Mesosaur

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Mesosaurs
Temporal range: Cisuralian, 299–270.6  Ma
Mesosaurus BW.jpg
Mesosaurus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Parareptilia
Order: Mesosauria
Seeley, 1892
Family: Mesosauridae
Baur, 1889
Genera

Mesosaurs ("middle lizards") were a group of small aquatic reptiles that lived during the early Permian period (Cisuralian), roughly 299 to 270 million years ago. Mesosaurs were the first known aquatic reptiles, having apparently returned to an aquatic lifestyle from more terrestrial ancestors. It is uncertain which and how many terrestrial traits these ancestors displayed; recent research cannot establish with confidence if the first amniotes were fully terrestrial, or only amphibious. [1] Most authors consider mesosaurs to have been aquatic, [1] [2] although adult animals may have been amphibious, rather than completely aquatic, as indicated by their moderate skeletal adaptations to a semiaquatic lifestyle. [3] [4] Similarly, their affinities are uncertain; they may have been among the most basal sauropsids [5] [6] or among the most basal parareptiles (in the case of which parareptiles were basal sauropsids). [7] [8]

Contents

Phylogeny

The phylogenetic position of mesosaurs has an important bearing on the definition of Reptilia. In one of the first major phylogenetic studies of amniotes (vertebrates laying eggs on land) Gauthier et al. (1988) placed Mesosauridae in a group called Parareptilia. [9] Parareptilia means "at the side of reptiles" and was placed outside the clade Reptilia, which was considered a crown group. As a crown group, Reptilia included the most recent common ancestor of the then believed to be the two main lineages of living reptiles -anapsids (specifically turtles) and diapsids (all other living reptiles)- and all descendants of that common ancestor. This view of placing turtles outside of diapsids is now outdated and the majority of modern paleontologists believe that the Testudines (turtles and allies) are descended from diapsid reptiles that lost their temporal fenestrae. More recent morphological phylogenetic studies with this in mind placed turtles firmly within diapsids, [10] [11] [12] [13] and, more commonly, as a sister taxon to Archosauria (made up of crocodiles, dinosaurs -including birds- and allies). [14] Furthermore, Anapsida is rarely considered a valid clade in recent phylogenetic analyses. [15] [16] In this sense, Reptilia was a node-based taxon because the first reptilian common ancestor would have been a "node" on the phylogenetic tree. Under this phylogeny, many extinct forms traditionally regarded as reptiles including mesosaurs were excluded from the group because they were outside the node. [17]

Gauthier et al., 1988 [9]
Amniota  

Synapsida

MESOSAURIDAE

Procolophonia

Millerettidae

Pareiasauria

  Reptilia  
  Anapsida  

Captorhinidae

Testudines

  Romeriida  

Paleothyris

  Diapsida  

Araeoscelidia

Sauria

Fossil of a South American Mesosaur Mesosaur Fossil.jpg
Fossil of a South American Mesosaur

The study of Laurin and Reisz (1995) was the second major phylogenetic analysis of amniotes. [18] Like Gauthier et al., Laurin and Reisz used Reptilia as a crown group and placed mesosaurs outside the group. Their phylogeny differed in that the parareptiles of Gauthier et al. were now regarded as close relatives of turtles, within crown group Reptilia. Laurin and Reisz adopted the name Sauropsida as a node-based taxon including the last common ancestor of mesosaurs and Reptilia. Traditionally, amniotes are divided into two groups: a mammal lineage called Synapsida and a reptile lineage called either Reptilia or Sauropsida. In fact, the study of Gauthier (1994) defined Sauropsida as all amniotes more closely related to reptiles than to mammals, which meant that Sauropsida was a stem-based taxon encompassing the entire reptilian lineage or reptilian "stem" of Amniota (Synapsida was the mammalian stem). Under this phylogeny, the only group that prevents Sauropsida from being equivalent to Reptilia is mesosaurs. [17]

