Trematopidae

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Trematopidae
Temporal range: 305.9–272.5  Ma
Acheloma cumminsi.jpg
Skeleton of Acheloma cumminsi in the Field Museum of Natural History.
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order:† Temnospondyli
Clade:† Olsoniformes
Family:† Trematopidae
Williston, 1910
Genera

Acheloma
Actiobates
Anconastes
Ecolsonia
Fedexia
Phonerpeton
Rotaryus
Tambachia
Mordex
Mattauschia

Contents

Trematopidae is a family of dissorophoid temnospondyls spanning the late Carboniferous to the early Permian. Together with Dissorophidae, the family forms Olsoniformes, a clade comprising the medium-large terrestrial dissorophoids. Trematopids are known from numerous localities in North America, primarily in New Mexico, Oklahoma, and Texas, and from the Bromacker quarry in Germany.

History of study

The clade Trematopidae was first proposed by American paleontologist S.W. Williston in 1910, although it was named as "Trematopsidae" following the historical (but inaccurate) derivation from the genus "Trematops" (now synonymized with Acheloma). [1] British paleontologist D.M.S. Watson proposed a related clade in 1919, Achelomidae, for Acheloma, based on perceived differences separating the taxa; [2] this is now considered a junior synonym of Trematopidae following guidelines of historical precedent.

19th century history

In 1882, American paleontologist Edward Drinker Cope named Acheloma cumminsi based on material collected from the early Permian of Texas. [3] This is technically the first trematopid to be named, although the holotype of Mordex calliprepes was named a year earlier as a species of Limnerpeton by Czech paleontologist Antonin Fritsch in 1881 (one year prior); [4] the status of Mordex as a trematopid remained debated however until the revision by Milner (2018). [5]

20th century history

In the first half of the 20th century, American paleontologist S.W. Williston named a new genus from the early Permian of Texas, Trematops. [6] At the time, most workers regarded it as only distantly related to Cope's Acheloma. [7] Two additional species of Trematops were named in short order, Trematops thomasi from Oklahoma, named by American paleontologist Maurice Mehl, [8] and Trematops willistoni from Texas, named by American paleontologist E.C. Olson in 1941. [9] Both are now regarded as junior synonyms of Acheloma cumminsi. Olson also named two species of Acheloma, A. whitei and A. pricei; both are now regarded as belonging to Phonerpeton. [10] In the same paper that he named these taxa, Olson also provided the first review of the Trematopidae, synthesizing all of the known material and providing updated taxonomic frameworks.

The second half of the 20th century saw an increase in trematopid research. In 1956, Olson named a new genus and species of trematopid from the Vale Formation of Texas, Trematopsis seltini; [11] this is now regarded as a junior synonym of the dissorophid Cacops aspidephorus . [12] In 1969, American paleontologist Peter Vaughn described the first trematopid from New Mexico, Ecolsonia cutlerensis, named for the contemporaneous Olson and the Cutler Formation from which the holotype was collected. A re-description of this taxon based on substantial new material, was completed in 1985 by a team led by American paleontologist David Berman from the Carnegie Museum of Natural History. [13] In 1970, Olson named a new species of Trematops from the early Permian of Ohio, "Trematops stonei"; [14] this is the only occurrence of Acheloma/Trematops outside of Texas and Oklahoma but is now accepted to be a junior synonym of Acheloma cumminsi. In 1973, American paleontologist Theodore Eaton named Actiobates peabodyi, the first and only trematopid from Kansas and the oldest trematopid in North America. [15] Eaton also challenged the longstanding separation of trematopids and dissorophids, synonymizing them under Dissorophidae, but this has not been supported or maintained by subsequent workers. Additional material from the Garnett quarry where Actiobates was discovered, assigned to "Hesperoherpeton garnettense" may also belong to Actiobates. [16] In 1974, American paleontologist John Bolt published two papers, one describing the first trematopid material from the fossil-rich site near Richards Spur, Oklahoma, [17] and the a second exploring the function of the elongate naris in trematopids. [18] In 1985, Olson described a purported larval specimen of a trematopid; [19] this was subsequently challenged by Canadian paleontologist David Dilkes in 1991, [20] and the specimen is now regarded to be an adult specimen of an amphibamiform dissorophoid. [21] [22] In 1987, the second trematopid from New Mexico, Anconastes vesperus, named for its discovery in El Cobre Canyon in western North America, was described by the same team of Berman, Reisz, and Eberth who described Ecolsonia. [23] Also in 1987, Dilkes and Robert R. Reisz of the University of Toronto re-described the holotypes of Acheloma cumminsi and Trematops milleri and identified them as synonyms; A. cumminsi takes precedence, having been described first. In 1990, Dilkes named a new taxon of trematopid from the early Permian of Texas, Phonerpeton pricei . [10] A study on developmental changes to the characteristic elongate naris of trematopids was undertaken using primarily material from Phonerpeton by Dilkes in 1993. [24] In 1998, the first trematopid from Europe, Tambachia trogallas , from the early Permian Bromacker quarry in Germany was named by a team led by American paleontologist Stuart Sumida of California State University, San Bernardino. [25]

