Dendromaia

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Dendromaia
Temporal range: Late Moscovian, 309–306  Ma
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Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Family: Varanopidae
Genus: Dendromaia
Maddin, Mann, & Hebert, 2019
Type species
Dendromaia unamakiensis
Maddin, Mann, & Hebert, 2019

Dendromaia is an extinct genus of varanopid from the Carboniferous of Nova Scotia. It contains a single species, Dendromaia unamakiensis. [1] Dendromaia is the oldest known varanopid, likely the oldest known synapsid, and the only member of the family Varanopidae to be discovered in Nova Scotia. Known from a large partial skeleton preserved with its tail wrapped around a much smaller partial skeleton, Dendromaia may also represent the oldest known occurrence of parental care in the fossil record. While the larger skeleton possessed certain mycterosaurine-like features, the smaller skeleton resembled basal varanopids such as Archaeovenator and Pyozia , creating uncertainty over whether characteristics at the base of Varanopidae have legitimate phylogenetic significance or instead reflect the immaturity of basal varanopid specimens. [1]

Contents

Discovery

Dendromaia unamakiensis is known from a slab and counterslab containing two skeletons. The specimen, NSM017GF020.001, was discovered in a petrified lycopod stump at Point Aconi on Cape Breton Island in Nova Scotia. This site is part of the Sydney Mines Formation, which is dated to the late Moscovian stage of the Carboniferous period, 309-306 million years ago. [2] "Dendromaia" roughly translates to "the mother in the tree", [3] according to its discovery in a stump and proposed parental care. The specific name references Unama'kik, the Mi'kmaq name for Cape Breton Island. The genus and species were described by Hillary Maddin, Arjan Mann, and Brian Hebert in 2019. [1] Hebert had discovered the specimen in 2017. [3] [4]

The two skeletons had different sizes and preserved different areas, with the larger skeleton (designated the holotype) incorporating a large portion of the rear half of the body and the smaller skeleton (designated the paratype) including a skull and only fragments of the postcranium, 1/4th the size of the larger one. These skeletons were inferred to represent two individuals of a single new taxon based on their similar overlapping anatomy, apparent varanopid ancestry, close association, and the fact that there are no other varanopids known from Nova Scotia. [1]

Description

The 6 preserved dorsal vertebrae of the larger skeleton were among the most characteristic bones in the animal. They shared several traits with Mycterosaurus , such as tall rectangular neural spines with slight depressions at their base. However, the rib facets at the tip of the transverse processes were not vertical, but instead oriented diagonally. Like other non-varanodontine varanopids, the lower edge of each centrum had a rounded keel. 10 incomplete caudal vertebrae preserved on the counterpart were elongated and had low neural spines. Holocephalous (single-headed) ribs and Heleosaurus -like gastralia were also present, though osteoderms were seemingly absent. The plate-like bones composing the pelvis were not fused to each other, and Dendromaia had a large pubic foramen like that of Heleosaurus. The femur was lightly-built and twisted, similar to mycterosaurines. The rest of the leg and foot was present but incomplete. [1] The larger specimen may have been 20 to 30 centimeters (7.9-12 inches) long from the tip of the snout to the base of the tail, and its full tail length is unknown but likely elongated. [4] [5]

The small skeleton's poorly-preserved skull was triangular and pointed like those of Archaeovenator , Pyozia , and Heleosaurus. It had thin, curved teeth on the maxilla (without a canine region) and smaller teeth on the palate, which also possessed a varanopid-like pterygoid and cultriform process. Isolated hyoid bones were also identifiable. Like the larger skeleton, the small skeleton's vertebrae had a rounded keel on their underside. It also possessed several limb bone fragments, including a twisted humerus. [1]

Classification

Dendromaia is the oldest member of a family of amniotes known as varanopids. Varanopids are traditionally considered early synapsids (part of the lineage of amniotes leading to mammals), although some studies alternatively propose that they are diapsid reptiles related to Orovenator . [6] [7] The describers of Dendromaia prefer a position for varanopids among synapsids, citing both biological characteristics (parental care) and arguing that traits which link varanopids to diapsids may be based on oversampling of juvenile specimens at the base of Varanopidae. [1]