Laurin and Reisz (1995) [18]
Amniota  

Synapsida

MESOSAURIDAE

  Reptilia  
  Anapsida  

Millerettidae

  Procolophonia  

Pareiasauria

Testudinomorpha

Procolophonidae

Testudines

  Eureptilia  

Captorhinidae

  Romeriida  

Paleothyris

  Diapsida  

Araeoscelidia

Sauria

More recent phylogenetic analyses, such as that of Modesto (1999), support that of Gauthier et al. (1988) by placing mesosaurs with parareptiles. [7] However, these phylogenies follow Laurin and Reisz (1995) in placing Parareptilia within crown-group Reptilia, meaning that mesosaurs are once again members of Reptilia. Using Laurin and Reisz's node-based definition of Sauropsida as "The last common ancestor of mesosaurs, testudines and diapsids, and all its descendants", [18] Sauropsida and Reptilia are equivalent groupings; mesosaurs and testudines are more closely related to each other than either group is to diapsids, [a] meaning that the clade containing testudines and diapsids (which would be crown-group Reptilia) must also contain mesosaurs. Since Reptilia was named earlier than Sauropsida, it is used most often in modern phylogenetic analyses. [17]

Modesto, 1999 [7]
Amniota  

Synapsida

  Reptilia  
  Anapsida  

MESOSAURIDAE

Millerettidae

  Procolophonia  

Pareiasauria

Testudinomorpha

Procolophonidae

Testudines

  Eureptilia  

Captorhinidae

  Romeriida  

Paleothyris

  Diapsida  

Araeoscelidia

Sauria

A 2017 phylogenetic analysis by Laurin (who had previously published the 1995 study) and Piñeiro recovered mesosaurs as a basal member of Sauropsida/Reptilia and no longer present within Parareptilia, with Parareptilia being redefined as including former members of Procolophonomorpha (found to be paraphyletic), Millerosauria, Pareiasauria, and Pantestudines, with the latter two being found to be sister groups to one another. Parareptilia was also found to actually nest inside Diapsida as the sister group to Neodiapsida. [19]

Laurin & Piñeiro, 2017 [19]
Amniota  

Synapsida

  Sauropsida  /  Reptilia  

MESOSAURIDAE

Captorhinidae

Diapsida

Araeoscelida

Paleothyris

Neodiapsida

Acleistorhinidae

Millerosauria

Owenettidae

Procolophonidae

Pareiasauria

Pantestudines

In 2012 it was revealed that Mesosaurus has holes at the back of the skull called lower temporal fenestrae, a characteristic once thought to be present only in synapsids and diapsids. [20] This confirmed the previous results of German paleontologist Friedrich von Huene, already published in 1941 [21] The condition in the skull of Mesosaurus is most similar to that in synapsid skulls because both lack the upper temporal fenestrae of diapsids. Lower temporal fenestrae are so far known only in Mesosaurus, but may be present in all mesosaurs. The presence or absence of temporal fenestrae is an important consideration in the phylogeny of mesosaurs and other amniotes because the three major groups of amniotes -Synapsida, Diapsida, and Anapsida- have been named after the number of holes in their skull; Diapsida means "two arches" in reference to the two bars that close off the upper and lower fenestra, Synapsida means "fused arch" in reference to a single bar at the bottom of the skull closing a single fenestra, and Anapsida means "no arch" in reference to skulls that lack any bars or fenestrae. Mesosaurs were traditionally classified as anapsids because they were thought to have lacked temporal fenestrae. However, the occurrence of fenestrae in amniotes has been recognized a highly variable feature within the group for many years prior to their discovery in Mesosaurus; many anapsids such as Candelaria , Bolosaurus , and lanthanosuchoids possess lower temporal fenestrae.

The skull of a generalized anapsid. Skull anapsida 1.svg
The skull of a generalized anapsid.
The skull of a generalized synapsid. Skull synapsida 1.svg
The skull of a generalized synapsid.

The phylogenetic position of mesosaurs influences the current understanding of how amniotes evolved temporal fenestrae. If the phylogeny produced by Laurin and Reisz (1995) is correct in that mesosaurs are basal sauropsids, the lower temporal fenestra may be a primitive feature in amniotes, present in amniote's most recent common ancestor. Synapsids would have retained their fenestrae, and so too would sauropsids except for turtles and most parareptiles. Another possibility under Laurin and Reisz's phylogeny is that lower temporal fenestrae evolved independently in mesosaurs, synapsids, diapsids, and some parareptiles, and that the lack of fenestrae is a primitive feature in amniotes. If instead mesosaurs are members of Parareptilia, the presence of temporal fenestrae is probably not a primitive feature in amniotes, and the lower temporal fenestrae in mesosaurs may be characteristic of a lineage of basal parareptiles that also includes fenestra-bearing lanthanosuchoids and Bolosaurus. [20]

