The 2010's in the eyes of temnospondyls

As many people know, I am an avid sports fan - comes with the territory growing up in L.A., I suppose. Unsurprisingly, I'm also a big fan of sports analysis shows and blogs, which form the inspirational backbone of the last #TemnospondylTuesday post of 2019. One of my favorites is Matthew Berry's Love/Hate column for fantasy football. I essentially grew up without NFL team since both the Rams and the Raiders skipped town after the '94 season (when I was born), and the Rams didn't move back until the 2016 season (when I moved to Canada). As most Canadians know, the CFL is about as popular in Canada as curling is in California, so there's not much going on with that front. So fantasy football is about as exciting as it gets for football; the Rams' current struggles aren't helping either, and does anybody actually know any Chargers fans who live in L.A.? Anyway, Berry often starts off his columns with a story of semi-relevance to his column highlighting which NFL players you should sit or start for the upcoming week, a model that I'll be following in the next section. There is A LOT in this post (perks of spending the post-holiday time cat-sitting for my labmate, Kayla), so buckle up and let's get after it!

I started this blog at the start of the new year, and I have to say, I'm pleasantly surprised that I both managed to keep blogging as semi-regularly as I did and that it got as much traction (a lot not from social media, I might add)! There is clearly an appetite for all things temnospondyl, and I am here for it! Stay tuned as I probably start the year pretty strong on a weekly basis and then fall off the face of the earth for a few weeks in the summer. And keep writing me! Sometimes I don't always respond or I'm real slow to do so, but it's not because I'm getting tired of hearing about how much you like temnospondyls - it's because I have a day job (and a night job) as a PhD student...

When I started out, I wrote up a list of topics that I thought I'd blog about. I've done a few of those, but mostly I've just been spitballing as things started coming to me. I always think you should write what you want, not what you said you would, so here we are. The end of the year auspiciously falls on a #TemnospondylTuesday, thereby imposing an entirely arbitrary sense of particular responsibility to send the blog off on a high note. I thought about doing a summary of temno research in 2019 (meh, sorta predictable) or highlighting stuff I'm working on for next year (too specific). As usual, as I asked my former labmate, Yara Haridy, what she thought I should write. She suggested I do "temnos of the last decade, 2000-2010." After I reminded her that it was in fact about to be 2020, and not 2011, she started spitballing. "What has changed"; "what do we know"; "who's been kicked out of the fam." The alternative prompt was "you just woke up from a coma and you hadn't read a single paper but wanted to be caught up - what do you have to know?" This situation doesn't actually require a coma since most people don't read temno papers. I'm not actually sure most of them want to be caught up either, but they say that overexposure (sometimes) builds an affinity for something, so might as well try it, right? 

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Ten Things I Love, Hate, or Love and Hate

One of my favourite features from the ESPN columns is the one that ESPN senior writer Zach Lowe puts out each week: "Ten Things I Like and Don't Like," which involves a lot of videos (sorry, I don't have any candids of my subject matter), detailed analysis, and give-it-to-me-straight honesty (which can still be complementary). I've come up with my own "Ten Things," which will probably be easy to figure out where each lies on the 'Love-Hate' spectrum and which are listed in absolutely no particular order. 

1. Stereospondyls are in vogue. Whether you buy it or not, and a lot of people I know do not buy it, the hypothesis that Chinlestegophis jenkensi is a stem caecilian and that modern amphibians derived from two temnospondyl clades, not just the dissorophoids, has put Stereospondyli on the map. I'd like to say "back on the map," since stereospondyls are the OG temnos going back to Richard Owen in the 1840s, but let's be real, they've never been on the map. But they should be! Stereospondyls are freaking amazing! They are your worst nightmare if you live near a body of water! Ned Colbert said they were like "queerly proportioned beasts such as we might encounter in mediaeval manuscripts." Who doesn't love that?!

I'll equivocate with my own personal opinion - I find the idea of lissamphibian diphyly attractive, and Chinlestegophis certainly looks like a caecilian (more so than 'lepospondyls' anyway), but I think we need some more fossils (I know of some ongoing projects) and analyses before we start rewriting textbooks. 

Digital renderings and reconstruction of Chinlestegophis jenkensi (figure from Pardo et al., 2017)

2. No temnospondyl is too good to be cut. ​​So we're not at the level of cutting holotypes just yet, but why bother when your sample size is in the dozens and nobody else cares about 'em? Temnospondyls have been getting cut for a long time; you can go back to Richard Owen who identified the labyrinthine infolding (plicidentine) of Mastodonsaurus that gave the 'labyrinthodonts' their hallmark name. Armand de Ricqlès also cut a lot of different temnospondyl elements back in the day, and now that histological methods and the pre-sampling conservation of target specimens has drastically improved, almost nothing is off limits. Nearly every major clade of temnospondyls has been sampled to date, although it continues to skew heavily toward stereospondyls, which are larger and more often preserved in mass death assemblages. Given that temnospondyls do nearly everything other than take to the skies, workers of other groups looking for a precedent or a comparative data point for more uncommon microanatomical / histological features (e.g., pachyostosis, non-pathological calcified cartilage, thin sections of skulls) should definitely give temnos a thorough look! You can find more on specific studies later in this blog.

Humeral thin sections of the branchiosaurid Apateon from the Permo-Carboniferous of Germany (figure from Sanchez et al., 2010b).

Vertebral thin sections of the plagiosaurids Plagiosuchus and Gerrothorax from the Triassic of Europe (figure from Danto et al., 2016).

3. Not enough independent testing of phylogenetic topologies. I'd like to see more independent workers producing novel phylogenetic matrices to test longstanding (or not so longstanding) topologies. A lot of the existing ones admittedly derive from a small handful of workers and are essentially passed along, nearly unmodified, with simple taxon additions (I'm guilty of this too; it's fast and also nice to assume that previously published work is rigorous). This isn't to say that I think that the base matrices are wrong or bad, but we also won't know how robust they are until somebody else tries to do it from scratch. At least independently rescoring of all sampled taxa to assess the rigor of codings would be nice. 

The phylogeny of temnospondyls from Schoch's (2013) analysis mapped onto stratigraphic occurrence; this matrix forms the skeleton for the majority of larger-scale temnospondyl analyses that have followed.

