Monotreme

Monotremes(/ˈmɒnətriːmz/) aremammalsof the orderMonotremata. They are the only group of living mammals thatlay eggs, rather than bearing live young. The extant monotreme species are theplatypusand the four species ofechidnas. Monotremes are typified by structural differences in their brains, jaws, digestive tract, reproductive tract, and other body parts, compared to the more common mammalian types. Although they are different from almost all mammals in that they lay eggs, like all mammals, the female monotremes nurse their young withmilk.
Monotremes have been considered by some authors to be members ofAustralosphenida, acladethat contains extinct mammals from theJurassicandCretaceousof Madagascar, South America, and Australia, but this categorization is disputed andtheir taxonomyis under debate.
All extant species of monotremes are indigenous toAustraliaandNew Guinea, although they were also present during the Late Cretaceous and Paleocene epochs in southern South America, implying that they were also present in Antarctica, though remains have not yet been found there.
The namemonotremederives from the Greek wordsμονός(monós'single') andτρῆμα(trêma'hole'), referring to thecloaca.
Like other mammals, monotremes areendothermicwith a high metabolic rate (though not as high as other mammals; see below); havehairon their bodies; produce milk through mammary glands to feed their young; have a single bone in their lower jaw; and have threemiddle earbones.
In common withmarsupials, monotremes lack the connective structure (corpus callosum) which inplacentalsis the primary communication route between the right and left brain hemispheres.Theanterior commissuredoes provide an alternate communication route between the two hemispheres, though, and in monotremes and marsupials it carries all thecommissural fibersarising from theneocortex, whereas in placental mammals the anterior commissure carries only some of these fibers.
Extant monotremes lack teeth as adults. Fossil forms and modern platypus young have a "tribosphenic" form ofmolars(with theocclusalsurface formed by threecuspsarranged in a triangle), which is one of the hallmarks of extant mammals. Some recent work suggests that monotremes acquired this form of molar independently of placentals and marsupials,although this hypothesis remains disputed.Tooth loss in modern monotremes might be related to their development ofelectrolocation.
Monotreme jaws are constructed somewhat differently from those of other mammals, and the jaw opening muscle is different. As in all true mammals, the tiny bones that conduct sound to the inner ear are fully incorporated into the skull, rather than lying in the jaw as in non-mammaliancynodontsand other pre-mammaliansynapsids; this feature, too, is now claimed to have evolved independently in monotremes andtherians,although, as with the analogous evolution of the tribosphenic molar, this hypothesis is disputed.Nonetheless, findings on the extinct speciesTeinolophosconfirm that suspended ear bones evolved independently among monotremes and therians.The external opening of the ear still lies at the base of the jaw.
The sequencing of the platypus genome has also provided insight into the evolution of a number of monotreme traits, such as venom andelectroreception, as well as showing some new unique features, such as monotremes possessing five pairs of sex chromosomes and that one of the X chromosomes resembles theZ chromosomeof birds,suggesting that the two sex chromosomes of marsupial and placentals evolved after the split from the monotreme lineage.Additional reconstruction through shared genes in sex chromosomes supports this hypothesis of independent evolution.This feature, along with some other genetic similarities with birds, such as shared genes related to egg-laying, is thought to provide some insight into themost recent common ancestorof thesynapsidlineage leading to mammals and thesauropsidlineage leading to birds and modern reptiles, which are believed to have split about 315 million years ago during theCarboniferous.The presence ofvitellogeningenes (a protein necessary for egg yolk formation) is shared with birds; the presence of thissymplesiomorphysuggests that the common ancestor of monotremes, marsupials, and placentals wasoviparous, and that this trait was retained in monotremes but lost in all other extant mammal groups. DNA analyses suggest that although this trait is shared and issynapomorphicwith birds, platypuses are still mammals and that the common ancestor of extant mammals lactated.
The monotremes also have extra bones in theshoulder girdle, including aninterclavicleandcoracoid, which are not found in other mammals. Monotremes retain a reptile-like gait, with legs on the sides of, rather than underneath, their bodies. The monotreme leg bears a spur in the ankle region; the spur is not functional in echidnas, but contains a powerfulvenomin the male platypus. This venom is derived fromβ-defensins, proteins that are present in mammals that create holes in viral and bacterial pathogens. Some reptile venom is also composed of different types of β-defensins, another trait shared with reptiles.It is thought to be an ancient mammalian characteristic, as many non-monotreme archaic mammal groups also possessvenomous spurs.