Biology

They have long been thought to have been coastal forms that probably inhabited relatively shallow water, [22] but recent research suggests that at least those from Uruguay inhabited a hypersaline environment, rather than a coastal marine environment. [23] Recently described embryos show that pachyostosis of the ribs (which were thicker and denser than in terrestrial tetrapods) developed even before hatching, which suggests that mesosaurs were able to swim at birth, or shortly thereafter. They were apparently not very fast swimmers, with an optimal swimming speed estimated to have been between 0.15 and 0.86 m/s, but this must have been somewhat faster than the speed of their main prey, the pygocephalomorph crustaceans. [24] Their reproductive mode is somewhat uncertain because association between adults and possible embryos in utero suggests viviparity, as in many aquatic reptiles, but a potentially isolated egg has also been found. [25]

Recently, evidence of predation on both pygocephalomorph crustaceans and members of their own species has been established. It is thought that mesosaurs were in general adapted to hypersaline habitats. [26]

A study on the vertebral column torso and tail proportions of Mesosaurus suggests that, while juveniles may have been fully aquatic, adults might have spent some time on land; this is further vindicated by the rarity of adult animals in aquatic settings and some faeces showing signs of drying fracture. However, how terrestrial they were is difficult to say, as this same study states that terrestrial foraging would have been difficult due to their specializations to an aquatic life. [4]

Notes

  1. ^ Studies using molecular phylogenetics, which examine the genes and proteins of living organisms, suggest that testudines (turtles) are diapsids. These studies show that mesosaurs do not form a clade with turtles that excludes diapsids, but fossil evidence still suggests that mesosaurs form a group with parareptiles. In most recent studies, Reptilia is not used as a crown group and still contains mesosaurs and Parareptilia.

Related Research Articles

<span class="mw-page-title-main">Anapsid</span> Subclass of reptiles

An anapsid is an amniote whose skull lacks one or more skull openings near the temples. Traditionally, the Anapsida are the most primitive subclass of amniotes, the ancestral stock from which Synapsida and Diapsida evolved, making anapsids paraphyletic. It is however doubtful that all anapsids lack temporal fenestra as a primitive trait, and that all the groups traditionally seen as anapsids truly lacked fenestra.

<span class="mw-page-title-main">Amniote</span> Clade of tetrapods including reptiles, birds and mammals

Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibian ancestors during the Carboniferous period and further diverged into two groups, namely the sauropsids and synapsids. They are distinguished from the other living tetrapod clade — the non-amniote lissamphibians — by the development of three extraembryonic membranes, thicker and keratinized skin, and costal respiration.

<span class="mw-page-title-main">Diapsid</span> Clade of amniote tetrapods with two holes in each side of their skulls

Diapsids are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The group first appeared about three hundred million years ago during the late Carboniferous period. All diapsids other than the most primitive ones in the clade Araeoscelidia are sometimes placed into the clade Neodiapsida. The diapsids are extremely diverse, and include birds and all modern reptile groups, including turtles, which were historically thought to lie outside the group. Although some diapsids have lost either one hole (lizards), or both holes, or have a heavily restructured skull, they are still classified as diapsids based on their ancestry. At least 17,084 species of diapsid animals are extant: 9,159 birds, and 7,925 snakes, lizards, tuatara, turtles, and crocodiles.

<span class="mw-page-title-main">Sauropsida</span> Taxonomic clade

Sauropsida is a clade of amniotes, broadly equivalent to the class Reptilia, though typically used in a broader sense to include both extinct stem-group relatives of modern reptiles, as well as birds. The most popular definition states that Sauropsida is the sibling taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early synapsids have historically been referred to as "mammal-like reptiles", all synapsids are more closely related to mammals than to any modern reptile. Sauropsids, on the other hand, include all amniotes more closely related to modern reptiles than to mammals. This includes Aves (birds), which are now recognized as a subgroup of archosaurian reptiles despite originally being named as a separate class in Linnaean taxonomy.

