SVP 2018: Thaisaurus, basal ichthyosaur

Liu, Samathi and Chanthasit 2018
study for the first time Thaisaurus (Fig. 1), a basal ichthyosaur in the large reptile tree (LRT, 1315 taxa). We first looked at Thaisaurus in April, 2015 here.

The authors report, “Since its first brief description, however, T. chonglakmanii has never been restudied in detail, and its exact stratigraphic and phylogenetic position remained elusive. Here we revisit the well prepared holotype specimen of T. chonglakmanii.  This is the earliest record of Mesozoic marine reptiles, two million years earlier
than the earliest previous record.” The authors do not record an outgroup for the Ichthyosauria. The LRT provides dozens in a lineage going back to Devonian tetrapods. Late surviving Wumengosaurus nests as the basalmost ichthyosaur in the LRT (Fig. 2) and mesosaurs are the sister clade appearing as early as the Early Permian. So that gives plenty of time for ichthyosaurs to diverge from primitive mesosaur/sauropterygians. And we should be finding basal ichthyosaurs throughout the Permian.

Figure 1. Thaisaurus in situ, traced using DGS, elements of tracing shifted using DGS and restored.

Figure 1. Thaisaurus in situ, traced using DGS, elements of tracing shifted using DGS and restored.

From 2015
Thaisaurus chonglakmanii (Mazin et al. 1991; Early Triassic; Fig. 1.) was considered the most basal ichthyosaur by Maisch (2010). That is largely confirmed in the large reptile tree where Thaisaurus nests between Wumengosaurus and the remainder of the Ichthyosauria (sensu Maisch 2010, Fig. 2).

Figure 3. Aquatic younginiform subset of the LRT demonstrating relationships within the Enaliosauria (=Sauropterygia + Ichthyosauria)

Figure 2. Aquatic younginiform subset of the LRT demonstrating relationships within the Enaliosauria (=Sauropterygia + Ichthyosauria)

Nice to see that everyone is in agreement
on the taxonomic nesting of Thaisaurus.

Figure 2. Click to enlarge. The origin of ichthyosaurs and thalattosaurs from basal diapsids and basal mesosaurs. Relationships are rather apparent when seen in this context.

Figure 2. Click to enlarge. The origin of ichthyosaurs and thalattosaurs from basal diapsids and basal mesosaurs. Relationships are rather apparent when seen in this context.

Thaisaurus was a late-survivor in the Early Triassic,
a time in which ichthyosaurs were diversifying rapidly. Or did ichthyosaurs just appear in the fossil record then, having diversified throughout the Permian?

References
Liu J, Samathi A and Chanthasit P 2018. The earliest ichthyosaur from the middle Lower Triassic of Thailand.
Maisch MW 2010. Phylogeny, systematics, and the origin of the Ichthyosauria – the state of the art. Palaeodiversity 3:151-214.
Mazin J-M et al. 1991. Preliminary description of Thaisaurus chonglakmanii n. g. n. sp. a new ichthyopterygian (Reptilia) from the Early Triassic of Thailand. – Comptes- Rendus des Séances de l’Académie de Sciences Paris, Série II, 313: 1207-1212.

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The ichthyosaur(s) with 4 nostrils: Musicasaurus

Maxwell et al. 2015
described a juvenile ophthalmosaur, Muiscasaurus catheti, from the Early Cretaceous of Columbia, and it had a bony process dividing its naris. Online press (BBC.com) described the specimen as having four nostrils (Fig. 1). It does not really have four nostrils, but wait, there’s more…

Figure 1. Muiscasaurus catheti prior to final prep, final prep and diagram. Naris is highlighted.

Figure 1. Muiscasaurus catheti prior to final prep, final prep and diagram. Naris is highlighted.. Compare to Ophthalmosaurus natans in figure 2.

The BBC site reported, 
“The fossil is of an infant only about 3m long. Adults may have reached 5m.” Maybe it is best described as “immature” or a “juvenile” when it is more than half the adult size. It is certainly not an infant.

“I could tell it was a juvenile based on the size of its eyes relative to the rest of the skull,” says author Erin Maxwell of the Natural History Museum in Stuttgart, Germany. “In reptiles, babies have very big eyes and heads compared to their body.”

Of course
adult ichthyosaurs with exceptionally large eyes, like Ophthalmosaurus (Fig. 2) have been known for over a century. Perhaps Dr. Maxwell was misquoted. That happens. Also when we look at Ophthalmosaurus, it has nearly the same naris shape as seen in Muiscasaurus catheter. 

Figure 2. Two variations on Ophthalmosaurus, both with large eyes and one with a peanut-shaped naris, similar to the four-nostril Muiscasaurus.

Figure 2. Two variations on Ophthalmosaurus, both with large eyes and one with a peanut-shaped naris, similar to the four-nostril Muiscasaurus.

