Protorosauria

Last updated

Protorosaurs
Temporal range: Latest Middle Permian - Late Triassic,
260–201.3  Ma
Protorosaurus speneri fossil.jpg
Fossil specimen of Protorosaurus speneri , Teyler's Museum
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Order:† Protorosauria
Huxley, 1871
Subtaxa

Protorosauria is an extinct, likely paraphyletic group of basal archosauromorph reptiles from the latest Middle Permian (Capitanian stage) to the end of the Late Triassic (Rhaetian stage) of Asia, Europe and North America. It was named by the English anatomist and paleontologist Thomas Henry Huxley in 1871 as an order, originally to solely contain Protorosaurus . Other names which were once considered equivalent to Protorosauria include Prolacertiformes and Prolacertilia. [1]

Contents

Protorosaurs are distinguished by their long necks formed by elongated cervical vertebrae, which have ribs that extend backward to the vertebrae behind them. Protorosaurs also have a gap between the quadrate bones and the jugal bones in the back of the skull near the jaw joint, making their skulls resemble those of lizards. [1] While previously thought to be monophyletic, the group is now thought to consist of various groups of basal archosauromorph reptiles that lie outside Crocopoda, [2] though some recent studies have recovered the group as monophyletic. [3] A number of members of Protosauria have been found to belong to a monophyletic group (though not including Protorosaurus) which was named Tanysauria in 2024. [4]

Classification

Protorosauria was considered to be a synonym of Prolacertiformes for many years. [5]

Since 1998, many phylogenetic analyses have found Protorosauria, as used in its widest sense, to be a polyphyletic or paraphyletic taxon. Protorosaurus, Macrocnemus , tanystropheids, and various other protorosaurs are usually placed near the base of Archosauromorpha, while Prolacerta and Pamelaria , two Gondwanan Triassic protorosaurs, are now thought to be in a more derived position as close relatives of Archosauriformes. [6] Most phylogenetic analyses since 1998 have found a strongly supported clade that includes only the genus Prolacerta and the Archosauriformes. [7]

For this reason Prolacerta, Pamelaria, and several other related forms (collectively called prolacertids) have been removed from Protorosauria. Because the name Prolacertiformes is defined based on the genus Prolacerta, the name Protorosauria is used for the remaining group.

Only recently has Protorosauria been defined in a phylogenetic sense as the most inclusive clade containing taxa such as Protorosaurus , Macrocnemus , and Tanystropheus . Analyses, such as Dilkes (1998), Sues (2003), Modesto & Sues (2004), Rieppel, Fraser & Nosotti (2003), Rieppel, Li & Fraser (2008), Gottmann-Quesada and Sander (2009) and Renesto et al. (2010), [7] [8] [9] [10] [11] recovered a large Protorosauria, that includes Protorosaurus , Drepanosauridae (and relatives) and Tanystropheidae (and relatives). However, some analysis found Protorosaurus (and sometimes the closely related Czatkowiella ) to be more advanced [12] or more basal [13] than the node Drepanosauridae+Tanystropheidae, but always more basal than Prolacerta.

Some studies still use the term Prolacertiformes to include prolacertids and traditional protorosaurs, while restricting the term Protorosauria to the smallest clade that includes Protorosaurus, Macrocnemus, and Tanystropheus; thus Protorosauria is a true clade, while Prolacertiformes is an evolutionary grade of early archosauromorphs. [14]

Pritchard et al. (2015), [15] Nesbitt et al. (2015), [16] Ezcurra (2016) [17] and Spiekman et al., 2021 [2] found that even this definition of Protorosauria, like Prolacertiformes, was an unnatural group of various non-Crocopodan archosauromorphs. These studies found that tanystropheids were archosauromorphs more closely related to crocopods than to Protorosaurus. Nevertheless, Ezcurra noted that archosauromorph systematics required further study, and that phylogenetic support for Protorosauria being a natural group was only barely weaker than the support for the group being unnatural.