<span class="mw-page-title-main">Archosauromorpha</span> Infraclass of reptiles

Archosauromorpha is a clade of diapsid reptiles containing all reptiles more closely related to archosaurs rather than lepidosaurs. Archosauromorphs first appeared during the late Middle Permian or Late Permian, though they became much more common and diverse during the Triassic period.

<span class="mw-page-title-main">Neodiapsida</span> Clade of reptiles

Neodiapsida is a clade, or major branch, of the reptilian family tree, typically defined as including all diapsids apart from some early primitive types known as the araeoscelidians. Modern reptiles and birds belong to the neodiapsid subclade Sauria.

<span class="mw-page-title-main">Pareiasauria</span> Extinct clade of reptiles

Pareiasaurs are an extinct clade of large, herbivorous parareptiles. Members of the group were armoured with osteoderms which covered large areas of the body. They first appeared in southern Pangea during the Middle Permian, before becoming globally distributed during the Late Permian. Pareiasaurs were the largest reptiles of the Permian, reaching sizes equivalent to those of contemporary therapsids. Pareiasaurs became extinct in the Permian–Triassic extinction event.

<span class="mw-page-title-main">Parareptilia</span> Subclass of reptiles

Parareptilia ("near-reptiles") is a subclass or clade of basal sauropsids/reptiles, typically considered the sister taxon to Eureptilia. Parareptiles first arose near the end of the Carboniferous period and achieved their highest diversity during the Permian period. Several ecological innovations were first accomplished by parareptiles among reptiles. These include the first reptiles to return to marine ecosystems (mesosaurs), the first bipedal reptiles, the first reptiles with advanced hearing systems, and the first large herbivorous reptiles. The only parareptiles to survive into the Triassic period were the procolophonoids, a group of small generalists, omnivores, and herbivores. The largest family of procolophonoids, the procolophonids, rediversified in the Triassic, but subsequently declined and became extinct by the end of the period.

<span class="mw-page-title-main">Procolophonia</span> Extinct suborder of reptiles

Procolophonia is an extinct suborder (clade) of herbivorous reptiles that lived from the Middle Permian till the end of the Triassic period. They were originally included as a suborder of the Cotylosauria but are now considered a clade of Parareptilia. They are closely related to other generally lizard-like Permian reptiles such as the Millerettidae, Bolosauridae, Acleistorhinidae, and Lanthanosuchidae, all of which are included under the Anapsida or "Parareptiles".

<span class="mw-page-title-main">Procolophonomorpha</span> Order of reptiles (fossil)

Procolophonomorpha is an order or clade containing most parareptiles. Many papers have applied various definitions to the name, though most of these definitions have since been considered synonymous with modern parareptile clades such as Ankyramorpha and Procolophonia. The current definition of Procolophonomorpha, as defined by Modesto, Scott, & Reisz (2009), is that of as a stem-based group containing Procolophon and all taxa more closely related to it than to Milleretta. It constitutes a diverse assemblage that includes a number of lizard-like forms, as well as more diverse types such as the pareiasaurs. Lee 1995, 1996, 1997 argues that turtles evolved from pareiasaurs, but this view is no longer considered likely. Rieppel and deBraga 1996 and deBraga and Rieppel, 1997 argue that turtles evolved from sauropterygians, and there is both molecular and fossil (Pappochelys) evidence for the origin of turtles among diapsid reptiles.

<i>Mesosaurus</i> Extinct genus of reptile from the early Permian of South Africa

Mesosaurus is an extinct genus of reptile from the Early Permian of southern Africa and South America. Along with it, the genera Brazilosaurus and Stereosternum, it is a member of the family Mesosauridae and the order Mesosauria. Mesosaurus was long thought to have been one of the first marine reptiles, although new data suggests that at least those of Uruguay inhabited a hypersaline water body, rather than a typical marine environment. In any case, it had many adaptations to a fully aquatic lifestyle. It is usually considered to have been anapsid, although Friedrich von Huene considered it to be a synapsid. Recent study of Mesosauridae phylogeny places the group as either the basal most clade within Parareptilia or the basal most clade within Sauropsida despite the skull of Mesosaurus possessing the "Synapsid condition" of one temporal fenestra.

<i>Claudiosaurus</i> Extinct genus of reptiles

Claudiosaurus is an extinct genus of diapsid reptiles from the Late Permian Sakamena Formation of the Morondava Basin, Madagascar. It has been suggested to be semi-aquatic.