4. The Temnospondyl Handbook part 2. The companion to the Paleoherpetology Handbook Part 3B: Stereospondyli, published in 2000 by Rainer Schoch and Andrew Milner, was published in 2014 (listing on publisher's website here). It's a nicely laid out book, printed on that glossy matte paper, and with a ton of figures, and since there's been more work (especially systematics-wise) on the stereospondyl side of Temnospondyli, it's not super outdated yet in comparison to the stereospondyl handbook. It's a fantastic reference if you need a side-by-side comparison of the osteology or taxonomic diagnoses of various taxa, the descriptions of some of which date back to the early 1900s and are thus either not very accessible or at least not "CTRL + F"-able. I've taken it to with me to at least five major U.S. cities, and it's probably the only book that I'm guaranteed to open every week.
    That being said, there's a few downsides – it's pricey (over $100 USD, even on used book sites), the binding isn't great (personal observation), and it's not exactly peer-reviewed like journal articles are. This creates a lot of problems because there are taxonomic acts, whether erection (or resurrection) of new taxa (e.g., Nanobamus macrorhinus​) and suggested placements of taxa in untraditional clades (e.g., Parioxys ​as a dissorophid), which even if correct, aren't properly tested in this book. That can make it challenging for researchers who don't have easy access to this (you're unlikely to get all 150 pages scanned vs. getting a temporary loan through systems like ILLIAD, if you have access to that) who would otherwise not be aware of a lot of these acts. 

5. Big comparative studies. Like most other groups of animals, temnospondyls are seeing a lot more "big-picture" work looking at trends and patterns within the group. Plenty of descriptive work to go around here, such as work on the functional anatomy of the interpterygoid vacuities (Lautenschlager et al., 2016; Witzmann & Werneburg, 2017; Witzmann & Ruta, 2019) and modeling of feeding habits (e.g., Fortuny et al., 2011), but there's also been stuff on macroevolutionary trends and diversity (e.g., Angielczyk & Ruta, 2012; Tarailo, 2018; ​Pérez-Ben & Gomez, 2019; Pérez-Ben et al., 2019) and of course on phylogeny (e.g., Schoch, 2013; Eltink et al., 2019). Several paleoecological papers explored topics such as metamorphosis and development (e.g., Schoch, 2010; 2014b), function of dermal ornamentation (e.g., Witzmann et al., 2010; Rinehart & Lucas, 2013), general physiology (Witzmann & Brainerd, 2017), pathologies (e.g., Witzmann et al., 2014), and trackways (e.g., Lucas et al., 2010; Bird et al., 2019). There have been a good number of broader histological studies, especially on limbs (e.g. Muhkerjee et al., 2010; Sanchez et al., 2010c), but also on osteoderms (Witzmann & Soler-Gijón, 2010) and vertebrae (Konietzko-Meier et al., 2014; Danto et al., 2016, 2017). One of the few downsides to these bigger picture studies is the increasing prevalence of supertrees that are used to map character distributions; these are often compiled by splicing together different matrices (or even just recovered topologies) from different studies that are obviously going to be the most convenient for whatever the authors are trying to test (if not show).

Deformation and stress loads and distribution in various temnospondyl taxa based on different models of bite style (figure from Fortuny et al., 2011).

6. Temno paleoart is flourishing!

The giant late-surviving temnospondyl Siderops in the spring. #SciArt https://t.co/frSOgGTAYX pic.twitter.com/BG4UMMBMl3

— John Conway (@thejohnconway) March 6, 2017

BEHOLD #CACOPS!! New paper by @Koskinonodon & Yara Haridy on this lil #amphibian death trap's teeth! https://t.co/EmnQvbuuv4#paleontology pic.twitter.com/Cx7Fil2GzC

— Brian Engh (@BrianEngh_Art) September 13, 2017

Quick morning warmup speedpaint (1 hr) of a pair of Platyhystrix (a Permian dissorphoid temnospondyl). It is based on a dream I had last night where I found a pair of them walking in a drying creek #paleoart #sciart #TemnospondylTuesday pic.twitter.com/ONbZlwg8it

— Gabriel N. U. (@SerpenIllus) March 13, 2019

I ended up with this "rotten banana" pattern of dull yellow, dark greyish-red, and tiny salmon-colored spots after closely referencing the tiger salamander and red-spotted newt. #paleoart #sciart #Eryops pic.twitter.com/R4F0r1gNrT

— Liam Elward Paleoart (@paleobyliam) January 27, 2019

New #Patreon post: a poster-sized version of my new @cpdinosaurs Mastodonsaurus art can be yours for just $1 a month! That's a lot of slimy, algae-infested water for your dollar, and you get access to heaps more high-res #paleoart, essays and videos too. https://t.co/RahECWQHjl pic.twitter.com/ayN9O9Xn8q

— Mark Witton (@MarkWitton) April 16, 2019

6B. Browers giving you issues with embedded tweets.
This has nothing to do with temnospondyls, but you'll notice that I embedded a lot of different tweets here. Depending on your browser and ad-block / tracking / cookie settings, you may not be able to view the associated images and instead just see things as a block of text. Sorry...it's not me or my website builder, it's the internet...

7. Participation in temnospondyl research by underrepresented minorities is not flourishing...
No surprises here, but most temnospondyl work has historically been done by white dudes. The earliest workers were guys like Alexey Bystrow, Robert Carroll, E.C. Case, E. D. Cope, Al Romer, D.M.S. Watson, and Samuel Williston, and all of those guys were from Europe or North America. Margaret Steen was one of the few prominent women to work on temnospondyls in the early 20th century, with a particular focus on coal swamp specimens (e.g., Steen, 1930, 1934, 1938). Temnospondyl research hasn't really diversified much in the late 20th century, whose most prominent workers were still dudes, such as Dave Berman, John Bolt, John Cosgriff, Andrew Milner, Ivan Novikov, and Mikhail Shishkin, although Eleanor Daly did her PhD dissertation (1981) on what ended becoming Eoscopus locklardi, an amphibamiform from Kansas, and both Anne Warren (now emeritus at LaTrobe University in Australia) and Claudia Marsicano (Universidad de Buenos Aires in Argentina) both joined the temnospondyl efforts starting in the 80s and 90s (arguably the majority of their work and their most prominent publications have been in the 2000s).

Representation from outside of Europe/North America and among women *has* improved in the 21st century - researchers such as Graciela Piñeiro (Universidad de la República), Dorota Konietzko-Meier (Opole University), and Sophie Sanchez (Uppsala University) are some of the most established female temnospondyl workers; Hillary Maddin (Carleton University) and Nadia Fröbisch (Museum für Naturkunde) also have a great track record, although they're both trending more toward evo-devo these days. Julia McHugh (Museums of Western Colorado) and Kat Pawley (formerly LaTrobe) both did their dissertations on temnospondyls but Julia hasn't published on temnos in a few years, and Kat is out of the field now. We're also starting to see some homegrown researchers outside of the Euramerica complex, particularly as temno research is picking up in South America; Estevan Eltink (Universidade Federal do Vale do São Francisco in Brazil), Sérgio Dias-da-Silva (Universidade Federal de Santa Maria in Brazil), and Dhurjati Sengupta (Indian Statistical Institute) are three who immediately come to mind. My buddy Calvin So (PhD student at George Washington), Ben Otoo (PhD student at UChicago), Elzbieta Teschner (Opole University), and Sanjukta Chakravorti (recent PhD grad at the Indian Statistical Institute) are some up-and-coming women and/or POC to keep an eye on, and please look for ways to be inclusive in your temno research teams! 