Included groups

The Protorosauria includes the Permian genus Protorosaurus , closely related to Czatkowiella . [18] A wide variety of Permian and Triassic reptiles have been classified within Protorosauria, including the arboreal gliding reptile Sharovipteryx and the aquatic tanystropheids, which have extremely long necks.

Another enigmatic group of Triassic reptiles, the Drepanosauromorpha, have often been classified as belonging to the Protorosauria. [19]

Pterosaurs have also been proposed as protorosaurs or close relatives of them, [20] although they are now regarded as a more derived group of archosaurs.

While Senter (2004) reassigned the bizarre, arboreal drepanosaurids and Longisquama to a group of more primitive diapsids called Avicephala, [21] subsequent studies failed to find the same result, instead supporting the hypothesis that they were protorosaurs.

Cladogram

The following cladogram shows the position of Protorosauria among the Sauria sensu Sean P. Modesto and Hans-Dieter Sues (2004). [7]

Most recent studies have recovered Protorosauria as a whole as a paraphyletic, cladogram after Spiekman et al. 2021 [2]

Diapsida

Although Protorosauria as a whole is often found to be a paraphyletic, a large group of former "protorosaurs" (excluding Protorosaurus) is frequently found to be monophyletic. This clade was given the name "Tanysauria" by Spiekman et al. in 2024. [4]

Related Research Articles

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

Dinocephalosaurus is a genus of long necked, aquatic protorosaur that inhabited the Triassic seas of China. The genus contains the type and only known species, D. orientalis, which was named by Chun Li in 2003. Unlike other long-necked protorosaurs, Dinocephalosaurus convergently evolved a long neck not through elongation of individual neck vertebrae, but through the addition of neck vertebrae that each had a moderate length. As indicated by phylogenetic analyses, it belonged in a separate lineage that also included at least its closest relative Pectodens, which was named the Dinocephalosauridae in 2021. Like tanystropheids, however, Dinocephalosaurus probably used its long neck to hunt, utilizing the fang-like teeth of its jaws to ensnare prey; proposals that it employed suction feeding have not been universally accepted. It was probably a marine animal by necessity, as suggested by the poorly-ossified and paddle-like limbs which would have prevented it from going ashore.

<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.

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

Tanystropheus is an extinct genus of archosauromorph reptile which lived during the Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than the torso and tail combined. The neck was composed of 13 vertebrae strengthened by extensive cervical ribs. Tanystropheus is one of the most well-described non-archosauriform archosauromorphs, known from numerous fossils, including nearly complete skeletons. Some species within the genus may have reached a total length of 6 meters (20 ft), making Tanystropheus the longest non-archosauriform archosauromorph as well. Tanystropheus is the namesake of the family Tanystropheidae, a clade collecting many long-necked Triassic archosauromorphs previously described as "protorosaurs" or "prolacertiforms".

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

Protorosaurus is an extinct genus of reptile. Members of the genus lived during the late Permian period in what is now Germany and Great Britain. Once believed to have been an ancestor to lizards, Protorosaurus is now known to be one of the oldest and most primitive members of Archosauromorpha, the group that would eventually lead to archosaurs such as crocodilians and dinosaurs.

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

Drepanosaurs are a group of extinct reptiles that lived between the Carnian and Rhaetian stages of the late Triassic Period, approximately between 230 and 210 million years ago. The various species of drepanosaurid were characterized by specialized grasping limbs and often prehensile tails, adaptions for arboreal (tree-dwelling) and fossorial (digging) lifestyles, with some having also been suggested to be aquatic. Fossils of drepanosaurs have been found in Arizona, New Mexico, New Jersey, Utah, England, and northern Italy. The name is taken from the family's namesake genus Drepanosaurus, which means "sickle lizard," a reference to their strongly curved claws.

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

Macrocnemus is an extinct genus of archosauromorph reptile known from the Middle Triassic of Europe and China. Macrocnemus is a member of the Tanystropheidae family and includes three species. Macrocnemus bassanii, the first species to be named and described, is known from the Besano Formation and adjacent paleontological sites in the Italian and Swiss Alps. Macrocnemus fuyuanensis, on the other hand, is known from the Zhuganpo Formation in southern China. A third species, Macrocnemus obristi, is known from the Prosanto Formation of Switzerland and is characterized by gracile limbs. The name Macrocnemus is Greek for "long tibia".

