Traditionally, questions still stir
about the origin of the clade Ichthyosauria and about interrelationships between clade members (Fig. 1). Wikipedia offers several variations of interrelations while reporting, “The origin of the ichthyosaurs is contentious.”
Those questions do not stir
in the large reptile tree (LRT, 1327 taxa, subset Fig. 1) which confidently nests ichthyosaurs arising from Wumengosaurus, mesosaurs + thalattosaurs and basal sauropterygians in order of increasing distance. All candidate taxa are tested here, minimizing uncertainty, increasing confidence. Taxon exclusion is the chief problem robbing paleontologists of the joy of knowing where ichthyosaurs come from. For some reason, Wumengosaurus never seems to make the ichthyosaur inclusion list. Wikipedia reports, “It is unknown exactly what Hupehsuchus is related to.”
The ichthyosaur mimics, Cartorhynchus and Sclerocomus, nest not as a basal ichthyosaurs, but as basal sauropterygians (contra Motani et al. 2014 and Jiang et al. 2016) in the LRT. More details here.
Wumengosaurus (Fig. 2) has been nested as the basalmost ichthyosaur here since 2011. Hupehsuchids and Thaisaurus have been linked to ichthyosaurs from their discovery on, but similar-looking outgroup taxa, like Wumengosaurus and mesosaurs, have been ignored for reasons unknown. Tall dorsal vertebrae topped by spine tables help link these taxa together.
Grippia and Late Triassic Mikadocephalus (Fig. 4) become more dolphin-like. Tiny Early Triassic Parvinatator and Chaohusaurus show that Early Triassic ichthyosaurs had already radiated widely having split from mesosaurs in the Early Permian. At present, no Permian ichthyosaurs are known, but someday they will be discovered.
Phalarodon and Contectopalatus developed a long narrow rostrum without elongating the limbs (Fig. 4). Moreover, the dorsal cranium develops large ridges, anchors for powerful jaw muscles not seen in prior taxa with flat-top skulls.
The Middle Triassic
sub-meter-long Mixosaurus had a semi-dolphin-like body that became more elongated with longer necks, tiny teeth and giant descendants like Middle Triassic Cymbospondylus and even more gigantic (20m) toothless Late Triassic Shonisaurus sikanniensis (Fig. 5, distinct from Shonisaurus popularis, Fig. 7). Late Triassic, short-snouted, toothless 10m long Guanlingsaurus had an odd twice-as-wide-as-all skull.
Speedy dolphin-like and dolphin-to-killer-whale-sized ichthyosaurs
like Wimanius, Platypterygius and Guizhouichthyosaurus (Fig. 6) split off next. Middle Triassic ‘Cymbospondylus’ buchseri with a 90cm long skull is basal to Late Triassic Shonisaurus popularis with a 3m skull and a 15m overall length with elongate flippers and a much longer rostrum.
This is another clade of increasingly gigantic taxa,
but only the basal taxa, like Platypterygius, survived past the end of the Triassic.
The final clade of ichthyosaurs: Thunnosauria
are truly highly derived, dolphin-like and the speedsters of the Ichthyosauria. These start with Icthyosaurus and continue through Ophthalmosaurus, Leptonectes, tiny Hauffiopteryx and swordfish-like Eurhinosaurus with the longest rostrum and control surfaces in the Ichthyosauria (Fig. 8). This clade also had the largest eye/cranium ratios.
Compare this cladogram
(Fig. 1) of ichthyosaurs to competing cladograms (the latest, so far as I have found, is Moon 2017) and see if the others provide the gradual accumulation of traits shown here. That’s how you know, after all the scores have been entered, if the cladogram makes sense. (And check to see if any include Wumengosaurus. Ji et al. 2016 does not.)
Moon et al. 2017 ran the most recent analysis
of the Ichthyosauria. Unequivocally resolved clades include Ichthyopterygia, Ichthyosauria, Shastasauria, Euichthyosauria, Parvipelvia and Neoichthyosauria, but with variation in their taxonomic components. Mixosauridae and Ophthalmosauridae are similarly recovered, but their definitions are modified to stem-based definitions to prevent substantial variation of included taxa. Several genera are not monophyletic in Moon et al.: Brachypterygius, Leptonectes, Mixosaurus, Ophthalmosaurus, Paraophthalmosaurus, Phalarodon, Platypterygius, Stenopterygius, Temnodontosaurus and Undorosaurus. Moon et al. conclude: “Complex and variable relationships suggest the need for new characters and a re-evaluation of the state of ichthyosaur phylogenetics.”
Here are a few current ichthyosaur clades and their definitions
with comments regarding their validity and membership in the LRT.
- Ichthyosauromorpha – The last common ancestor (LCA) of Ichthyosaurus + Hupesuchia and all descendants. That LCA in the LRT is Wumengosaurus.
- Ichthyosauriformes – All ichthyosauromorphs closer to Ichthyosaurus than to Hupehsuchus. Whenever Cartorhynchus is a basal member, as it is within the current definition, this clade is a junior synonym for Enaliosauria in the LRT. That LCA in the LRT is Thaisaurus (and kin).
- Ichthyopterygia – last common ancestor of Ichthyosaurus, Utatsusaurus and Parvinatator (Motani 1999). That LCA in the LRT and Motani 1999 is Utatsusaurus.
- Eoichthyosauria – The LCA of Grippia and Ichthyosaurus (Motani 1999). That LCA in the LRT and Motani 1999 the LCA is Grippia.
- Ichthyosauria – all eoichthyosaurs more closely related to Ichthyosaurus than to Grippia (Motani 1999). That LCA in the LRT is Mikadocephalus (and kin). That LCA is Cymbospondylus (both species) in Motani 1999,
- Merriamosauria – The LCA of Shastasaurus and Ichthyosaurus. That LCA in the LRT is Utatsusaurus (junior synonym of Ichthyopterygia). That LCA in Motani 1999 is Shastasaurus.
- Shastasauria – All merriamosaurs more closely related to Shastasaurus than to Ichthyosaurus. In Motani 1999, this clade includes Besanosaurus, Shonisaurus and Shastasaurus.
- Parvipelvia – the LCA of Hudsonelpidia, Macgowania, Ichthyosaurus and all of its descendants.
- Thunnosauria – the LCA of Ichthyosaurus communis and Stenopterygius quadriscissus and all of its descendants
- Eurhinosauria – The LCA of Eurhinosaurus and Leptonectes (Motani 1999). The LCA in the LRT is Leptonectes. In Motani 1999 this clade nests outside the Thunnosauria.
2016. Phylogeny of the Ichthyopterygia incorporating recent discoveries from South China, Journal of Vertebrate Paleontology, 36:1, Maisch MW 2010. Phylogeny, systematics, and origin of the Ichthyosauria – the state of the art. Palaeodiversity 3:151-214.
Moon BC 2017. A new phylogeny of ichthyosaurs (Reptilia: Diapsida). Journal of Systematic Palaeontology
Motani R 1999. Phylogeny of the Ichthyopterygia. Journal of Vertebrate Paleontology 19:3, 473-496,
Motani R et al. 2014. A basal ichthyosauriform with a short snout from the Lower Triassic of China. Nature doi:10.1038/nature13866