​8. Almost 200 years of temnospondyl research and there's still zero consistency in what to call enlarged palatal teeth in temnospondyls.
Fangs? Tusk? "Fangs?" "Tusks?" I've already blogged about this before, but it sure it irritating (not that I'm any better at maintaining consistency) to have a bunch of terms, really none of which are appropriate in a strict sense anyway, floating around in the literature. Just one of those pet peeves things that make academics special... [Zach Lowe often has one short toss-away in his "10 Things," so this is my equivalent]

9. Jurassic World - The Game depictions of temnospondyls.
Temnospondyls haven't made it into a documentary since BBC's 2005 documentary "Walking with Monsters" in which an aestivating Rhinesuchus got eviscerated by a giant gorgonopsid (it's episode 6 if you want to watch the horror). They didn't make it into the Jurassic Park reboot either, but they did make it into the 2015 mobile game, the one that's like zoo tycoon, not the one where you use a drone to somehow collect DNA from AR dinosaurs and that destroys your battery and your data plan simultaneously. There are a bunch of probable temnospondyls - Koolasuchus, Microposaurus, 'Labyrinthodontia,' which I can only assume is supposed to be Mastodonsaurus (but also Mastodonsaurus is its own thing), Metoposaurus (!), Prionosuchus . Since Mastodonsaurus is an actual animal, it's not clear what 'Labyrinthodontia' is supposed to be? But it's in the "Common" tier, so evidently not a very good one. The weird little factoid for the first level states: "Did you know that it's believed that the Labyrinthodontia is the progenitor of all modern amphibians?" I'm pleasantly surprised that Metoposaurus is not only in the "Legendary" tier (the highest one) but can also be collected only through VIP packs (pay-to-play). 

All things considered, the base models look okay. Mastodonsaurus....something weird's going on with that dude's eyes, and Microposaurus' head just doesn't look right. But other than that, proportions and anatomy aren't totally off-base. It's when you get to the "evolved" (powered-up) forms that things get really weird...whoever designed this game decided that "evolution" meant you got amazingly flamboyant, and as far as the temnospondyls go, grew some wild dorsal fins! The one on Prionosuchus is particularly...magical.

Maximum level Metoposaurus.

Maximum level Prionosuchus.

Maximum level Koolasuchus.

Maximum level Mastodonsaurus.

On the off-chance that anyone from Ludia is reading this blog...I can help you fix your temnospondyls!

10. Newly outdated signage in newly renovated exhibits.
​Nothing says "new year" like new exhibits with signs that are already out of date (the revamped University of Michigan Natural History Museum is amazing in spite of this - check it out for sure!). 

Congratulations to @Koskinonodon for being the first person to render a label in the new Smithsonian Deep Time exhibit obsolete. pic.twitter.com/hk6iJVGwoN

— Christian Kammerer (@Synapsida) June 10, 2019

Let the renaming tour begin...I even went all out on my revised signage @Synapsida @Friedman_Lab #TemnospondylTuesday #metoposaurid #taxonomy pic.twitter.com/kGJdTMeciS

— Bryan Gee (@Koskinonodon) August 13, 2019

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POWER RANKINGS

As some people know, I am a HUGE basketball fan as well (Laker fan for life!), and I am an avid reader of the regular NBA power rankings that are put out by ESPN. These are almost entirely arbitrary rankings (although this is not a "list of Bryan's favourite and least favourite temnospondyls") and are based on zero quantitative metrics beyond my own perceptions of the body of the work that's been put out there, but it did add up to a nice, round 20 clades.

Stars (Lebron James tier)

1. Capitosaurs

Nothing says 'dynasty' like including the first temnospondyl to be named (Mastodonsaurus) and then still being so beloved that there are more than a dozen papers on your clade. Capitosaurs have had more new taxa described than in any other temno clade, and it's not even close; in the 2010s, we got Antarctosuchus polyodon Sidor, Steyer, & Hammer, 2014; Calmasuchus acri Fortuny, Galobart, & De Santisteban, 2011; Cyclotosaurus buechneri Witzmann, Sachs, & Nyhuis, 2016; Eocyclotosaurus appetolatus Rinehart, Lucas, & Schoch, 2015; Parotosuchus ptaszynskii Sulej & Niedzwiedzki, 2013; Samarabatrachus bjerringi Novikov, 2016; Selenocara rossica Novikov, 2016; Stanocephalus amenasensis Dahoume et al., 2016; Syrtosuchus morkovini Novikov, 2016; Tomeia witecki  Eltink, Stock Da-Rosa, & Dias-da-Silva, 2016; and Yuanansuchus maopingchangensis Liu, 2016. Additionally, there was the first report of any temnospondyl (a capitosaur) from Japan (Nakajima & Schoch, 2011), further description of material of E. appetolatus (Rinehart & Lucas, 2016), revision of capitosaur material from the Arctic (Kear et al., 2016), and revision of the well-known Parotuschus nasutus (Schoch, 2018). Capitosaurs were also the "test subjects" for a lot of the finite-element analysis (FEA) work done by Fortuny and colleagues (Fortuny et al., 2012, 2016; Marcé-Nogué et al., 2015), and some recent histo work suggested that some might have been more terrestrially capable than traditionally conceived (Mukherjee et al., 2019). 

Holotype of Yuanansuchus maopingchangensis from the Middle Triassic of China (figure from Liu, 2016).

Holotype of Antarctosuchus polyodon from the Middle Triassic of Antarctica (figure from Sidor et al., 2014).

2. Metoposaurids

No personal bias here, I promise! For reasons entirely unknown to myself or probably to anybody else, there's a massive surge in the popularity of metoposaurids. While they haven't enjoyed the same taxonomic explosion as some other clades (only three new taxa, Metoposaurus algarvensis Brusatte, Butler, Mateus, & Steyer, 2015; Panthasaurus maleriensis Chakravorti & Sengupta, 2018, Arganasaurus azerouali ​Buffa, Jalil, & Steyer, 2019, the last of which is really just re-allocation between genera), arguably no other "lower tetrapod" clade has sourced (or fallen victim to; you decide) more histological studies in the 2010s. Whether it's limbs (Konietzko-Meier & Sander, 2013; Konietzko-Meier & Klein, 2013; Teschner et al., 2018), vertebra (Konizetko-Meier et al., 2012, 2014; Danto et al., 2016, 2017; Gee et al., 2017; Gee & Parker, 2018), jaws (Gruntmejer et al,. 2019), or even an entire freaking skull (Gruntmejer et al., 2016), people all over Laurasia have been slicing and dicing these flat-headed, forward-eyed stereospondyls. We've also seen the first computer-assisted phylogenetic analyses of the clade in the form of three studies, all published within just seven months of each other (Chakravorti & Sengupta, 2018; Buffa et al., 2019; Gee et al., 2019). Tack on taphonomic studies (Lucas et al., 2010; Bodzioch, 2015; Bodzioch & Kowal-Linka, 2015), reappraisals (Fortuny et al., 2018), a little finite-element analysis to pair with your histology sampler (Fortuny et al., 2017, Konietzko-Meier et al., 2018), and the mass bonebed at Rotten Hill in the Texas Panhandle (Lucas et al., 2016), and we've got - dare I say it - a veritable buffet served up on a flat-headed platter. Not to mention I wracked up an obscene number of likes on Twitter for a Valentine's Day metoposaurid tweet... 