The key anatomical difference between monotremes and other mammals gives them their name;monotrememeans "single opening" in Greek, referring to the single duct (thecloaca) for their urinary, defecatory, and reproductive systems. Like birds and reptiles, monotremes have a single cloaca.Marsupials have aseparate genital tract, whereas most placental females have separate openings for reproduction (thevagina), urination (theurethra), and defecation (theanus). In monotremes, only semen passes through the penis while urine is excreted through the male's cloaca.The monotreme penis is similar to that of turtles and is covered by a preputial sac.Male monotremes do not have aprostateorseminal vesicles.
Monotremeeggsare retained for some time within the mother and receive nutrients directly from her, generally hatching within ten days after being laid – much shorter than the incubation period ofsauropsideggs.Much like newborn marsupials (and perhaps all non-placentals), newborn monotremes, called "puggles",are larval- and fetus-like and have relatively well-developed forelimbs that enable them to crawl around. Monotremes lackteats, so puggles crawl about more frequently than marsupial joeys in search of milk. This difference raises questions about the supposed developmental restrictions on marsupial forelimbs.
Rather than through teats, monotremes lactate from theirmammary glandsvia openings in their skin. All five extant species show prolonged parental care of their young, with low rates of reproduction and relatively long life-spans.
Monotremes are also noteworthy in their zygotic development: most mammalianzygotesgo through holoblasticcleavage, where the ovum splits into multiple, divisible daughter cells. In contrast, monotreme zygotes, like those of birds and reptiles, undergomeroblastic(partial) division. This means that the cells at the yolk's edge have cytoplasm continuous with that of the egg, allowing the yolk and embryo to exchange waste and nutrients with the surrounding cytoplasm.
Monotremes' metabolic rate is remarkably low by mammalian standards. The platypus has an averagebody temperatureof about 31 °C (88 °F) rather than the averages of 35 °C (95 °F) formarsupialsand 37 °C (99 °F) forplacentals.Research suggests this has been a gradual adaptation to the harsh, marginal environmental niches in which the few extant monotreme species have managed to survive, rather than a general characteristic of extinct monotremes.
Monotremes may have less developedthermoregulationthan other mammals, but recent research shows that they easily maintain a constant body temperature in a variety of circumstances, such as the platypus in icy mountain streams. Early researchers were misled by two factors: firstly, monotremes maintain a lower average temperature than most mammals; secondly, theshort-beaked echidna, much easier to study than the reclusive platypus, maintains normal temperature only when active; during cold weather, it conserves energy by "switching off" its temperature regulation. Understanding of this mechanism came when reduced thermal regulation was observed in thehyraxes, which are placentals.
The echidna was originally thought to experience norapid eye movement sleep(REM).However, a more recent study showed that REM sleep accounted for about 15% of sleep time observed on subjects at an environmental temperature of 25 °C (77 °F). Surveying a range of environmental temperatures, the study observed very little REM at reduced temperatures of 15 °C (59 °F) and 20 °C (68 °F), and also a substantial reduction at the elevated temperature of 28 °C (82 °F).
Monotreme milk contains a highly expressed antibacterial protein not found in other mammals, perhaps to compensate for the more septic manner of milk intake associated with the absence of teats.
During the course of evolution, the monotremes have lost thegastric glandsnormally found in mammalianstomachsas an adaptation to their diet.As such, by some definitions, they do not have stomachs as an organ,although the term is widely used in studies of monotreme anatomy.Monotremes synthesize L-ascorbic acidonly in the kidneys.
Both the platypus and echidna species havespurson their hind limbs. The echidna spurs are vestigial and have no known function, while the platypus spurs contain venom.Molecular data show that the main component of platypus venom emerged before the divergence of platypus and echidnas, suggesting that the most recent common ancestor of these taxa was also possibly a venomous monotreme.
The traditional "Theriahypothesis" states that the divergence of the monotreme lineage from theMetatheria(marsupial) andEutheria(placental) lineages happened prior to the divergence between marsupials and placentals, and this explains why monotremes retain a number ofprimitivetraits presumed to have been present in thesynapsidancestors of later mammals, such as egg-laying.Most morphological evidence supports the Theria hypothesis, but one possible exception is a similar pattern of tooth replacement seen in monotremes and marsupials, which originally provided the basis for the competing "Marsupionta" hypothesis in which the divergence between monotremes and marsupials happened later than the divergence between these lineages and the placentals. Van Rheede (2005) concluded that the genetic evidence favors the Theria hypothesis,and this hypothesis continues to be the more widely accepted one.