21st century history

The 21st century has seen a renewed flurry of research into trematopids. In 2010, a team led by Berman described a new taxon from the late Carboniferous of Pennsylvania, Fedexia striegeli , named after the FedEx Corporation on whose land the holotype was found and after Adam Striegel, the discoverer. [26] In 2011, another team led by Berman described a new taxon from the Bromacker quarry, Rotaryus gothae, named after the Rotary Club of Gotha's contributions to the excavation of the locality. [27] Also in 2011, Canadian paleontologists Brendan Polley and Reisz named a new species of Acheloma from the early Permian Richards Spur locality in Oklahoma, Acheloma dunni, named for Brent Dunn, one of the collectors at the locality. [28] A study examining development changes in this species was published the previous year by Canadian paleontologists Hillary C. Maddin, Jason S. Anderson, and Reisz, [29] although the species name was not formalized by the time of that publication. In their 2014 review of the earlier temnospondyls, Schoch & Milner resurrected a second species of Phonerpeton, P. whitei, [21] following Olson's original species distinctions. [9] Data from the pes of Acheloma cumminsi was included in a broader survey of the carpus and tarsus in temnospondyls by Dilkes in 2015. [30] In 2018, British paleontologist Andrew R. Milner revised the Nýřany trematopids, clarifying the complicated history of Mordex calliprepes and erecting a new taxon, Mattauschia laticeps. [5] In 2019, a team led by American paleontologist Bryan M. Gee published the first description of computed tomography (CT) data of a trematopid in the form of a small specimen from the Richards Spur locality that they referred to cf. Acheloma. [31]

Anatomy

Trematopids have typically been identified by the presence of a noticeably enlarged naris that is often sub-divided in Permian forms such as Acheloma. They are also among the largest of the dissorophoids, with some specimens of Acheloma exceeding 18 cm in skull length and thus being rivaled only by middle Permian dissorophids such as Anakamacops . Schoch & Milner (2014) diagnosed trematopids by the following features: (1) greatly expanded naris replacing much of the lacrimal; (2) medially situated narial flange meeting antorbital bar; (3) otic notch with a ventral margin sloping at less than 45-degrees in large individuals; (4) a medial inflection of the rim of the adductor fossa; (5) a pterygoid-vomer contact; (6) a triangular patch of denticles on the basal plate of the parasphenoid; and (7) a humerus with a supinator process. [21] Milner (2018) further refined this based on his restudy of Mordex, including only characters 1, 3, and 5 from Schoch & Milner (2014), noting that some typical trematopid features are either not known or are not present in the primitive Mattauschia and Mordex. [5]

The function of the naris remains largely unresolved. Olson (1941) suggested that the anterior half served the typical function of smell detection and that the posterior half housed some kind of gland. [9] Bolt (1974) conjectured that the well-developed nasal flange was probably for distribution of stresses throughout the skull and might be a hallmark of terrestrial dissorophoids. [18] He then suggested that there might be a gland related to salt excretion (the "glandula nasalis externa") that would produce the enlargement of the naris. Dilkes (1993) did not discount this hypothesis, but suggested alternatives, namely that the nasal flange and the expanded naris may have been for another physiological function, such as improving respiratory efficiency and water retention, an important attribute for a terrestrial amphibian to have. [24] However, all of these hypotheses remain speculative in the absence of much complexity of the hard tissues surrounding the naris and given the relatively vague paleoecological information available for trematopids (e.g., diet).