Roses are red
I'm kinda obsolete
My head's really flat
And opens like a toilet seat#ValentineASpecies #temnospondyl #metoposaurid pic.twitter.com/ixIqsVJWS6

— Bryan Gee (@Koskinonodon) February 14, 2019

Sidekicks (CJ McCollum tier)

3. Amphibamiforms

Some name changes and their perennial status as probable stem-lissamphibians keep amphibamiforms in the spotlight. For starters, the clade traditionally termed 'Amphibamidae' that encompassed small-bodied terrestrial dissorophoids is now restricted to just Amphibamus and Doleserpeton (Schoch, 2019a); 'Amphibamiformes' now encompasses all of the traditional amphibamids and branchiosaurids. There were a few new taxa: Rubeostratilia texensis Bourget & Anderson, 2011; Nanobamus macrorhinus Schoch & Milner, 2014 (then re-described by Gee & Reisz, 2019); Platyrhinops fritschi Werneburg, 2012; and Tersomius dolesensis Anderson & Bolt, 2013; as well as a thorough redescription of Doleserpeton annectens (Sigurdsen & Bolt, 2010), revisions of Tersomius texensis (Maddin et al., 2013), Milnererpeton huberi (Werneburg et al., 2013), Branchiosaurus salamandroides ​(Werneburg, 2012) and additional reports of Leptorophus (Schoch, 2014a). Werneburg (2019) revised a number of Permo-Carboniferous specimens from France, erecting the terrestrial amphibamiforms Anthracobamus fayoli and Montceaubatrachus platyrynchus and a branchiosaurid Branchiosaurus commentryensis. A complete but badly weathered branchiosaurid was also reported from Turkey by Fortuny et al. (2015). There were three great histological studies by Sanchez et al. (2010a, 2010b, 2010d) that used Apateon ​to examine different paleoecological and paleoenvironmental attributes, and Eichler & Werneburg (2010) examined a number of branchiosaurids using UV light. Schneider & Werneburg (2012) provided an updated stratigraphic framework of the Rotliegend branchiosaurids. Even if they prove to not be ancestral to all modern amphibians, their diminutive body size (referred to by some as 'miniaturization'), extensive sample size for some taxa, and exceptional preservation of soft tissue will keep them in the limelight for a long time.

Reconstruction of Doleserpeton annectens from the early Permian of Oklahoma (figure from Sigurdsen & Bolt, 2010).

WIP. This is a reconstruction of the stem-caecilian Chinlestegophis jenkinsi from the Triassic of Colorado that I am working on for my upcoming book (hopefully out by early 2019). This is a key fossil in the evolution of caecilians #paleoart #sciart #FossilFriday #caecilians pic.twitter.com/SJ5m7WhoCY

— Gabriel N. U. (@SerpenIllus) July 13, 2018

4. 'Latiscopids'

Latiscopids? Almasaurids? Rileymillerids? Who knows what to call them, but the long-known and eternally poorly understood small stereospondyls of the Late Triassic have a new skip in their step (or whatever you call the gait of these weird animals) following the description of Chinlestegophis jenkensi Pardo, Small, & Huttenlocker, 2017, another one of these small-bodied forms that has generated major shakeups (and corresponding controversy) in the debate over origins of modern amphibians. Pardo et al. recovered C. jenkensi as stereospondyls most closely related to brachyopoids and on the way to caecilians, the weird noodles of the amphibian family. If correct, this would indicate that modern amphibians are not, in fact, closely related to each other and that they have a much deeper common ancestor than previously thought, which might account for some of the peculiarities between caecilians and the grouping of frogs/toads + salamanders/newts.

5. Dissorophids

Dissorophids have had one heck of a renaissance in the 21st century. In the last decade, there have been five new taxa (Broiliellus reiszi Holmes, Berman, & Anderson, 2013; Cacops woehri Fröbisch & Reisz, 2012; Nooxobeia gracilis Gee, Scott, & Reisz, 2018; Reiszerpeton renascentis Maddin, Fröbisch, Evans, & Milner, 2013; and Scapanops neglecta Schoch & Sues, 2013), as well as the first phylogenetic analysis of the clade (Schoch, 2012), detailed descriptions of several eucacopines (Gee & Reisz, 2018; Liu, 2018), and some promising reports of Carboniferous-age material from the midcontinent of North America (May et al., 2011). Numerous sources indicate exciting forthcoming work from areas outside of the classic collecting localities for the clade...

Three specimens of Cacops morrisi from the early Permian of Oklahoma (figure from Gee & Reisz, 2018).

Sixth men (Lou Williams tier)

6. Trematosaurs

Although trematosaurs have a wide-ranging distribution, their study in the last decade has been largely carried by the Europeans. Time for someone to get back out to the Moenkopi in North America... Trematosaurs have quietly ended up with a lot of descriptive work, with several new taxa described: Aphaneramma gavialimimus Fortuny et al. 2018; Benthosuchus guseae Novikov 2012b; Microposaurus averyi Warren, 2012, Qantas samarensis Novikov, 2012; and Trematosaurus galae Novikov, 2010. Yes, Qantas is spelled just like the airline; no, it is not named after the airline. ​Several other taxa were re-examined, including Calamops paludosus (Sues & Schoch, 2013) and Benthosuchus (Novikov, 2012b), with reports of new material from Russia (Novikov, 2018) and Jordan (Schoch, 2011) and some of the first CT work on stereospondyls (Fernandez-Coll et al., 2019). 

Just a reminder that Trematosaurid Temnospondyls are awesome! There were two subfamilies, the shorter-snouted trematosaurines and the gharial-like, longirostrine lonchorhynchines. many of these amazing animals will appear in my book #paleoart #temnospondyls #sciart pic.twitter.com/X4Y24ElnEI

— Gabriel N. U. (@SerpenIllus) October 22, 2018

7. Plagiosaurs

Turns out that the OG "toilet seat heads" weren't just a one-hit wonder. Thanks to German researchers (mostly Rainer Schoch and Florian Witzmann), there's been a flurry of plagiosaur publications, including new taxon Megalophthalma ockerti Schoch, Milner, & Witzmann, 2014, a detailed survey of the cranial anatomy of the wide-ranging Gerrothorax pulcherrimus (Schoch & Witzmann, 2011; Witzmann et al., 2012), reconstruction of the cranial musculature in G. pulcherrimus (Witzmann & Schoch, 2013), a limb histology study of G. pulcherrimus (yes, people really like Gerrothorax; Sanchez & Schoch, 2013), and a comparative histo-vs.-CT study on Plagiosuchus (Konietzko-Meier & Schmitt, 2013).  Dias-da-Silva & Milner (2010) further clarified a plagiosaurid record from Brazil. 