To investigate the position of Dendromaia among other varanopids, the describers adapted a phylogenetic analysis matrix previously utilized by Brocklehurst & Fröbisch (2018). [8] The resulting strict consensus tree (average result of most parsimonious trees) of the parsimony analysis placed Dendromaia in a polytomy near the base of Varanopidae, along with Pyozia and a clade which forks into Varanodontinae and Mycterosaurinae. The Bayesian analysis recovers a similar result, albeit with the polytomy resolved, placing Dendromaia as the sister taxon to Pyozia. [1] The following cladogram is based on the results of the Bayesian analysis in Maddin, Mann, & Hebert (2019): [1]

Varanopidae

Archaeovenator hamiltonensis ArchaeovenatorDB.jpg

Apsisaurus witteri

Dendromaia unamakiensis

Pyozia mesenensis

Mycterosaurinae

Mycterosaurus longiceps

Mesenosaurus romeri

Heleosaurus scholtzi

Elliotsmithia longiceps ? (BP/1/5678)

Varanodontinae

Varanops brevirostris Varanops brevirostris2DB.jpg

Watongia meieri

Varanodon agilis Varanodon1DB.jpg

Aerosaurus wellesi Aerosaurus wellesi.jpg

Aerosaurus greenleorum

Ruthiromia elcobriensis

Paleobiology

The delicate preservation of the skeletons indicated that they likely died and were quickly buried at the same place and time. They were positioned with the smaller skeleton encircled by the tail of the larger skeleton. These taphonomic qualities led the paleontologists who described them to propose that the two skeletons were denning together under the roots of a lycopod tree, with the smaller skeleton likely representing the offspring of the larger skeleton. This may be the oldest fossil evidence of parental care, predating the previously oldest evidence, an aggregation of varanopids from the Permian of South Africa which may represent their own species ( Microvaranops parentis ) [9] or specimens of Heleosaurus scholtzi . [1] On the other hand, it remains a possibility that the two skeletons were not close relatives, and instead simply sheltered from a storm in the same stump. [3] The anatomy of the juvenile Dendromaia skeleton is remarkably similar to that of basal varanopids such as Archaeovenator and Pyozia , indicating that these genera may be based on juvenile specimens of larger varanopids. Likewise, certain characteristics (such as reduced dentition and limb development) found to link basal varanopids with diapsids may be a consequence of juvenile specimen sampling, rather than valid phylogenetic signals. [1]

Related Research Articles

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

Synapsids are one of the two major clades of vertebrate animals in the group Amniota, the other being the sauropsids, which include reptiles and birds. The synapsids were once the dominant land animals in the late Paleozoic and early Mesozoic, but the only extant group that survived into the Cenozoic are the 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">Pelycosaur</span> Informal grouping composed of basal or primitive Late Paleozoic synapsids

Pelycosaur is an older term for basal or primitive Late Paleozoic synapsids, excluding the therapsids and their descendants. Previously, the term mammal-like reptile had been used, and pelycosaur was considered an order, but this is now thought to be incorrect, and seen as outdated.

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

Hylonomus is an extinct genus of reptile that lived 312 million years ago during the Late Carboniferous period. It is the earliest unquestionable reptile. The only species is the type species Hylonomus lyelli. Despite being amongst the oldest known reptiles, it is not the most primitive member of group, being a eureptile more derived than either parareptiles or captorhinids.

<span class="mw-page-title-main">Eupelycosauria</span> Clade of synapsids

Eupelycosauria is a large clade of animals characterized by the unique shape of their skull, encompassing all mammals and their closest extinct relatives. They first appeared 308 million years ago during the Early Pennsylvanian epoch, with the fossils of Echinerpeton and perhaps an even earlier genus, Protoclepsydrops, representing just one of the many stages in the evolution of mammals, in contrast to their earlier amniote ancestors.

<span class="mw-page-title-main">Caseasauria</span> Extinct clade of synapsids

Caseasauria is one of the two main clades of early synapsids, the other being the Eupelycosauria. Caseasaurs are currently known only from the Late Carboniferous and the Permian, and include two superficially different families, the small insectivorous or carnivorous Eothyrididae, and the large, herbivorous Caseidae. These two groups share a number of specialised features associated with the morphology of the snout and external naris.