Relationships

The placement of trematopids within Dissorophoidea has long been accepted, as has their close relationship to dissorophids, although Olsoniformes was not formalized until 2007. [32] Ecolsonia and Mordex have been more uncertainly placed in their history of study, [13] [5] but both are now accepted as unequivocal trematopids. The two most recent phylogenetic analyses of Trematopidae are those by Berman et al. (2011) and Polley & Reisz (2011):

Topology of Berman et al. (2011): [27]

  Amphibamidae  

Eoscopus

Amphibamus

Doleserpeton

Micropholis

Tersomius

  Olsoniformes  

Ecolsonia

  Dissorophidae  

Broiliellus

Dissorophus

Aspidosaurus

Cacops

 Trematopidae 

Fedexia

Anconastes

Tambachia

Rotaryus

Acheloma

Phonerpeton

Topology of Polley & Reisz (2011): [28]

Sclerocephalus haeuseri

Micromelerpeton credneri

Micropholis stowi

Eoscopus lockardi

  Olsoniformes  

Cacops morrisi

 Trematopidae 

Phonerpeton pricei

Acheloma cumminsi

Acheloma dunni

Actiobates peabodyi

Ecolsonia cutlerensis

Anconastes vesperus

Tambachia trogallas

Related Research Articles

<i>Cacops</i> Extinct genus of amphibians

Cacops, is a genus of dissorophid temnospondyls from the Kungurian stage of the early Permian of the United States. Cacops is one of the few olsoniforms whose ontogeny is known. Cacops fossils were almost exclusively known from the Cacops Bone Bed of the Lower Permian Arroyo Formation of Texas for much of the 20th century. New material collected from the Dolese Brothers Quarry, near Richards Spur, Oklahoma in the past few decades has been recovered, painting a clearer picture of what the animal looked and acted like.

<span class="mw-page-title-main">Dissorophidae</span> Extinct family of temnospondyls

Dissorophidae is an extinct family of medium-sized temnospondyls that flourished during the late Carboniferous and early Permian periods. The clade is known almost exclusively from North America.

<span class="mw-page-title-main">Dissorophoidea</span> Extinct superfamily of amphibians

Dissorophoidea is a clade of medium-sized, temnospondyl amphibians that appeared during the Moscovian in Euramerica, and continued through to the Late Permian and the Early Triassic of Gondwana. They are distinguished by various details of the skull, and many species seem to have been well adapted for life on land.

<span class="mw-page-title-main">Temnospondyli</span> Ancestors of modern amphibians adapted to life on land

Temnospondyli or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carboniferous, Permian and Triassic periods, with fossils being found on every continent. A few species continued into the Jurassic and Early Cretaceous periods, but all had gone extinct by the Late Cretaceous. During about 210 million years of evolutionary history, they adapted to a wide range of habitats, including freshwater, terrestrial, and even coastal marine environments. Their life history is well understood, with fossils known from the larval stage, metamorphosis and maturity. Most temnospondyls were semiaquatic, although some were almost fully terrestrial, returning to the water only to breed. These temnospondyls were some of the first vertebrates fully adapted to life on land. Although temnospondyls are amphibians, many had characteristics such as scales and armour-like bony plates that distinguish them from the modern soft-bodied lissamphibians.

<i>Acheloma</i> Extinct genus of amphibians

Acheloma is an extinct genus of temnospondyl that lived during the Early Permian. The type species is A. cumminsi.

<i>Anconastes</i> Extinct genus of amphibians

Anconastes is an extinct genus of dissorophoid temnospondyl within the family Trematopidae. It is known from two specimens from the Late Carboniferous Cutler Formation of north-central New Mexico in the southwestern United States. The genus name derives from two Greek roots, ankos and nastes ("inhabitant"), which refers to the type locality of El Cobre Canyon where the specimens were found. The specific name is derived from the Latin word vesperus ("western"). The more complete specimen, the holotype, is a partial skull with articulated mandibles and a substantial amount of the postcranial skeleton. The less complete specimen, the paratype, consists only of the right margin of the skull with an articulated mandible.

Actiobates is an extinct genus of trematopid temnospondyl that lived during the Late Carboniferous. It is known from the Garnett Quarry in Kansas.

<i>Broiliellus</i> Extinct genus of amphibians

Broiliellus is an extinct genus of dissorophoid temnospondyl within the family Dissorophidae. Broiliellus is most closely related to the genus Dissorophus, and both have been placed in the subfamily Dissorophinae. Broiliellus is known from five species from the Early Permian: the type species is Broiliellus texensis, and the other species are Broiliellus brevis,Broiliellus olsoni, Broiliellus arroyoensis, and Broiliellus reiszi. An additional species, Broiliellus novomexicanus, which was originally named Aspidosaurus novomexicanus, is now thought to fall outside the genus as a member of the subfamily Eucacopinae.