Comparison of different skull morphologies of Gerrothorax pulcherrimus (figure from Schoch & Witzmann, 2011).

8. Rhinesuchids

This clade lives and dies by the Brazilian rhinesuchid Australerpeton cosgriffi, which has been a real stand-out among the rhinesuchids, with a cranial osteology packaged in with a phylogenetic analysis of Rhinesuchidae (Eltink et al., 2016) and two reports of new specimens (Eltink & Langer, 2014; Azevedo et al., 2019). A second phylogenetic analysis was published by Marsicano et al. (2017) just a short while later. Piñeiro et al. (2012) described a new taxon from Uruguay, Arachana nigra, and there are also promising reports of indeterminate rhinesuchids among the early Permian Brazilian assemblages (Cisneros et al., 2015). McHugh (2014) sliced and diced the eponymous Rhinesuchus. Arguably the ace-in-the-hole is the poorly known Broomistega, an individual of which was found cuddling a therapsid in a burrow (Fernandez et al., 2013).

Various specimens of Australerpeton cosgriffi from the Middle-Late Permian of Brazil (figure from Eltink et al., 2016).

The famous Broomistega-Thrinaxodon interspecies couple from the Early Triassic of South Africa (figure from Fernandez et al., 2013)

9. Trematopids

Trematopids live and die by the mantra of 'gen. et sp. nov.,' and I mean it very literally. In the last decade, we've seen four new taxa, Acheloma dunni Polley & Reisz, 2011; Fedexia striegeli Berman, Henrici, Brezinski, & Kollar, 2010; Mattauschia laticeps Milner, 2018; and Rotaryus gothae Berman, Henrici, Martens, Sumida, & Anderson, 2011, only one of which belongs to either a previously known genus or to a genus with more than one species in it (Acheloma). Outside of that, the only main work was a description of neurocranial ontogeny in A. dunni ​by Maddin et al. (2010) and the first CT analysis of a trematopid, published by Gee et al. (2019).

Holotype skull of Acheloma dunni from the early Permian of Oklahoma (figure from Polley & Reisz, 2011).

Role players (PJ Tucker tier)

10. Eryopoids

The most iconic Paleozoic temnospondyl (and arguably, tetrapod, other than Dimetrodon) might never have a heyday as great as the early 20th century when everybody and their mother (or father, let's be real) published papers about every specific region of the skeleton of Eryops. The 21st century has mostly seen simple reports / documentation of various taxa (mostly Eryops) from different parts of the world (mostly North America) (e.g., Werneburg et al., 2010; Witzmann, 2013; Aber et al., 2014; Voigt et al., 2014; Witzmann & Voigt, 2015; Huttenlocker et al., 2018). Keep your eyes peeled for a new species of Eryops ​though (Rasmussen et al., 2016)! On the descriptive side, Čerňanský et al. (2016) published a CT analysis of the quadratojugal of Eryops, and Werneburg & Berman (2012) touched up the osteology of Glaukerpeton avinoffi.

11. Dvinosaurs

Although there hasn't been a ton of work on dvinosaurs in the last decade, there remain exciting prospects for work on the clade as a whole. Cisneros et al. (2015) described two new taxa, Timonya anneae and Procuhy nazariensis, from the early Permian of northern Brazil, a historically under-surveyed area that's starting to produce a lot of really exciting new finds for Paleozoic tetrapods in general. Schoch & Voigt (2019) described Trypanognathus remigiusbergensis (try saying that five times fast) from an immaculate specimen collected in Germany, the first definitive dvinosaur report from central Europe in the lengthy history of this diverse and remarkably widespread clade.

Holotype of Trypanognathus remigiusbergensis from the Permo-Carboniferous boundary of Germany (figure from Schoch & Voigt, 2019).

12. Stereospondylomorphs (in the paraphyletic sense)

A pretty low-key decade for the early stereospondylomorphs (the non-stereospondyl stereospondylomorphs to be technical). A few new taxa popped up as well, such as Konzhukovia sangabrielensis Pacheco, Eltink, Müller, & Dias-da-Silva, 2017 and Sclerocephalus stambergi Klembara & Steyer, 2012. The holotype of the type species of Sclerocephalus, S. haeuseri, which was long thought to be lost, also reappeared this decade (Schoch et al., 2019). Stereospondylomorphs also got a brand new phylogenetic analysis (Eltink et al., 2019), with the erection of a few new clades. Witzmann & Brainerd (2017) modeled the physiology of Archegosaurus decheni. Like dvinosaurs, the early stereospondylomorph record from South America is developing really quickly and in very interesting ways, with a lot of connections to eastern Eurasia (particularly Russia) that are changing a lot of the ways in which we historically conceived of biogeographic distribution and radiation of stereospondylomorphs.

13. Micromelerpetids

Like rhinesuchids, micromelerpetid research rests largely on the eponymous Micromelerpeton; the only other major descriptive work is the first report of the clade in Africa (Werneburg et al., 2019) and a re-description of Limnogyrinus ​with a phylogenetic analysis (Schoch & Witzmann, 2018). Unlike rhinesuchids, there's no micromelerpetid cozying up with an amniote. BUT, micromelerpetids do show evidence of limb regeneration and preaxial polarity during limb formation (Frobisch et al., 2014, 2015), which makes them super interesting from the evo-devo side of things.

Riding the bench ("pick any obsolete rookie" tier)

14. Brachyopoids

The two brachyopoid clades, Brachyopidae and Chigutisauridae, have been pretty meager. The main work on the brachyopid front was Warren et al.'s (2011) documentation of tupilakosaur-like vertebrae (better known as the diplospondylous type) in Bothriceps. Chigutisaurids faired a little better. Compsocerops, originally described from India, was found in Brazil (Dias-da-Silva et al., 2012). Some chigutisaurid material from India was also sampled in a comparative histological analysis (Muhkerjee et al., 2010); word on the street is there's more histo coming out soon! There's hope on the horizon for brachyopoids if future work bears out their relationship to 'latiscopids,' and in turn, 'latiscopids'' close relationship to caecilians.

15. Rhytidosteids

Next to nothing on these small-bodied and typically horned temnos, which occupy an important position in the context of the early radiation of Mesozoic stereospondyls. Dias-da-Silva & Marsicano (2011) did provide the most comprehensive phylogenetic analysis of the group to date, although many of their results were challenged by Maganuco et al. (2014) who re-described Mahavisaurus dentatus. Sennikov & Novikov (2018) discussed possible trophic adaptations / specialization for hard-bodied invertebrates. 