Varanopidae is an extinct family of amniotes that resembled monitor lizards and may have filled a similar niche, hence the name. Typically, they are considered synapsids that evolved from an Archaeothyris-like synapsid in the Late Carboniferous. However, some recent studies have recovered them being taxonomically closer to diapsid reptiles. A varanopid from the latest Middle Permian Pristerognathus Assemblage Zone is the youngest known varanopid and the last member of the "pelycosaur" group of synapsids.

<i>Ophiacodon</i> Extinct genus of synapsids

Ophiacodon is an extinct genus of synapsid belonging to the family Ophiacodontidae that lived from the Late Carboniferous to the Early Permian in North America and Europe. The genus was named along with its type species O. mirus by paleontologist Othniel Charles Marsh in 1878 and currently includes five other species. As an ophiacodontid, Ophiacodon is one of the most basal synapsids and is close to the evolutionary line leading to mammals.

<i>Mycterosaurus</i> Extinct genus of tetrapods

Mycterosaurus is an extinct genus of synapsids belonging to the family Varanopidae. It is classified in the varanopid subfamily Mycterosaurinae. Mycterosaurus is the most primitive member of its family, existing from 290.1 to 272.5 MYA, known to Texas and Oklahoma. It lacks some features that its advanced relatives have.

<i>Archaeovenator</i> Extinct genus of tetrapods

Archaeovenator is an extinct genus of Late Carboniferous varanopid synapsids known from Greenwood County, Kansas of the United States. It was first named by Robert R. Reisz and David W. Dilkes in 2003 and the type species is Archaeovenator hamiltonensis. Archaeovenator hamiltonensis is known from the holotype KUVP 12483, a three-dimensionally preserved, nearly complete and articulated skeleton, including the skull, with limbs and girdles slightly separated from postcranial skeleton. It was collected in the Hamilton Quarry, from the Calhouns Shale Formation of the Shawnee Group, dating to the Virgilian stage of the Late Pennsylvanian Series, about 300 million years ago. The generic name is derived from the Latin Archaeo and venator, meaning "ancient hunter". The specific name is named after its finding place Hamilton Quarry. Archaeovenator is the oldest and the basalmost known varanopid, as it is the sister taxon to all other known varanopsids.

<i>Echinerpeton</i> Extinct genus of synapsids

Echinerpeton is an extinct genus of synapsid, including the single species Echinerpeton intermedium from the Late Carboniferous of Nova Scotia, Canada. The name means 'spiny lizard' (Greek). Along with its contemporary Archaeothyris, Echinerpeton is the oldest known synapsid, having lived around 308 million years ago. It is known from six small, fragmentary fossils, which were found in an outcrop of the Morien Group near the town of Florence. The most complete specimen preserves articulated vertebrae with high neural spines, indicating that Echinerpeton was a sail-backed synapsid like the better known Dimetrodon, Sphenacodon, and Edaphosaurus. However, the relationship of Echinerpeton to these other forms is unclear, and its phylogenetic placement among basal synapsids remains uncertain.

<i>Mesenosaurus</i> Extinct genus of synapsids

Mesenosaurus is an extinct genus of amniote. It belongs to the family Varanopidae. This genus includes two species: the type species Mesenosaurus romeri from the middle Permian Mezen River Basin of northern Russia, and Mesenosaurus efremovi from the early Permian (Artinskian) Richards Spur locality. M. romeri’s stratigraphic range is the middle to late Guadalupian while M. efremovi’s stratigraphic range is the Cisuralian.

<i>Ianthodon</i> Extinct genus of synapsids

Ianthodon is an extinct genus of basal haptodontiform synapsids from the Late Carboniferous about 304 million years ago. The taxon was discovered and named by Kissel & Reisz in 2004. The only species in the taxon, Ianthodon schultzei, was found by separating it from a block that also contained the remains of Petrolacosaurus and Haptodus. The evolutionary significance of the taxon wasn't realized until a publication in 2015. The fossil of this organism was discovered in Garnett, Kansas.