<span class="mw-page-title-main">Araeoscelidia</span> Extinct clade of reptiles

Araeoscelidia or Araeoscelida is a clade of extinct diapsid reptiles superficially resembling lizards, extending from the Late Carboniferous to the Early Permian. The group contains the genera Araeoscelis, Petrolacosaurus, the possibly aquatic Spinoaequalis, and less well-known genera such as Kadaliosaurus and Zarcasaurus. This clade is usually considered to be the sister group to all later diapsids.

<i>Stereosternum</i>

Stereosternum tumidum is an extinct genus of mesosaur marine reptile from the Early Permian of Brazil and also the Great Karoo Basin of South Africa. The taxon mesosaur is a monophyletic group containing Brazilosaurus sanpauloensis and Mesosaurus tenuidens.

<i>Eunotosaurus</i> Extinct genus of reptiles

Eunotosaurus is an extinct genus of amniote, possibly a close relative of turtles. Eunotosaurus lived in the late Middle Permian and fossils can be found in the Karoo Supergroup of South Africa. Eunotosaurus resided in the swamps of southern Africa. Its ribs were wide and flat, forming broad plates similar to a primitive turtle shell, and the vertebrae were nearly identical to those of some turtles. Accordingly, it is often considered as a possible transitional fossil between turtles and their prehistoric ancestors. However, it is possible that these turtle-like features evolved independently of the same features in turtles, since other anatomical studies and phylogenetic analyses suggest that Eunotosaurus may instead have been a parareptile, an early-diverging neodiapsid unrelated to turtles, or a synapsid.

<i>Acleistorhinus</i> Extinct genus of reptiles

Acleistorhinus (ah-kles-toe-RYE-nuss) is an extinct genus of parareptile known from the Early Permian of Oklahoma. It is notable for being the earliest known anapsid reptile yet discovered. The morphology of the lower temporal fenestra of the skull of Acleistorhinus bears a superficial resemblance to that seen in early synapsids, a result of convergent evolution. Only a single species, A. pteroticus, is known, and it is classified in the Family Acleistorhinidae, along with Colobomycter.

<i>Owenetta</i> Extinct genus of reptiles

Owenetta is an extinct genus of owenettid procolophonian parareptile. Fossils have been found from the Beaufort Group in the Karoo Basin of South Africa. Although most procolophonians lived during the Triassic, Owenetta existed during the Wuchiapingian and Changhsingian stages of the Late Permian as well as the early Induan stage of the Early Triassic. It is the type genus of the family Owenettidae, and can be distinguished from other related taxa in that the posterior portion of the supratemporal bears a lateral notch and that the pineal foramen is surrounded by a depressed parietal surface on the skull table.

<span class="mw-page-title-main">Evolution of reptiles</span> Origin and diversification of reptiles through geologic time

Reptiles arose about 320 million years ago during the Carboniferous period. Reptiles, in the traditional sense of the term, are defined as animals that have scales or scutes, lay land-based hard-shelled eggs, and possess ectothermic metabolisms. So defined, the group is paraphyletic, excluding endothermic animals like birds that are descended from early traditionally-defined reptiles. A definition in accordance with phylogenetic nomenclature, which rejects paraphyletic groups, includes birds while excluding mammals and their synapsid ancestors. So defined, Reptilia is identical to Sauropsida.

<span class="mw-page-title-main">Pantestudines</span> Clade of reptiles

Pantestudines or Pan-Testudines is the group of all reptiles more closely related to turtles than to any other living animal. It includes both modern turtles and all of their extinct relatives. Pantestudines with a complete shell are placed in the clade Testudinata.

<i>Pappochelys</i> Extinct genus of reptiles

Pappochelys is an extinct genus of diapsid reptile possibly related to turtles. The genus contains only one species, Pappochelys rosinae, from the Middle Triassic of Germany, which was named by paleontologists Rainer Schoch and Hans-Dieter Sues in 2015. The discovery of Pappochelys provides strong support for the placement of turtles within Diapsida, a hypothesis that has long been suggested by molecular data, but never previously by the fossil record. It is morphologically intermediate between the definite stem-turtle Odontochelys from the Late Triassic of China and Eunotosaurus, a reptile from the Middle Permian of South Africa.

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