<i>Ecolsonia</i> Extinct genus of amphibians

Ecolsonia is an extinct genus of trematopid temnospondyl. Its phylogenetic position within Olsoniformes has been historically debated, but it is presently considered to be a trematopid.

<i>Dissorophus</i> Extinct genus of amphibians

Dissorophus (DI-soh-ROH-fus) is an extinct genus of temnospondyl amphibian that lived during the Early Permian Period about 273 million years ago. Its fossils have been found in Texas and in Oklahoma in North America. Its heavy armor and robust build indicate Dissorophus was active on land, similar to other members of the clade Dissorophidae that are known from the Late Carboniferous to the Early Permian periods. Dissorphus is distinguished by its small body size, disproportionately large head and short trunk.

Limnerpeton is an extinct genus of dissorophoidean euskelian temnospondyl within the family Amphibamidae.

<i>Phonerpeton</i> Extinct genus of amphibians

Phonerpeton is an extinct genus of dissorophoid temnospondyl within the family Trematopidae that is known from the early Permian of Texas.

Tersomius is an extinct genus of dissorophoid temnospondyl within the family Micropholidae. It is known from the early Permian of North America.

<i>Fedexia</i> Extinct genus of amphibians

Fedexia is an extinct genus of carnivorous temnospondyl within the family Trematopidae. It lived 300 million years ago during the late Carboniferous period. It is estimated to have been 2 feet (0.61 m) long, and likely resembled a salamander. Fedexia is known from a single skull found in Moon Township, Pennsylvania. It is named after the shipping service FedEx, which owned the land where the holotype specimen was first found.

<i>Tambachia</i> Extinct genus of amphibians

Tambachia is an extinct genus of dissorophoid temnospondyl within the family Trematopidae. It is known from the Early Permian Tambach Formation near the town of Tambach-Dietharz in Thuringia, Germany. Tambachia is the first trematopid to have been discovered outside the United States.

<span class="mw-page-title-main">Olsoniformes</span> Taxon of temnospondyl amphibians (fossil)

Olsoniformes is an extinct clade of dissorophoid temnospondyls, including the families Dissorophidae and Trematopidae. Most members of the clade were highly adapted to a terrestrial lifestyle. The clade was named in 2008 and is defined as the least inclusive clade containing Dissorophus multicinctus and Acheloma cumminsi but not Amphibamus grandiceps, Micromelerpeton credneri, or Apateon pedestris. Olsoniforms share various features such as a stout and low ilium and a thin cultriform process. The earliest-branching olsoniform is Palodromeus bairdi, from the Late Carboniferous of Ohio.

Perryella is an extinct genus of dvinosaurian(?) temnospondyl from the Permian of Oklahoma.

Parioxys is an extinct genus of temnospondyl amphibian from the Early Permian of Texas.

Nanobamus is an extinct genus of amphibamiform temnospondyl amphibian. The type species is Nanobamus macrorhinus. The type specimen is UCLAVP 3686, a skull recovered from the Lower Permian Arroyo Formation.

<span class="mw-page-title-main">Amphibamiformes</span> Extinct clade of temnospondyls

Amphibamiformes is an unranked clade with Dissorophoidea created by Schoch (2018). It encompasses all of the taxa traditionally considered to be "amphibamids", branchiosaurids, and hypothetically lissamphibians under the traditional temnospondyl hypothesis of lissamphibian origins. These taxa are typically small-bodied dissorophoids and form the sister group to Olsoniformes, which comprises dissorophids and trematopids.