<i>Heleosaurus</i> Extinct genus of tetrapods

Heleosaurus scholtzi is an extinct species of basal synapsids, known as pelycosaurs, in the family of Varanopidae during the middle Permian. At first H. scholtzi was mistakenly classified as a diapsid. Members of this family were carnivorous and had dermal armor, and somewhat resembled monitor lizards. This family was the most geologically long lived, widespread, and diverse group of early amniotes. To date only two fossils have been found in the rocks of South Africa. One of these fossils is an aggregation of five individuals.

<span class="mw-page-title-main">Tuditanidae</span> Extinct family of amphibians

Tuditanidae is an extinct family of tuditanomorph microsaurs. Fossils have been found from Nova Scotia, Ohio, and the Czech Republic and are Late Carboniferous in age.

<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.

<i>Orovenator</i>

Orovenator is an extinct genus of diapsid from Lower Permian deposits of Oklahoma, United States. It is known from two partial skulls from the Richards Spur locality in Oklahoma. The holotype OMNH 74606 consists of a partial skull preserving snout and mandible, and the referred specimen, OMNH 74607, a partial skull preserving the skull roof, vertebrae and palatal elements. It was first named by Robert R. Reisz, Sean P. Modesto and Diane M. Scott in 2011 and the type species is Orovenator mayorum. The generic name means "mountain", oro, in Greek in reference to the Richards Spur locality, which was mountainous during the Permian period and "hunter", venator, in Latin. The specific name honours Bill and Julie May. Orovenator is the oldest and most basal neodiapsid to date.

Ascendonanus is an extinct genus of varanopid amniote from the Early Permian of Germany. It is one of the earliest specialized arboreal (tree-living) tetrapods currently known and outwardly resembled a small lizard. The animal was about 40 cm long, with strongly curved claws, short limbs, a slender, elongated trunk, and a long tail. It would have preyed on insects and other small arthropods.

<i>Cabarzia</i> Extinct genus of lizard-like animals

Cabarzia is an extinct genus of varanopid from the Early Permian of Germany. It contains only a single species, Cabarzia trostheidei, which is based on a well-preserved skeleton found in red beds of the Goldlauter Formation. Cabarzia shared many similarities with Mesenosaurus romeri, although it did retain some differences, such as more curved claws, a wide ulnare, and muscle scars on its sacral ribs. With long, slender hindlimbs, a narrow body, an elongated tail, and short, thick forelimbs, Cabarzia was likely capable of running bipedally to escape from predators, a behavior shared by some modern lizards. It is the oldest animal known to have adaptations for bipedal locomotion, predating Eudibamus, a bipedal bolosaurid parareptile from the slightly younger Tambach Formation.

Hillary Catherine Maddin is a Canadian paleontologist and developmental biologist known for her work on development in extinct and extant amphibians. She is currently an associate professor in the Department of Earth Sciences at Carleton University.

References

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  2. Allen, Jonathan P.; Fielding, Christopher R.; Gibling, Martin R.; Rygel, Michael C. (2013). "Recognizing products of palaeoclimate fluctuation in the fluvial stratigraphic record: An example from the Pennsylvanian to Lower Permian of Cape Breton Island, Nova Scotia". Sedimentology. 61 (5): 1332–1381. doi:10.1111/j.1365-3091.2013.12102.x. ISSN   1365-3091.
  3. 1 2 3 Wu, Katherine J. (23 December 2019). "Lizard-Like Fossil May Represent 306-Million-Year-Old Evidence of Animal Parenting". Smithsonian Magazine.
  4. 1 2 Davis, Nicola (2019-12-23). "300m-year-old fossil is early sign of creatures caring for their young". The Guardian. ISSN   0261-3077.
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  8. Brocklehurst, Neil; Fröbisch, Jörg (2018-09-03). "A reexamination of Milosaurus mccordi, and the evolution of large body size in Carboniferous synapsids". Journal of Vertebrate Paleontology. 38 (5): e1508026. doi:10.1080/02724634.2018.1508026. ISSN   0272-4634. S2CID   91487577.
  9. Spindler, Frederik; Werneburg, Ralf; Schneider, Joerg W.; Luthardt, Ludwig; Annacker, Volker; Rößler, Ronny (2018-06-01). "First arboreal 'pelycosaurs' (Synapsida: Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny". PalZ. 92 (2): 315–364. doi:10.1007/s12542-018-0405-9. ISSN   1867-6812. S2CID   133846070.