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Tetrapod Zoology 3/27/2018 THE AMAZING WORLD OF SALAMANDERS - Scientific American Blog Network Tetrapod Zoology THE AMAZING WORLD OF SALAMANDERS Tet Zoo loves amphibians* (that’s anurans, salamanders, caecilians and their close relatives), and since 2008 I’ve been making a concerted effort to get through all the amphibian groups of the world. By Darren Naish on October 1, 2013 https://blogs.scientificamerican.com/tetrapod-zoology/the-amazing-world-of-salamanders/?print=true 1/29 3/27/2018 THE AMAZING WORLD OF SALAMANDERS - Scientific American Blog Network The salamanders are here. Clockwise from top left: Chinese giant salamander (Andrias davidianus), Jefferson salamander (Ambystoma jeffersonianum), Blue‑spotted salamander (A. laterale), Eastern newt (Notophthalmus viridescens). Andrias photo by Markus Bühler, remainder by Rosemary Mosco, used with permission. Tet Zoo loves amphibians* (that’s anurans, salamanders, caecilians and their close relatives), and since 2008 I’ve been making a concerted effort to get through all the amphibian groups of the world. I’ve failed, and I’m blaming that entirely on the fact that I can’t put the time I need to into blogging. Sigh, always busy spending my time working my backside off for stupid money, sigh… Anyway, back in 2008 I did succeed in producing a couple of articles that review the caudates – the salamanders – of the world. * I have to say at this point that I really prefer to use the term lissamphibian when referring to extant amphibian lineages. Mostly, this is because I somewhat dislike the use of the term ‘amphibian’: its perpetuation makes people think that non-amniote tetrapods – or anamniotes – are all close relatives, all closely related or ancestral to lissamphibians. However, since the term amphibian is in such wide use and is overwhelmingly popular among biologists and conservationists, I’ll stick with it here. Oh, and no, we haven't forgotten you, albanerpetontids (round these parts, they're affectionately known as albies). https://blogs.scientificamerican.com/tetrapod-zoology/the-amazing-world-of-salamanders/?print=true 2/29 3/27/2018 THE AMAZING WORLD OF SALAMANDERS - Scientific American Blog Network Sadly, there aren't that many salamander‑themed books to track down and obtain: here's an (admittedly Eurocentric) collection of volumes from the TetZoo Library. There are about 655 recognised living salamander species, and many aspects of their diversity, biology and behaviour are fascinating. There are (or were) giant salamanders more than 2 m long, tiny, tree-climbing salamanders with prehensile tails, partially herbivorous salamanders that possess beaks, cave-dwelling salamanders, long-bodied, aquatic eel-like salamanders, and armoured salamanders with defensive spikes. Claws, intra-uterine cannibalism, neoteny, the defensive shedding of tails and limbs, weird developmental shifts in digital development, bizarre warning displays, lekking and nuptial dancing, facultative herbivory and fungivory, dedicated cave-dwelling… it’s all here. So shame on you if the thought ever occurred to you that salamanders might be plain or boring. A lot of new stuff on these animals has appeared since 2008, meaning that I’ve had to substantially update my original text. Without further ado, let’s get on with it… The term caudate means ‘possessing a tail’ and, in contrast to anurans and caecilians, salamanders retain the body shape that seems to have been plesiomorphic for amphibians. Having said that, even salamanders exhibit a list of anatomical specialisations that make them unusual compared to other tetrapods: they’ve lost a long list of skull bones that are present in other anamniotes (including the postorbitals, jugals tabular supraoccipital https://blogs.scientificamerican.com/tetrapod-zoology/the-amazing-world-of-salamanders/?print=true 3/29 3/27/2018 THE AMAZING WORLD OF SALAMANDERS - Scientific American Blog Network jugals, tabular, supraoccipital and ectopterygoids), and also lack a middle ear (though they’re not deaf). Bizarrely, salamanders have disproportionately large amounts of DNA. Salamanders are generally Laurasian and the groups that Mudpuppy (Necturus maculosus), one of many salamanders occur on the southern continents that retains external gills and a fully aquatic lifestyle into (there are plethodontids in South adulthood. More on mudpuppies below. Photo by Matt Keevil, America and some salamandrids used with permission. in northern Africa) have only gotten there comparatively recently. An important theme which has cropped up several times independently within different salamander clades is neoteny: the retention of juvenile characters into sexual maturity. It’s been widely suspected that neoteny results in the retention of confusing plesiomorphic characters, and in