Langobardisaurus is an extinct genus of tanystropheid archosauromorph reptile, with one valid species, L. pandolfii. Its fossils have been found in Italy and Austria, and it lived during the Late Triassic period, roughly 228 to 201 million years ago. Langobardisaurus was initially described in 1994, based on fossils from the Calcare di Zorzino Formation in Northern Italy. Fossils of the genus are also known from the Forni Dolostone of Northern Italy and the Seefeld Formation of Austria.

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

Cosesaurus is a genus of archosauromorph reptiles likely belonging to the family Tanystropheidae. It is known from fossil imprints of a single small skeleton, MGB V1, which was found in Muschelkalk outcrops near the municipalities of Mont-ral and Alcover in Spain. These outcrops are dated to the Ladinian age of the middle Triassic about 242 to 237 million years ago. The specimen is stored at the Museu Martorell, which is now part of the Museu de Ciències Naturals de Barcelona. The poor preservation and likely juvenile nature of the specimen has led to the anatomy of Cosesaurus being misidentified by several different sources. For example, Paul Ellenberger claimed that it was an ancestor to birds in the 1970s, while David Peters claimed that it was a pterosaur ancestor in 2000. Both of these claims contrast with mainstream scientific theories on the origins of either group, and other paleontologists who study the specimen are unable to find the features which Ellenberger or Peters reported to be present. The Ellenberger and Peters hypotheses are thus considered fringe theories with questionable scientific soundness due to their low reproducibility. Mainstream hypotheses for the relations of Cosesaurus generally agree that it is a "protorosaur", specifically a tanystropheid closely related to long-necked reptiles such as Macrocnemus, Tanytrachelos, Tanystropheus, or Langobardisaurus.

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

Jesairosaurus is an extinct genus of early archosauromorph reptile known from the Illizi Province of Algeria. It is known from a single species, Jesairosaurus lehmani. Although a potential relative of the long-necked tanystropheids, this lightly-built reptile could instead be characterized by its relatively short neck as well as various skull features.

<i>Prolacerta</i> Extinct genus of reptile from the lower Triassic

Prolacerta is a genus of archosauromorph from the lower Triassic of South Africa and Antarctica. The only known species is Prolacerta broomi. The generic name Prolacerta is derived from Latin meaning “before lizard” and its species name broomi is in commemoration of the famous paleontologist Robert Broom, who discovered and studied many of the fossils found in rocks of the Karoo Supergroup. When first discovered, Prolacerta was considered to be ancestral to modern lizards, scientifically known as lacertilians. However, a study by Gow (1975) instead found that it shared more similarities with the lineage that would lead to archosaurs such as crocodilians and dinosaurs. Prolacerta is considered by modern paleontologists to be among the closest relatives of the Archosauriformes.

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

Pamelaria is an extinct genus of allokotosaurian archosauromorph reptile known from a single species, Pamelaria dolichotrachela, from the Middle Triassic of India. Pamelaria has sprawling legs, a long neck, and a pointed skull with nostrils positioned at the very tip of the snout. Among early archosauromorphs, Pamelaria is most similar to Prolacerta from the Early Triassic of South Africa and Antarctica. Both have been placed in the family Prolacertidae. Pamelaria, Prolacerta, and various other Permo-Triassic reptiles such as Protorosaurus and Tanystropheus have often been placed in a group of archosauromorphs called Protorosauria, which was regarded as one of the most basal group of archosauromorphs. However, more recent phylogenetic analyses indicate that Pamelaria and Prolacerta are more closely related to Archosauriformes than are Protorosaurus, Tanystropheus, and other protorosaurs, making Protorosauria a polyphyletic grouping.