References

  1. ↑ Williston, S. W. (1910). "Dissorophus Cope". The Journal of Geology. 18 (6): 526–536. Bibcode:1910JG.....18..526W. doi:10.1086/621768. ISSN   0022-1376. S2CID   224836685.
  2. ↑ Watson, D. M. S. (1920-01-01). "The Structure, Evolution and Origin of the Amphibia. The "Orders' Rachitomi and Stereospondyli". Philosophical Transactions of the Royal Society B: Biological Sciences. 209 (360–371): 1–73. Bibcode:1920RSPTB.209....1W. doi: 10.1098/rstb.1920.0001 . ISSN   0962-8436.
  3. ↑ Cope, E. D. (1882). "Third Contribution to the History of the Vertebrata of the Permian Formation of Texas". Proceedings of the American Philosophical Society. 20 (112): 447–461. ISSN   0003-049X. JSTOR   982692.
  4. ↑ Fritsch, Antonin (1881). "Fauna der Gaskohle und der Kalksteine der Permformations Bohmens". Self-published. 1 (3).
  5. 1 2 3 4 Milner, Andrew R. (2018-11-27). "Two primitive trematopid amphibians (Temnospondyli, Dissorophoidea) from the Upper Carboniferous of the Czech Republic". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 109 (1–2): 201–223. doi:10.1017/s1755691018000725. ISSN   1755-6910. S2CID   133895158.
  6. ↑ Williston, S. W. (1909). "New or Little-Known Permian Vertebrates: Trematops, New Genus". The Journal of Geology. 17 (7): 636–658. Bibcode:1909JG.....17..636W. doi: 10.1086/621665 . ISSN   0022-1376.
  7. ↑ Moodie, Roy L. (1911). "Recent Contributions to a Knowledge of the Extinct Amphibia". The American Naturalist. 45 (534): 375–384. doi: 10.1086/279219 . ISSN   0003-0147.
  8. ↑ Mehl, Maurice G. (1926). "Trematops Thomasi, a New Amphibian Species from the Permian of Oklahoma". The Journal of Geology. 34 (5): 466–474. Bibcode:1926JG.....34..466M. doi:10.1086/623333. ISSN   0022-1376. S2CID   140610416.
  9. 1 2 3 Olson, Everett Claire (1941). "The Family Trematopsidae". The Journal of Geology. 49 (2): 149–176. Bibcode:1941JG.....49..149O. doi:10.1086/624952. ISSN   0022-1376. S2CID   129647184.
  10. 1 2 Dilkes, David W. (1990-06-21). "A new trematopsid amphibian (Temnospondyli: Dissorophoidea) from the Lower Permian of Texas". Journal of Vertebrate Paleontology. 10 (2): 222–243. doi:10.1080/02724634.1990.10011809. ISSN   0272-4634.
  11. ↑ Olson, Everett C. (Everett Claire), 1910-1993. (1956). Fauna of the Vale and Choza : 12, a new Trematopsid amphibian from the Vale formation. Chicago Natural History Museum. OCLC   910497526.{{cite book}}: CS1 maint: multiple names: authors list (link) CS1 maint: numeric names: authors list (link)
  12. ↑ Milner, Andrew R. (1985). "On the identity of Trematopsis seltini (Amphibia: Temnospondyli) from the Lower Permian of Texas". Neues Jahrbuch für Geologie und Paläontologie, Monatshefte. 1985 (6): 357–367. doi:10.1127/njgpm/1985/1985/357.
  13. 1 2 Berman, David S.; Reisz, Robert R.; Eberth, David A. (1985). "Ecolsonia cutlerensis, an early Permian dissorophid amphibian from the Cutler Formation of north-central New Mexico". New Mexico Bureau of Mines and Minerals Research Circular. 191: 1–31.
  14. ↑ Olson, Everett C. (Everett Claire), 1910-1993. (1970). Trematops stonei sp. nov. (Temnospondyli: amphibia) from the Washington Formation, Dunkard Group, Ohio. Cleveland Museum of Natural History. OCLC   36073554.{{cite book}}: CS1 maint: multiple names: authors list (link) CS1 maint: numeric names: authors list (link)
  15. ↑ Eaton, Theodore Hildreth, Verfasser (1973). A Pennsylvanian Dissorophid Amphibian from Kansas. OCLC   1068422466.{{cite book}}: |last= has generic name (help)CS1 maint: multiple names: authors list (link)
  16. ↑ Milner, Andrew R. (1985). "On the identity of the amphibian Hesperoherpeton garnettense from the Upper Pennsylvanian of Kansas". Palaeontology. 28 (4): 767–776.
  17. ↑ Bolt, John R. (1974). A trematopsid skull from the Lower Permian, and analysis of some characters of the dissorophoid (Amphibia, Labyrinthodontia) otic notch / John R. Bolt. Chicago: Field Museum of Natural History. doi:10.5962/bhl.title.3432.