16. Edopoids

Apparently little appetite for the "gluttonous faces" of the Paleozoic. Essentially no descriptive work beyond some new material of the relict Nigerpeton (Sidor, 2013) and only brief discussion in broader comparative papers (e.g., Fortuny et al., 2011; Schoch, 2019b). Bird et al. (2019) did squeeze a paper on a trackway attributed to an edopoid from the lower Carboniferous of the UK into 2019.

17. Dendrerpetids

Other than having the honour of being utilized as the outgroup for just about every Paleozoic temnospondyl study, dendrerpetids have been on the down and out for some time. New material and dedicated re-analysis to the ages-old question of "just how many dendrerpetids are there at Joggins" may inspire some work going forward (Maddin et al., 2018).

18. Lydekkerinids

What happens when you're small, fairly abundant, not closely related to modern amphibians, and phylogenetically well-resolved? You guessed it - you get sliced and diced (Canoville & Chinsamy, 2015; McHugh, 2015) and that's about it...

19. Zatracheidids

Does anybody still work on zatracheidids? Zero publications on this clade in the 2010s aside from their sporadic inclusion in broader analyses of temnospondyl macroevolutionary patterns (e.g., Angielczyk & Ruta, 2012; Witzmann & Ruta, 2019).

20. Lapillopsids

Guess what happens when you're small, one-of-a-kind, and a phylogenetic wildcard? You guessed it - no research in the 2010's! In fact, you only get six hits on Google Scholar from 2010 onward.

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Five predictions for the 2020's

I will freely admit that I am totally spitballing here, but it'll be interesting to see how these pan out.

1. There will be no fewer than 15 new genera and 40 new species described, but at least 5 genera and 10 species will be sunk into synonymy or formalized as nomina dubia (the ones that are new and the ones that are sunk are not necessarily mutually exclusive). 
2. Temnospondyl phylogeny will be shaken up enough, or at least sufficiently questioned, to force everyone to stop using Dendrysekos helogenes as the outgroup for the vast majority of studies.
3. South America will prove to be the temnospondyl hotspot of the decade (in the sense of an appreciable number of new taxa with important macroevolutionary and biogeographic implications). But at the same time...
4. ...the classic coal swamp localities and their constituents will come back into fashion, as workers look to re-assess some of the earliest records of many of the major clades. Expect no fewer than a dozen papers specifically on these types of taxa (this is not a foreshadowing of my own predicted work).
5. Somebody will finally name a temnospondyl after a famous, living non-scientist (this may be foreshadowing my predicted work).

Five things I wish for but don't reasonably expect in the 2020's

1. A fossilized temnospondyl tadpole, whatever that would look like.
2. A metoposaurid with dinosaur remains in its gut. #StickItToTheAmniotes
3. A temnospondyl (I'll take literally any clade) from California.
4. Strong evidence for identifying sexual dimorphism in some taxon.
5. Taylor Swift to follow me on Twitter (yes, I know she doesn't follow anyone on Twitter).

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​Social Media Round-up

If you're looking to get more doses of temnospondyls on Twitter, be sure to follow some of the following folks who actively post content:

• Aaron Kufner (PhD student at University of Wisconsin-Madison): @Nomen_Oblitum
• Ben Otoo (PhD student at University of Chicago): @baeleozoic
• Calvin So (PhD student at George Washington University): @calvincfso
• Gabriel Ugueto (Paleoartist): @SerpenIllus
• Hillary Maddin (Prof at Carleton University): @Evo_Deva
• Jason Pardo (PhD student at University of Calgary): @incisorial

And of course, make sure that you're following me @Koskinonodon!

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(Long list of references for this post, so I've hidden them below the "Read More" line). Note that this is not at all an exhaustive list of all temnospondyl publications since 2010; I haven't even cited some of my own here.