<span class="mw-page-title-main">Tanystropheidae</span> Extinct family of reptiles

Tanystropheidae is an extinct family of archosauromorph reptiles that lived throughout the Triassic Period, often considered to be "protorosaurs". They are characterized by their long, stiff necks formed from elongated cervical vertebrae with very long cervical ribs. Members of the group include both terrestrial and aquatic forms. While some tanystropheids were small lizard-like animals, other tanystropheids such as Tanystropheus were large animals that had necks that were several meters long, longer than the rest of their bodies.

Fuyuansaurus is an extinct genus of "protorosaur" reptiles known from the Middle Triassic Zhuganpo Formation of southern China. Fuyuansaurus was first named by Nicholas C. Fraser, Olivier Rieppel and Li Chun in 2013 and the type species is Fuyuansaurus acutirostris.

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

Eorasaurus is an extinct genus of archosauromorph reptile known from the middle Late Permian of Tatarstan, European Russia. It contains a single species, Eorasaurus olsoni. When originally described by Sennikov (1997), Eorasaurus was identified as an early archosauromorph and assigned to the family Protorosauridae, Ezcurra et al. (2014) and Ezcurra (2016) later reclassified Eorasaurus and placed it within the group Archosauriformes. Eorasaurus is based solely on scant fossil material from the neck region, and is thus considered an unstable taxon in phylogenetic analyses. If Eorasaurus is an archosauriform, it would be the oldest known member of the group and would pre-date the previous record holder.

<span class="mw-page-title-main">Prolacertidae</span> Extinct family of reptiles

Prolacertidae is an extinct family of archosauromorph reptiles that lived during the Early Triassic epoch. It was named in 1935 by the British palaeontologist Francis Rex Parrington to include the species Prolacerta broomi of South Africa and Antarctica. In 1979 a second species, Kadimakara australiensis, was described from Australia. Several other genera, such as Macrocnemus, Pamelaria and Prolacertoides, have also been assigned to this family in the past, but these have been placed elsewhere by later studies, leaving Prolacerta and Kadimakara as the only well-supported members.

<i>Ozimek volans</i> Extinct species of reptile

Ozimek is a genus of sharovipterygid archosauromorph reptile, known from Late Triassic deposits in Poland and closely related to the Kyrgyzstani Sharovipteryx. It contains one species, O. volans, named in 2016 by Jerzy Dzik and Tomasz Sulej. Like Sharovipteryx, Ozimek had long, slender limbs with the hindlimbs longer than the forelimbs; the hindlimbs likely supported gliding membranes as fossilized in Sharovipteryx. Another unusual characteristic was the shoulder girdle, where the massive coracoids formed a shield-like structure covering the bottom of the shoulder region that would have limited mobility. In other respects, such as its long neck, it was a typical member of the non-natural grouping Protorosauria. Phylogenetic analysis has indicated that it, possibly along with Sharovipteryx, may have been an unusual member of the protorosaur group Tanystropheidae, although further study of its anatomy is needed to resolve its precise relationships.

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

Pectodens is an extinct genus of archosauromorph reptile which lived during the Middle Triassic in China. The type and only species of the genus is P. zhenyuensis, named by Chun Li and colleagues in 2017. It was a member of the Archosauromorpha, specifically part of the unnatural grouping Protorosauria. However, an unusual combination of traits similar and dissimilar to other protorosaurs initially led to confusion over its evolutionary relationships. In 2021, it was placed in a newly-established group, Dinocephalosauridae, along with its closest relative Dinocephalosaurus.

<i>Kadimakara australiensis</i> Extinct species of reptile

Kadimakara is an extinct genus of early archosauromorph reptile from the Arcadia Formation of Queensland, Australia. It was seemingly a very close relative of Prolacerta, a carnivorous reptile which possessed a moderately long neck. The generic name Kadimakara references prehistoric creatures from Aboriginal myths which may have been inspired by ice-age megafauna. The specific name K. australiensis relates to the fact that it was found in Australia. Prolacerta and Kadimakara were closely related to the Archosauriformes, a successful group which includes archosaurs such as crocodilians, pterosaurs, and dinosaurs.