  18. 1 2 Bolt, John R. (1974). Osteology, function, and evolution of the Trematopsid (Amphibia: Labyrinthodontia) nasal region. Field Museum of Natural History. OCLC   914664.
  19. ↑ Olson, Everett C. (1985). "A Larval Specimen of a Trematopsid (Amphibia: Temnospondyli)". Journal of Paleontology. 59 (5): 1173–1180. ISSN   0022-3360. JSTOR   1305010.
  20. ↑ Dilkes, David W. (1991). "Reinterpretation of a larval dissorophoid amphibian from the Lower Permian of Texas". Canadian Journal of Earth Sciences. 28 (9): 1488–1492. Bibcode:1991CaJES..28.1488D. doi:10.1139/e91-130. ISSN   0008-4077.
  21. 1 2 3 Schoch, Rainer R.; Milner, Andrew R. (2014). Sues, Hans-Dieter (ed.). Handbuch der Paläoherpetologie Part 3A2. Temnospondyli I. Stuttgart: Verlag Dr. Friedrich Pfeil. ISBN   978-3-931516-26-0. OCLC   580976.
  22. ↑ Gee, Bryan M.; Reisz, Robert R. (2019-09-10). "The amphibamiform Nanobamus macrorhinus from the early Permian of Texas". Journal of Paleontology. 94 (2): 366–377. doi:10.1017/jpa.2019.72. ISSN   0022-3360. S2CID   203119272.
  23. ↑ Berman, David S; Reisz, Robert R.; Eberth, David A. (1987-09-16). "A new genus and species of trematopid amphibian from the Late Pennsylvanian of north-central New Mexico". Journal of Vertebrate Paleontology. 7 (3): 252–269. doi:10.1080/02724634.1987.10011659. ISSN   0272-4634.
  24. 1 2 Dilkes, David W. (1993). "Biology and evolution of the nasal region in trematopid amphibians". Palaeontology. 36 (4): 839–853.
  25. ↑ Sumida, Stuart S.; Berman, David S.; Martens, Thomas (1998). "A new trematopid amphibian from the Lower Permian of central Germany". Palaeontology. 41 (4): 605–629.
  26. ↑ Berman, David S; Henrici, Amy C.; Brezinski, David K.; Kollar, Albert D. (2009). "A New Trematopid Amphibian (Temnospondyli: Dissorophoidea) from the Upper Pennsylvanian of Western Pennsylvania: Earliest Record of Terrestrial Vertebrates Responding to a Warmer, Drier Climate". Annals of Carnegie Museum. 78 (4): 289–318. doi:10.2992/007.078.0401. ISSN   0097-4463. S2CID   85974488.
  27. 1 2 Berman, David S; Henrici, Amy C.; Martens, Thomas; Sumida, Stuart S.; Anderson, Jason S. (2011). "Rotaryus gothae , a New Trematopid (Temnospondyli: Dissorophoidea) from the Lower Permian of Central Germany". Annals of Carnegie Museum. 80 (1): 49–65. doi:10.2992/007.080.0106. ISSN   0097-4463. S2CID   84780478.
  28. 1 2 Polley, Brendan P.; Reisz, Robert R. (2011-03-21). "A new Lower Permian trematopid (Temnospondyli: Dissorophoidea) from Richards Spur, Oklahoma". Zoological Journal of the Linnean Society. 161 (4): 789–815. doi: 10.1111/j.1096-3642.2010.00668.x . hdl: 1807/18982 . ISSN   0024-4082.
  29. ↑ Maddin, Hillary C.; Reisz, Robert R.; Anderson, Jason S. (2010-07-02). "Evolutionary development of the neurocranium in Dissorophoidea (Tetrapoda: Temnospondyli), an integrative approach: Dissorophoid neurocranial development". Evolution & Development. 12 (4): 393–403. doi:10.1111/j.1525-142X.2010.00426.x. PMID   20618435. S2CID   10724489.
  30. ↑ Dilkes, David (2015-10-22). "Carpus and tarsus of Temnospondyli". Vertebrate Anatomy Morphology Palaeontology. 1: 51. doi: 10.18435/B5MW2Q . ISSN   2292-1389.
  31. ↑ Gee, Bryan M.; Bevitt, Joseph J.; Reisz, Robert R. (2019-03-12). "A Juvenile Specimen of the Trematopid Acheloma From Richards Spur, Oklahoma and Challenges of Trematopid Ontogeny". Frontiers in Earth Science. 7: 38. Bibcode:2019FrEaS...7...38G. doi: 10.3389/feart.2019.00038 . ISSN   2296-6463.
  32. ↑ Anderson, Jason S.; Henrici, Amy C.; Sumida, Stuart S.; Martens, Thomas; Berman, David S. (2008-03-12). "Georgenthalia clavinasica, a new genus and species of dissorophoid temnospondyl from the Early Permian of Germany, and the relationships of the family Amphibamidae". Journal of Vertebrate Paleontology. 28 (1): 61–75. doi:10.1671/0272-4634(2008)28[61:gcanga]2.0.co;2. ISSN   0272-4634. S2CID   55943106.
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