span class="mw-page-title-main">Eutriconodonta</span> Extinct order of mammals

Eutriconodonta is an order of early mammals. Eutriconodonts existed in Asia, Africa, Europe, North and South America during the Jurassic and the Cretaceous periods. The order was named by Kermack et al. in 1973 as a replacement name for the paraphyletic Triconodonta.

<span class="mw-page-title-main">Haramiyida</span> Extinct order of mammaliaforms

Haramiyida is a possibly polyphyletic order of mammaliaform cynodonts or mammals of controversial taxonomic affinites. Their teeth, which are by far the most common remains, resemble those of the multituberculates. However, based on Haramiyavia, the jaw is less derived; and at the level of evolution of earlier basal mammals like Morganucodon and Kuehneotherium, with a groove for ear ossicles on the dentary. Some authors have placed them in a clade with Multituberculata dubbed Allotheria within Mammalia. Other studies have disputed this and suggested the Haramiyida were not crown mammals, but were part of an earlier offshoot of mammaliaformes instead. It is also disputed whether the Late Triassic species are closely related to the Jurassic and Cretaceous members belonging to Euharamiyida/Eleutherodontida, as some phylogenetic studies recover the two groups as unrelated, recovering the Triassic haramiyidians as non-mammalian cynodonts, while recovering the Euharamiyida as crown-group mammals closely related to multituberculates.

<span class="mw-page-title-main">Morganucodonta</span> Extinct order of mammaliaforms

Morganucodonta 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 and China. The morganucodontans were probably insectivorous and nocturnal, though like eutriconodonts some species attained large sizes and were carnivorous. Nocturnality is believed to have evolved in the earliest mammals in the Triassic as a specialisation that allowed them to exploit a safer, night-time niche, while most larger predators were likely to have been active during the day.

<span class="mw-page-title-main">Shuotheriidae</span> Extinct family of mammaliaforms

Shuotheriidae is a small family of Jurassic mammaliaforms whose remains are found in China, England and possibly Russia. They have been proposed to be close relatives of Australosphenida, together forming the clade Yinotheria. However, some studies suggest shuotheres are closer to therians than to monotremes, or that australosphenidans and therians are more closely related to each other than either are to shuotheres, with a 2024 study suggesting that shuotheriids were closely related to Docodonta outside of the Mammalia crown group.

<i>Kuehneotherium</i> Extinct genus of mammaliaforms

Kuehneotherium is an early mammaliaform genus, previously considered a holothere, that lived during the Late Triassic-Early Jurassic Epochs and is characterized by reversed-triangle pattern of molar cusps. Although many fossils have been found, the fossils are limited to teeth, dental fragments, and mandible fragments. The genus includes Kuehneotherium praecursoris and all related species. It was first named and described by Doris M. Kermack, K. A. Kermack, and Frances Mussett in November 1967. The family Kuehneotheriidae and the genus Kuehneotherium were created to house the single species Kuehneotherium praecursoris. Modeling based upon a comparison of the Kuehneotherium jaw with other mammaliaforms indicates it was about the size of a modern-day shrew between 4 and 5.5 g at adulthood.

<i>Megaconus</i> Extinct genus of mammaliaforms

Megaconus is an extinct genus of allotherian mammal from the Middle Jurassic Tiaojishan Formation of Inner Mongolia, China. The type and only species, Megaconus mammaliaformis was first described in the journal Nature in 2013. Megaconus is thought to have been a herbivore that lived on the ground, having a similar posture to modern-day armadillos and rock hyraxes. Megaconus was in its initial description found to be member of a group called Haramiyida. A phylogenetic analysis published along its description suggested that haramiyidans originated before the appearance of true mammals, but in contrast, the later description of the haramiyidan Arboroharamiya in the same issue of Nature indicated that haramyidans were true mammals. If haramiyidans are not mammals, Megaconus would be one of the most basal ("primitive") mammaliaforms to possess fur, and an indicator that fur evolved in the ancestors of mammals and not the mammals themselves. However, later studies cast doubt on the euharamiyidan intrepretation, instead finding it to be a basal allotherian mammal.