Monotremes are conventionally treated as comprising a single orderMonotremata. The entire grouping is also traditionally placed into a subclassPrototheria, which was extended to include severalfossilorders, but these are no longer seen as constituting a group allied to monotreme ancestry. A controversial hypothesis now relates the monotremes to a different assemblage of fossil mammals in acladetermedAustralosphenida, a group of mammals from the Jurassic and Cretaceous of Madagascar, South America and Australia, that sharetribosphenic molars.However, in a 2022 review of monotreme evolution, it was noted thatTeinolophos, the oldest (Barremian~ 125 million years ago) and the most primitive monotreme differed substantially from non-monotreme australosphenidans in having five molars as opposed to the three present in non-monotreme australosphenidians. Aptian and Cenomanian monotremes of the family Kollikodontidae (113–96.6 ma) have four molars. This suggests that the monotremes are likely to be unrelated to the australosphenidan tribosphenids.
The time when the monotreme line diverged from other mammalian lines is uncertain, but one survey of genetic studies gives an estimate of about 220 million years ago,while others have posited younger estimates of 163 to 186 million years ago (though the already eutherianJuramaiais dated to 161–160 million years ago).Teinolophoslike modern monotremes displays adaptations to elongation and increased sensory perception in the jaws, related tomechanoreceptionorelectroreception.
Molecular clockand fossil dating give a wide range of dates for the split between echidnas and platypuses, with one survey putting the split at 19–48 million years ago,but another putting it at 17–89 million years ago.It has been suggested that both the short-beaked and long-beaked echidna species are derived from a platypus-like ancestor.
The precise relationships among extinct groups of mammals and modern groups such as monotremes are uncertain, butcladisticanalyses usually put thelast common ancestor(LCA) of placentals and monotremes close to the LCA of placentals andmultituberculates, whereas some suggest that the LCA of placentals and multituberculates was more recent than the LCA of placentals and monotremes.
The firstMesozoicmonotreme to be discovered was theCenomanian(100–96.6 Ma)Steropodon galmanifrom Lightning Ridge, New South Wales.Biochemical and anatomical evidence suggests that the monotremes diverged from the mammalian lineage before the marsupials and placentals arose. The only Mesozoic monotremes areTeinolophos(Barremian, 126 Ma),SundriusandKryoryctes(Albian, 113–108 Ma), andDharragarra,Kollikodon,Opalios,Parvopalus,Steropodon, andStirtodon(all Cenomanian, 100.2–96.6 Ma) from Australian deposits, andPatagorhynchus(Maastrichtian) fromPatagoniandeposits in theCretaceous, indicating that monotremes were diversifiying by the early Late Cretaceous.Monotremes have been found in the latest Cretaceous and Paleocene of southern South America, so one hypothesis is that monotremes arose in Australia in theLate JurassicorEarly Cretaceous, and that some migrated acrossAntarcticatoSouth America, both of which were still united with Australia at that time.This direction of migration is the opposite of that hypothesized for Australia's other dominant mammal group, themarsupials, which likely migrated across Antarctica to Australia from South America.
In 2024, a prominent assemblage of early monotremes was described from the Cenomanian deposits (100–96.6 Ma) of theGriman Creek Formationin Lightning Ridge, New South Wales. One of these, the fossil jaw fragment ofDharragarra, is the oldest known platypus-like fossil.The durophagousKollikodon, the pseudotribosphenicSteropodon, andStirtodon,Dharragarra,Opalios, andParvopalusoccur in the same Cenomanian deposits. Oligo-Miocene fossils of the toothed platypusObdurodonhave also been recovered from Australia, and fossils of a 63 million-year old platypus relative occur in southern Argentina (Monotrematum), see fossil monotremes below. The extant platypus genusOrnithorhynchusin also known from Pliocene deposits, and the oldest fossil tachyglossids are Pleistocene (1.7 Ma) in age.
ExceptingOrnithorhynchus anatinus, all the animals listed in this section are known only from fossils. Some family designations are hesitant, given the fragmentary nature of the specimens.