Boreopricea is an extinct genus of archosauromorph reptile from the Early Triassic of arctic Russia. It is known from a fairly complete skeleton discovered in a borehole on Kolguyev Island, though damage to the specimen and loss of certain bones has complicated study of the genus. Boreopricea shared many similarities with various other archosauromorphs, making its classification controversial. Various studies have considered it a close relative of Prolacerta, tanystropheids, both, or neither. Boreopricea is unique among early archosauromorphs due to possessing contact between the jugal and squamosal bones at the rear half of the skull.

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

Trachelosauridae is an extinct clade of archosauromorph reptiles that lived throughout the Triassic period. Like their close relatives the tanystropheids, they were "protorosaur"-grade archosauromorphs characterized by their long necks. Unlike tanystropheids, which lengthen their neck primarily by elongating the individual cervical (neck) vertebrae, trachelosaurids achieved their long necks by the addition of more vertebrae. The most extreme example of this trend was Dinocephalosaurus, which had at least 32 vertebrae in the neck alone, far more than the 13 neck vertebrae of Tanystropheus.

References

  1. 1 2 Sues, H.-D.; Fraser, N.C. (2010). "Early and early Middle Triassic in Gondwana". Triassic Life on Land: The Great Transition. New York: Columbia University Press. ISBN   9780231135221.
  2. 1 2 3 Spiekman, Stephan N. F.; Fraser, Nicholas C.; Scheyer, Torsten M. (2021-05-03). "A new phylogenetic hypothesis of Tanystropheidae (Diapsida, Archosauromorpha) and other "protorosaurs", and its implications for the early evolution of stem archosaurs". PeerJ. 9: e11143. doi: 10.7717/peerj.11143 . ISSN   2167-8359. PMC   8101476 . PMID   33986981.
  3. ↑ Simões, T. R.; Kammerer, C. F.; Caldwell, M. W.; Pierce, S. E. (2022). "Successive climate crises in the deep past drove the early evolution and radiation of reptiles". Science Advances. 8 (33): eabq1898. doi: 10.1126/sciadv.abq1898 . PMC   9390993 . PMID   35984885.
  4. 1 2 Spiekman, Stephan N. F.; Ezcurra, Martín D.; Rytel, Adam; Wang, Wei; Mujal, Eudald; Buchwitz, Michael; Schoch, Rainer R. (2024-03-15). "A redescription of Trachelosaurus fischeri from the Buntsandstein (Middle Triassic) of Bernburg, Germany: the first European Dinocephalosaurus-like marine reptile and its systematic implications for long-necked early archosauromorphs". Swiss Journal of Palaeontology . 143 (1): 10. doi: 10.1186/s13358-024-00309-6 . ISSN   1664-2384.
  5. ↑ Benton, M.J.; Allen, J.L. (1997). "Boreopricea from the Lower Triassic of Russia, and the relationships of the prolacertiform reptiles" (PDF). Palaeontology. 40 (4): 931–953.
  6. ↑ Gottmann-Quesada, A.; Sander, P.M. (2009). "A redescription of the early archosauromorph Protorosaurus speneri Meyer, 1832, and its phylogenetic relationships". Palaeontographica Abteilung A. 287 (4–6): 123–200. doi:10.1127/pala/287/2009/123.
  7. 1 2 3 Modesto, S. P.; Sues, H. D. (2004). "The skull of the Early Triassic archosauromorph reptile Prolacerta broomi and its phylogenetic significance". Zoological Journal of the Linnean Society. 140 (3): 335–351. doi: 10.1111/j.1096-3642.2003.00102.x .
  8. ↑ Dilkes, David M. (1998). "The Early Triassic rhynchosaur Mesosuchus browni and the interrelationships of basal archosauromorph reptiles". Philosophical Transactions of the Royal Society of London, Series B. 353 (1368): 501–541. doi:10.1098/rstb.1998.0225. PMC   1692244 .