<span class="mw-page-title-main">Yinotheria</span> Subclass of mammals

Yinotheria is a proposed basal subclass clade of crown mammals uniting the Shuotheriidae, an extinct group of mammals from the Jurassic of Eurasia, with Australosphenida, a group of mammals known from the Jurassic to Cretaceous of Gondwana, which possibly include living monotremes. Today, there are only five surviving species of monotremes which live in Australia and New Guinea, consisting of the platypus and four species of echidna. Fossils of yinotheres have been found in Britain, China, Russia, Madagascar and Argentina. Contrary to other known crown mammals, they retained postdentary bones as shown by the presence of a postdentary trough. The extant members (monotremes) developed the mammalian middle ear independently.

<i>Ichthyoconodon</i> Extinct family of mammals

Ichthyoconodon is an extinct genus of eutriconodont mammal from the Lower Cretaceous of Morocco. It is notable for having been found in a unique marine location, and the shape of its teeth suggests an unusual, potentially fish-eating ecological niche. Analysis suggests it is part of a group of gliding mammals that includes Volaticotherium.

Vilevolodon is an extinct, monotypic genus of volant, arboreal euharamiyids from the Oxfordian age of the Late Jurassic of China. The type species is Vilevolodon diplomylos. The genus name Vilevolodon references its gliding capabilities, Vilevol, while don is a common suffix for mammalian taxon titles. The species name diplomylos refers to the dual mortar-and-pestle occlusion of upper and lower molars observed in the holotype; diplo, mylos.