Refs

• Aber SW, Peterson N, May WJ, Johnston P, Aber JS. 2014. First report of vertebrate fossils in the Snyderville Shale (Oread Formation; Upper Pennsylvanian), Greenwood County, Kansas. Transactions of the Kansas Academy of Science, 117(3-4): 193-203.
• Anderson JS, Bolt JR. 2013. New information on amphibamids (Tetrapoda, Temnospondyli) from Richards Spur (Fort Sill), Oklahoma. Journal of Vertebrate Paleontology, 33(3): 553-567.
• Angielczyk KD, Ruta M. 2012. The roots of amphibian morphospace: a geometric morphometric analysis of Paleozoic temnospondyls. Fieldiana Life and Earth Sciences, 2012(5): 40-59.
• Azevedo KL, Vega CS, Soares MB. 2019. A new specimen of Australerpeton cosgriffi Barberena, 1998 (Stereospondyli: Rhinesuchidae) from the Middle/Upper Permian Rio do Rasto Formation, Paraná Basin, Brazil. Revista Brasileira de paleontologia, 20(3): 333-344.
• Berman DS, Henrici AC, Brezinski DK, Kollar AD. 2010. 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): 289-319.
• Berman DS, Henrici AC, Martens T, Sumida SS, Anderson JS. 2011. Rotaryus gothae, a new trematopid (Temnospondyli: Dissorophoidea) from the Lower Permian of Central Germany. Annals of Carnegie Museum, 80(1): 49-65.
• Bird HC, Milner AC, Shillito AP, Butler RJ. 2019. A lower Carboniferous (Visean) tetrapod trackway represents the earliest record of an edopoid amphibian from the UK. Journal of the Geological Society.
• ​Bodzioch A, Kowal-Linka M. 2012. Unraveling the origin of the Late Triassic multitaxic bone accumulation at Krasiejów (S Poland) by diagenetic analysis. Palaeogeography, Palaeoclimatology, Palaeoecology, 346: 25-36.
• Bodzioch A. 2015. Idealized model of mineral infillings in bones of fossil freshwater animals, on the example of Late Triassic metoposaurs from Krasiejów (Poland). Austin Journal of Earth Science, 2(1): 1-6.
• Bourget H, Anderson JS. 2011. A new amphibamid (Temnospondyli: Dissorophoidea) from the Early Permian of Texas. Journal of Vertebrate Paleontology, 31(1): 32-49.
• Brusatte SL, Butler RJ, Mateus O, Steyer JS. 2015. A new species of Metoposaurus from the Late Triassic of Portugal and comments on the systematics and biogeography of metoposaurid temnospondyls. Journal of Vertebrate Paleontology, 35(3): e912988.
• Buffa V, Jalil NE, Steyer JS. Redescription of Arganasaurus (Metoposaurus) azerouali (Dutuit) comb. nov. from the Upper Triassic of the Argana Basin (Morocco), and the first phylogenetic analysis of the Metoposauridae (Amphibia, Temnospondyli). Papers in Palaeontology.
• Canoville A, Chinsamy A. 2015. Bone microstructure of the stereospondyl Lydekkerina huxleyi reveals adaptive strategies to the harsh post Permian‐extinction environment. The Anatomical Record, 298(7): 1237-1254.
• Čerňanský A, Witzmann F, Klembara J, Van Heteren AH. 2016. The quadratojugal of Eryops studied by computed tomography and the morphological variability of foramina and canals in the quadratojugal of basal tetrapods. The Anatomical Record, 299(8): 1073-1079.
• Chakravorti S, Sengupta DP. 2019. Taxonomy, morphometry and morphospace of cranial bones of Panthasaurus gen. nov. maleriensis from the Late Triassic of India. Journal of Iberian Geology, 45(2): 317-340.
• Cisneros JC, Marsicano C, Angielczyk KD, Smith RM, Richter M, Fröbisch J, Kammerer CF, Sadleir RW. 2015. New Permian fauna from tropical Gondwana. Nature communications, 6: 8676.
• ​Danto M, Witzmann F, Fröbisch NB. 2016. Vertebral development in Paleozoic and Mesozoic tetrapods revealed by paleohistological data. PloS One, 11(4): e0152586.
• Danto M, Witzmann F, Pierce SE, Fröbisch NB. 2017. Intercentrum versus pleurocentrum growth in early tetrapods: A paleohistological approach. Journal of Morphology, 278(9): 1262-1283.
• Dias-da-Silva S, Milner AR. 2010. The pustulated temnospondyl revisited—a plagiosternine plagiosaurid from the Lower Triassic of Brazil. Acta Palaeontologica Polonica, 55(3): 561-564.
• Dias-da-Silva S, Marsicano C. 2011. Phylogenetic reappraisal of Rhytidosteidae (Stereospondyli: Trematosauria), temnospondyl amphibians from the Permian and Triassic. Journal of Systematic Palaeontology, 9(2): 305-325.
• Dias‐da‐Silva S, Sengupta DP, Cabreira SF, Da Silva LR. 2012. The presence of Compsocerops (Brachyopoidea: Chigutisauridae)(Late Triassic) in southern Brazil with comments on chigutisaurid palaeobiogeography. Palaeontology, 55(1): 163-172.
• Eichler B, Werneburg R. 2010. Neufunde von Branchiosauriern aus dem Rotliegend von Freital bei Dresden im UV-Licht [New specimens of branchiosaurs from the Rotliegend (Sakmarian) of Freital near Dresden in ultra-violet light]. Journal of Central European Geology, 56(2): 137-57.
• Eltink E, Langer MC. 2014. A new specimen of the temnospondyl Australerpeton cosgriffi from the Late Permian of Brazil (Rio do Rasto Formation, Paraná Basin): comparative anatomy and phylogenetic relationships. Journal of Vertebrate Paleontology, 34(3): 524-538.
• Eltink E, Dias EV, Dias-da-Silva S, Schultz CL, Langer MC. 2016. The cranial morphology of the temnospondyl Australerpeton cosgriffi (Tetrapoda: Stereospondyli) from the Middle-Late Permian of Paraná Basin and the phylogenetic relationships of Rhinesuchidae. Zoological Journal of the Linnean Society, 176(4): 835-860.
• ​Eltink E, Schoch RR, Langer MC. 2019. Interrelationships, palaeobiogeography and early evolution of Stereospondylomorpha (Tetrapoda: Temnospondyli). Journal of Iberian Geology, 45(2): 251-267.
• Fernandez V, Abdala F, Carlson KJ, Rubidge BS, Yates A, Tafforeau P. 2013. Synchrotron reveals Early Triassic odd couple: injured amphibian and aestivating therapsid share burrow. PLoS One, 8(6): e64978.
• Fernández-Coll, M., Arbez, T., Bernardini, F. and Fortuny, J., 2019. Cranial anatomy of the Early Triassic trematosaurine Angusaurus (Temnospondyli: Stereospondyli): 3D endocranial insights and phylogenetic implications. Journal of Iberian Geology, 45(2), pp.269-286.
• Fortuny J, Galobart À, De Santisteban C. 2011. A new capitosaur from the Middle Triassic of Spain and the relationships within the Capitosauria. Acta Palaeontologica Polonica, 56(3): 553-567.
• Fortuny J, Marcé‐Nogué J, de Esteban‐Trivigno S, Gil L, Galobart À. 2011. Temnospondyli bite club: ecomorphological patterns of the most diverse group of early tetrapods. Journal of Evolutionary Biology, 24(9): 2040-2054.
• Fortuny J, Marcé‐Nogué J, Gil L, Galobart À. 2012. Skull mechanics and the evolutionary patterns of the otic notch closure in capitosaurs (Amphibia: Temnospondyli). The Anatomical Record, 295(7): 1134-1146.
• Fortuny J, Steyer JS, Hoşgör I. 2015. First occurrence of temnospondyls from the Permian and Triassic of Turkey: Paleoenvironmental and paleobiogeographic implications. Comptes Rendus Palevol, 14(4): 281-289.
• Fortuny J, Marcé‐Nogué J, Steyer JS, de Esteban-Trivigno S, Mujal E, Gil L. 2016. Comparative 3D analyses and palaeoecology of giant early amphibians (Temnospondyli: Stereospondyli). Scientific Reports, 6: 30387.