  9. ↑ Sues, H.-D. (2003). "An unusual new archosauromorph reptile from the Upper Triassic Wolfville Formation of Nova Scotia". Canadian Journal of Earth Sciences. 40 (4): 635–649. Bibcode:2003CaJES..40..635S. doi:10.1139/e02-048.
  10. ↑ Rieppel, O.; Li, C.; Fraser, N. C. (2008). "The skeletal anatomy of the triassic protorosaur Dinocephalosaurus orientalis Li, from the Middle Triassic of Guizhou Province, southern China". Journal of Vertebrate Paleontology. 28: 95–110. doi:10.1671/0272-4634(2008)28[95:TSAOTT]2.0.CO;2. S2CID   86026836.
  11. ↑ Renesto, Silvio; Spielmann, Justin A.; Lucas, Spencer G.; Tarditi Spagnoli, Giorgio (2010). "The taxonomy and paleobiology of the Late Triassic (Carnian-Norian: Adamanian-Apachean) drepanosaurs (Diapsida: Archosauromorpha: Drepanosauromorpha)". New Mexico Museum of Natural History and Science Bulletin. 46: 1–81.
  12. ↑ Li, C.; Rieppel, O.; Labarbera, M. C. (2004). "A Triassic Aquatic Protorosaur with an Extremely Long Neck". Science. 305 (5692): 1931. doi:10.1126/science.1100498. PMID   15448262. S2CID   38739295.
  13. ↑ Borsuk−Białynicka, Magdalena; Evans, Susan E. (2009). "A long−necked archosauromorph from the Early Triassic of Poland" (PDF). Paleontologica Polonica. 65: 203–234.
  14. ↑ Hone, D. W. E.; Benton, M. J. (2007). "An evaluation of the phylogenetic relationships of the pterosaurs among archosauromorph reptiles". Journal of Systematic Palaeontology. 5 (4): 465–469. doi:10.1017/S1477201907002064. S2CID   86145645.
  15. ↑ Pritchard, Adam C.; Turner, Alan H.; Nesbitt, Sterling J.; Irmis, Randall B.; Smith, Nathan D. (2015-03-04). "Late Triassic tanystropheids (Reptilia, Archosauromorpha) from northern New Mexico (Petrified Forest Member, Chinle Formation) and the biogeography, functional morphology, and evolution of Tanystropheidae". Journal of Vertebrate Paleontology. 35 (2): e911186. doi:10.1080/02724634.2014.911186. ISSN   0272-4634. S2CID   130089407.
  16. ↑ J., Nesbitt, Sterling; Flynn, John J.; Pritchard, Adam C.; J. Michael, Parrish; Lovasoa, Ranivoharimanana; Wyss, André R. (2015-12-07). "Postcranial osteology of Azendohsaurus madagaskarensis (?Middle to Upper Triassic, Isalo Group, Madagascar) and its systematic position among stem archosaur reptiles. (Bulletin of the American Museum of Natural History, no. 398)". hdl:2246/6624.{{cite journal}}: Cite journal requires |journal= (help)CS1 maint: multiple names: authors list (link)
  17. ↑ Ezcurra, Martín D. (2016-04-28). "The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms". PeerJ. 4: e1778. doi: 10.7717/peerj.1778 . ISSN   2167-8359. PMC   4860341 . PMID   27162705.
  18. ↑ Borsuk–Białynicka, M.; Evans, S.E. (2009). "A long–necked archosauromorph from the Early Triassic of Poland" (PDF). Palaeontologia Polonica. 65: 203–234.
  19. ↑ Renesto, S (1994). "Megalancosaurus, a possibly arboreal archosauromorph (Reptilia) from the Upper Triassic of northern Italy". Journal of Vertebrate Paleontology. 14 (1): 38–52. doi:10.1080/02724634.1994.10011537.
  20. ↑ Peters, D (2000). "A Redescription of Four Prolacertiform Genera and Implications for Pterosaur Phylogenesis". Rivista Italiana di Paleontologia e Stratigrafia. 106 (3): 293–336.
  21. ↑ Senter, P (2004). "Phylogeny of Drepanosauridae (Reptilia: Diapsida)". Journal of Systematic Palaeontology. 2 (3): 257–268. doi:10.1017/S1477201904001427. S2CID   83840423.
This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.