References

  1. ↑ Abdala, F. (2007). "Redescription of Platycraniellus Elegans (Therapsida, Cynodontia) from the Lower Triassic of South Africa, and the cladistic relationships of eutheriodonts". Palaeontology. 53 (3): 591–618. Bibcode:2007Palgy..50..591A. doi: 10.1111/j.1475-4983.2007.00646.x . S2CID   83999660.
  2. 1 2 Rowe, T. S. (1988). "Definition, diagnosis, and origin of Mammalia" (PDF). Journal of Vertebrate Paleontology. 8 (3): 241–264. Bibcode:1988JVPal...8..241R. doi:10.1080/02724634.1988.10011708.
  3. 1 2 Luo, Zhe-Xi; Martin, Thomas (2007). "Analysis of Molar Structure and Phylogeny of Docodont Genera" (PDF). Bulletin of Carnegie Museum of Natural History. 39 (39): 27–47. doi:10.2992/0145-9058(2007)39[27:AOMSAP]2.0.CO;2. S2CID   29846648. Archived from the original (PDF) on 2016-03-03. Retrieved 2012-10-14.
  4. ↑ Datta, P. M. (2005). "Earliest mammal with transversely expanded upper molar from the Late Triassic (Carnian) Tiki Formation, South Rewa Gondwana Basin, India". Journal of Vertebrate Paleontology. 25 (1): 200–207. doi:10.1671/0272-4634(2005)025[0200:EMWTEU]2.0.CO;2. S2CID   131236175.
  5. ↑ Colbert, Edwin H.; Morales, Michael; Minkoff, Eli C. (2001). Colbert's evolution of the vertebrates: a history of the backboned animals through time (5th ed.). New York: Wiley. ISBN   978-0-471-38461-8.
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  7. ↑ Panciroli E., Benson RBJ., and Walsh S. 2017. The dentary of Wareolestes rex (Megazostrodontidae): a new specimen from Scotland and implications for morganucodontan tooth replacement. Papers in Palaeontology
  8. ↑ Hu, Yaoming; Meng, Jin; Clark, James M. "A New Tritylodontid from the Upper Jurassic of Xinjiang, China". Acta Palaeontologica Polonica. 54 (3): 385–391. doi : 10.4202/app.2008.0053.
  9. ↑ Oftedal, O.T. (2002). "The mammary gland and its origin during synapsid evolution". Journal of Mammary Gland Biology and Neoplasia. 7 (3): 225–252. doi:10.1023/A:1022896515287. PMID   12751889. S2CID   25806501.
  10. ↑ Ruben, J.A. & Jones, T.D. (2000). "Selective Factors Associated with the Origin of Fur and Feathers". American Zoologist. 40 (4): 585–596. doi: 10.1093/icb/40.4.585 .
  11. ↑ Qiang Ji; et al. (2006). "A Swimming Mammaliaform from the Middle Jurassic and Ecomorphological Diversification of Early Mammals" (PDF). Science. 311 (5764): 1123–27. Bibcode:2006Sci...311.1123J. doi:10.1126/science.1123026. PMID   16497926. S2CID   46067702.
  12. ↑ "Your Inner Fish: Episode Guide". PBS. 2014. Retrieved August 7, 2014.
  13. 1 2 Mitchinson, B.; Grant, R. A.; Arkley, K.; Rankov, V.; Perkon, I.; Prescott, T.J. (12 November 2011). "Active vibrissal sensing in rodents and marsupials". Phil. Trans. R. Soc. B. 366 (1581): 3037–3048. doi:10.1098/rstb.2011.0156. PMC   3172598 . PMID   21969685.
  14. ↑ Tamatsu, Yuichi; Tsukahara, Kazue; Hotta, Mitsuyuki; Shimada, Kazuyuki (August 2007). "Vestiges of vibrissal capsular muscles exist in the human upper lip". Clin Anat. 20 (6): 628–31. doi:10.1002/ca.20497. PMID   17458869. S2CID   21055062.
  15. ↑ Zofia Kielan-Jaworowska, Richard L. Cifelli, Zhe-Xi Luo (2004). "Chapter 7: Eutriconodontans". Mammals from the Age of Dinosaurs: origins, evolution, and structure . New York: Columbia University Press. pp.  216–248. ISBN   0-231-11918-6.{{cite book}}: CS1 maint: multiple names: authors list (link)
  16. ↑ Kielan−Jaworowska, Z. and Hurum, J.H. 2006. Limb posture in early mammals: Sprawling or parasagittal. Acta Palae− ontologica Polonica 51 (3): 393–406.
  17. ↑ Hurum, J.H.; Luo, Z-X; Kielan-Jaworowska, Z. (2006). "Were mammals originally venomous?" (PDF). Acta Palaeontologica Polonica. 51 (1): 1–11.
  18. ↑ Kemp, T. S. (2005). The Origin and Evolution of Mammals . Oxford University Press. p.  149. ISBN   0-19-850760-7.
  19. ↑ Jason A. Lillegraven, Zofia Kielan-Jaworowska, William A. Clemens, Mesozoic Mammals: The First Two-Thirds of Mammalian History, University of California Press, 17/12/1979 - 321
  20. ↑ Stephen Reily and Thomas White, Hypaxial Motor Patterns and the Function of Epipubic Bones in Primitive Mammals, ARTICLE in SCIENCE 299(5605):400-2 · FEBRUARY 2003, Department of Biological Sciences, Ohio University, Athens, OH 45701, USA. Impact Factor: 33.61 · doi : 10.1126/science.1074905 · Source: PubMed
  21. ↑ Michael L. Power, Jay Schulkin. The Evolution Of The Human Placenta. pp. 68–.
  22. ↑ Luo, Zhe-Xi; Gates, Stephen M.; Jenkins Jr., Farish A.; Amaral, William W.; Shubin, Neil H. (16 November 2015). "Mandibular and dental characteristics of Late Triassic mammaliaform Haramiyavia and their ramifications for basal mammal evolution". PNAS . 112 (51): E7101-9. Bibcode:2015PNAS..112E7101L. doi: 10.1073/pnas.1519387112 . PMC   4697399 . PMID   26630008.
  23. ↑ Rougier, G. W.; Wible, J. R.; Hopson, J. A. (1996). "Basicranial Anatomy of Priacodon fruitaensis (Triconodontidae, Mammalia) from the Late Jurassic of Colorado, and a Reappraisal of Mammaliaform Interrelationships" (PDF). American Museum Novitates (3183). American Museum of Natural History. ISSN   0003-0082.
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