• ​Fortuny J, Marcé‐Nogué J, Konietzko‐Meier D. 2017. Feeding biomechanics of Late Triassic metoposaurids (Amphibia: Temnospondyli): a 3D finite element analysis approach. Journal of Anatomy, 230(6): 752-765.
• Fortuny J, Gastou, S., Escuillié, F., Ranivoharimanana, L. and Steyer, J.S., 2018. A new extreme longirostrine temnospondyl from the Triassic of Madagascar: phylogenetic and palaeobiogeographical implications for trematosaurids. Journal of Systematic Palaeontology, 16(8), pp.675-688.
• Fortuny J, Arbez T, Mujal E, Steyer JS. 2019. Reappraisal of ‘Metoposaurus hoffmani’ Dutuit, 1978, and description of new temnospondyl specimens from the Middle–Late Triassic of Madagascar (Morondava Basin). Journal of Vertebrate Paleontology: e1576701.
• Fröbisch NB, Bickelmann C, Witzmann F. 2014. Early evolution of limb regeneration in tetrapods: evidence from a 300-million-year-old amphibian. Proceedings of the Royal Society B: Biological Sciences, 281(1794): 20141550.
• Fröbisch NB, Bickelmann C, Olori JC, Witzmann F. 2015. Deep-time evolution of regeneration and preaxial polarity in tetrapod limb development. Nature, 527(7577): 231-234.
• Gee BM, Parker WG. 2018. Morphological and histological description of small metoposaurids from Petrified Forest National Park, AZ, USA and the taxonomy of Apachesaurus. Historical Biology, 31 pp.
• Gee BM, Parker WG, Marsh AD. 2017. Microanatomy and paleohistology of the intercentra of North American metoposaurids from the Upper Triassic of Petrified Forest National Park (Arizona, USA) with implications for the taxonomy and ontogeny of the group. PeerJ, 5: e3183.
• Gee BM, Reisz RR. 2018. Cranial and postcranial anatomy of Cacops morrisi, a eucacopine dissorophid from the early Permian of Oklahoma. Journal of Vertebrate Paleontology, 38(2): e1433186.
• Gee BM, Reisz RR. 2019. The amphibamiform Nanobamus macrorhinus from the early Permian of Texas. Journal of Paleontology, 12 pp.
• Gee BM, Bevitt JJ, Reisz RR. 2019. A juvenile specimen of the trematopid Acheloma from Richards Spur, Oklahoma and challenges of trematopid ontogeny. Frontiers in Earth Science, 7: 38.
• Gee BM, Parker WG, Marsh AD. 2019. Redescription of Anaschisma (Temnospondyli: Metoposauridae) from the Late Triassic of Wyoming and the phylogeny of the Metoposauridae. Journal of Systematic Palaeontology, 26 pp.
• ​Gruntmejer K, Konietzko-Meier D, Bodzioch A. 2016. Cranial bone histology of Metoposaurus krasiejowensis (Amphibia, Temnospondyli) from the Late Triassic of Poland. PeerJ, 4: e2685.
• ​Gruntmejer K, Konietzko-Meier D, Bodzioch A, Fortuny J. 2019. Morphology and preliminary biomechanical interpretation of mandibular sutures in Metoposaurus krasiejowensis (Temnospondyli, Stereospondyli) from the Upper Triassic of Poland. Journal of Iberian Geology, 45(2): 301-316.​​
• ​Huttenlocker AK, Henrici A, Nelson WJ, Elrick S, Berman DS, Schlotterbeck T, Sumida SS. 2018. A multitaxic bonebed near the Carboniferous–Permian boundary (Halgaito Formation, Cutler Group) in Valley of the Gods, Utah, USA: Vertebrate paleontology and taphonomy. Palaeogeography, Palaeoclimatology, Palaeoecology, 499: 72-92.
• Kear BP, Poropat SF, Bazzi M. 2016. Late Triassic capitosaurian remains from Svalbard and the palaeobiogeographical context of Scandinavian Arctic temnospondyls. Geological Society, London, Special Publications, 434(1): 113-126.
• Klembara J, Steyer JS. 2012. A New Species of Sclerocephalus (Temnospondyli: Stereospondylomorpha) from the Early Permian of the Boskovice Basin (Czech Republic). Journal of Paleontology, 86(2): 302-310.
• Konietzko-Meier D, Klein N. 2013. Unique growth pattern of Metoposaurus diagnosticus krasiejowensis (Amphibia, Temnospondyli) from the Upper Triassic of Krasiejów, Poland. Palaeogeography, Palaeoclimatology, Palaeoecology, 370: 145-157.
• Konietzko-Meier D, Sander PM. 2013. Long bone histology of Metoposaurus diagnosticus (Temnospondyli) from the Late Triassic of Krasiejów (Poland) and its paleobiological implications. Journal of Vertebrate Paleontology, 33(5): 1003-1018.
• ​Konietzko-Meier D, Schmitt A. 2013. A histological study of a femur of Plagiosuchus, a Middle Triassic temnospondyl amphibian from southern Germany, using thin sections and micro-CT scanning∙. Netherlands Journal of Geosciences, 92(2-3): 97-108.
• ​Konietzko-Meier D, Bodzioch A, Sander PM. 2013 (for 2012). Histological characteristics of the vertebral intercentra of Metoposaurus diagnosticus (Temnospondyli) from the Upper Triassic of Krasiejów (Upper Silesia, Poland). Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 103(3-4): 237-250.
• Konietzko-Meier D, Danto M, Gądek K. 2014. The microstructural variability of the intercentra among temnospondyl amphibians. Biological Journal of the Linnean Society, 112(4): 747-764.
• Konietzko-Meier D, Gruntmejer K, Marcé-Nogué J, Bodzioch A, Fortuny J. 2018. Merging cranial histology and 3D-computational biomechanics: a review of the feeding ecology of a Late Triassic temnospondyl amphibian. PeerJ, 6: e4426.
• Lautenschlager S, Witzmann F, Werneburg I. 2016. Palate anatomy and morphofunctional aspects of interpterygoid vacuities in temnospondyl cranial evolution. The Science of Nature, 103(9-10): 79.
• Liu J. 2016. Yuanansuchus maopingchangensis sp. nov., the second capitosauroid temnospondyl from the Middle Triassic Badong Formation of Yuanan, Hubei, China. PeerJ, 4: e1903.
• Lucas SG, Fillmore DL, Simpson EL. 2010. Amphibian body impressions from the Mississippian of Pennsylvania, USA. Ichnos, 17(3): 172-176.
• Lucas SG, Rinehart LF, Krainer K, Spielmann JA, Heckert AB. 2010. Taphonomy of the Lamy amphibian quarry: a Late Triassic bonebed in New Mexico, USA. Palaeogeography, Palaeoclimatology, Palaeoecology, 298(3-4): 388-398.
• Lucas SG, Rinehart LF, Heckert AB, Hunt AP, Spielmann JA. 2016. Rotten Hill: a Late Triassic bonebed in the Texas Panhandle. New Mexico Museum of Natural History and Science Bulletin, 72.
• Maddin HC, Reisz RR, Anderson JS. 2010. Evolutionary development of the neurocranium in Dissorophoidea (Tetrapoda: Temnospondyli), an integrative approach. Evolution & development, 12(4): 393-403.
• Maddin HC, Mann A, Atkins J, Paterson R, Mcfeeters B, Hebert B, Adams G, Thompson M. 2018. Recent tetrapod discoveries and the changing view of the Carboniferous.
• Maddin HC, Fröbisch NB, Evans DC, Milner AR. 2013. Reappraisal of the Early Permian amphibamid Tersomius texensis and some referred material. Comptes Rendus Palevol, 12(7-8): 447-461.​
• ​Marcé-Nogué J, Fortuny J, de Esteban-Trivigno S, Sánchez M, Gil L, Galobart À. 2015. 3D computational mechanics elucidate the evolutionary implications of orbit position and size diversity of early amphibians. PloS one, 10(6): e0131320.
• Marsicano CA, Latimer E, Rubidge B, Smith RM. 2017. The Rhinesuchidae and early history of the Stereospondyli (Amphibia: Temnospondyli) at the end of the Palaeozoic. Zoological Journal of the Linnean Society, 181(2): 357-384.
• McHugh JB. 2014. Paleohistology and histovariability of the Permian stereospondyl Rhinesuchus. Journal of Vertebrate Paleontology, 34(1): 59-68.
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