extensive convergence, so working out the relationships of neotenous salamander clades has been difficult (Wiens et al. 2005). Phylogenetic studies on salamanders have in fact differed pretty radically in the relationships they infer (particularly with respect to the position of sirens), though a rough consensus has emerged. The oldest salamanders The oldest salamanders we know of are Middle Jurassic forms from central Asia (Kokartus honorarius) and England (two species of Marmorerpeton). What we know indicates that these early forms looked superficially like stout- bodied living ones, but they lacked various bony and muscular characters present in the crown-group. The best known early salamander – Karaurus sharovi from the Upper Jurassic of Kazakhstan – has been compared to living mole salamanders https://blogs.scientificamerican.com/tetrapod-zoology/the-amazing-world-of-salamanders/?print=true 4/29 3/27/2018 THE AMAZING WORLD OF SALAMANDERS - Scientific American Blog Network living mole salamanders (ambystomatids) and perhaps lived in a similar manner (Estes 1981). [Adjacent image by David Meloni.] Whether the name Urodela is restricted to the crown-group, with Caudata used for the total-group, or whether the name Skeleton of Karaurus sharovi (total length 20 cm). Caudata is restricted to the crown- Image by Davide Meloni, licensed under Creative group, with Urodela used for the total- Commons Attribution‑Share Alike 2.0 Generic license. group, depends on which authors you pay attention to. Evans & Milner (1996) argued that it makes better sense to use Caudata for the total-group, and this seems to have been mostly followed and is used here. So Marmorerpeton and the karaurids Kokartus and Karaurus are stem-caudates but not urodeles. A few other Mesozoic taxa, like Pangerpeton and Jeholotriton from the Jurassic or Cretaceous of Liaoning Province in China, are regarded by some authors as additional stem-caudates (Wang & Evans 2006). [Image below by internet hero Nobu Tamura.] Life reconstruction of a swimming Chunerpeton, by Nobu Tamura. Licensed under Creative Commons Attribution 3.0 Unported license. https://blogs.scientificamerican.com/tetrapod-zoology/the-amazing-world-of-salamanders/?print=true 5/29 3/27/2018 THE AMAZING WORLD OF SALAMANDERS - Scientific American Blog Network Various fossil salamanders from the Middle and Upper Jurassic seem to be crown- group salamanders: that is, early members of the lineages that include the cryptobranchids (giant salamanders) or hynobiids (Asian salamanders) (together, cryptobranchids and hynobiids are termed cryptobranchoids), or members of Salamandroidea, the 'advanced', internally fertilising salamander clade. Chunerpeton from the Daohugou Beds of Inner Mongolia (Gao & Shubin 2003, Carroll & Zheng 2012), and – according to Carroll & Zheng (2012) – Pangerpeton, Jeholotriton, Regalerpeton, Liaoxitriton and Iridotriton are all Mesozoic cryptobranchoids. If this is correct, and if there’s a sister-group relationship between cryptobranchoids and salamandroids, then members of both of these major salamander clades had appeared before the end of the Jurassic. The recent description of Beiyanerpeton jianpingensis from the Jurassic Tiaojishan Formation (Gao & Shubin 2012) seems to confirm the predicted presence of salamandroids in the Jurassic. Iridotriton hechti from the Morrison Formation has also been interpreted as an early salamandroid (Evans et al. 2005) but this was contested by Gao & Shubin (2012) who found it to be a cryptobranchoid. By the Upper Cretaceous, members of most or all of the living salamander ‘families’ had probably appeared. Cryptobranchoids: giant salamanders and Asiatic salamanders I'm not gonna lie to you: some hynobiids look a bit boring. This is a Fischer's clawed salamander (Onychodactylus fischeri). Image by Pierre Fidenci, licensed under Creative Commons Attribution‑Share Alike 2.5 Generic license. Two particularly ancient groups of salamanders are still around today: the hynobiids and the cryptobranchids. Hynobiids, generally just called Asiatic salamanders, are a poorly known group of about 50 species that occur from Afghanistan and Iran https://blogs.scientificamerican.com/tetrapod-zoology/the-amazing-world-of-salamanders/?print=true 6/29 3/27/2018 THE AMAZING WORLD OF SALAMANDERS - Scientific American Blog Network eastwards to Japan, though between the Miocene and Pleistocene they also occurred in Europe (Venczel 1999). Some hynobiids occur in cold parts of northern Asia and are particularly cold-tolerant, being able to withstand freezing at temperatures below -50°C for months at a time (that is, they literally get frozen alive and can stay dormant in a block of ice). [Image of Onychodactylus above by Pierre Fidenci.] Some hynobiids employ aquatic suction feeding
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