Another news source,
the Ulyanovsk Chronicles, recently published a story and image of another “ichthyosaur with four nostrils,” (Fig. 3) from the Aptian (Early Cretaceous, 120 mya) of Sengileevsky paleontological reserve. The site reported [after Google translation], “A preliminary study of a new Museum exhibit conducted by Valentin Fischer (University of Liege, Belgium), [AND] Maxim Arkhangelsky (Saratov state technical University) showed that he loved aikataulu [referred the specimen to?] (Muiscasaurus).” 

Figure 3. A Russian four-nostril ichthyosaur with the pencil resting in the posterior naris.

Figure 3. A Russian four-nostril ichthyosaur with the pencil resting in the posterior naris.

In this new specimen
the anterior and posterior portions of the naris are more completely divided. I wonder if all ichthyosaurs had such a dual naris in soft tissue, but only in these specimens can we find bony support?

References
Maxwell EE, Dick D, Padilla S and Parra ML 2015. A new ophthalmosaurid ichthyosaur from the Early Cretaceous of Columbia. Papers in Palaeontology 2015:1-12.

Ichthyosaur phylogeny: Ji et al. 2016

Ji et al. 2016
present us with an updated cladogram of ichthyosaur interrelationships. The only problem is they punted on the providing an outgroup. Instead they used several basal diapsids and marine younginiforms with no proto-icchthyosaur traits.

From the Ji et al. 2016 text: 
“We adopted five outgroup taxa from Motani (1999) for character polarization, whereas the position of the Ichthyoptergia within the Amniota is beyond the scope of the current study.”

That’s not the way to start a valid phylogenetic analysis
You really should know what your taxon is before attempting to figure out interrelationships. The authors chose as outgroup taxa:

  1. Petrolacosaurus
  2. Hovasaurus
  3. Claudiosaurus
  4. Thadeosaurus
  5. Hupehsuchus

Unfortunately none of these taxa
are suitable/decent/proximal outgroup taxa for the Ichthyopterygia, according to the large reptile tree. When you don’t have a decent proximal outgroup, how can you know which is the basalmost taxon? And how can you determine a gradual evolution of character traits?

The large reptile tree recovers
the following outgroup taxa in order of increasing distance:

  1. Xinminosaurus
  2. Thaisaurus
  3. Wumengosaurus
  4. the clade Thalattosauria + Mesosauria (Serpianosaurus and Psilotrachelosaurus basal taxa)
  5. the clade Sauropterygia (Diandongosaurus and Pachypleurosaurus basal taxa)
  6. Anarosaurus
Figure 1. Subset of the LRT focusing on the clade Ichthyosauria.

Figure 1. Subset of the LRT focusing on the clade Ichthyosauria.

Where are hupehsuchids in this phylogeny?
As you can tell by looking at hupehsuchids (Fig. 1), they are quite derived, so derived that the authors don’t realize they actually nest within the Ichthyopterygia. In the large reptile tree they are derived from sisters to Wumengosaurus.

Figure 5. Shastasaurus

Figure 5. Shastasaurus

When you apply tested outgroup taxa
then Xinminosaurus no longer nests with Cymbospondlylus. ChaohusaurusCymbospondylus, Mixosaurus and Phalarodon no longer nest as a basal ichthyopterygians.

As with pterosaur workers
it would be nice if ichthyosaur workers would take the time to figure out what ichthyosaurs are. They have been nested with Wumengosaurus, thalattosaurs and mesosaurs since the origin of ReptileEvolution.com in 2011.

On a happier note
Earlier we looked at two odd bedfellow den mates (Fig. 2): an early Triassic amphibian and a cynodont, evidently getting cozy on a sleepover.

The rarest of rare fossils finds: Two more-than-friends having a sleepover. Credit to Fernandez et al. 2013.

Figure 2. The rarest of rare fossils finds: Two more-than-friends having a sleepover. Credit to Fernandez et al. 2013.

A reader suggested
I take a look at this modern equivalent on YouTube (Fig. 3). I encourage you to do the same. I think you’ll enjoy it. Wish I knew about this one on Valentine’s Day.

Video 1. Click to play on YouTube. Puppet the cat and Puff the bearded dragon are evidently soul mates.

Figure 3. Click to play on YouTube. Puppet the cat and Puff the bearded dragon are evidently soul mates.

References
Ji C, Jiang D-Y, Motani R, Rieppel O, Hao W-C and Sun Z-Y 2016. Phylogeny of the Ichthyopterygia incorporating recent discoveries from South China. Journal of Vertebrate Paleontology 36(1):e1025956. doi: http://dx.doi.org/10.1080/02724634.2015.1025956

New ichthyosaur family tree by Ji et al. 2015

A recent paper on ichthyosaur systematics
(Ji et al. 2015, Fig. 1) adds newly discovered taxa and the tree is getting nice and big.

Unfortunately,
at the base of their cladogram Ji et al. place a distinctly different proximal outgroup for ichthyosaurs than what was recovered in the large reptile tree (subset shown in Fig. 1, click to enlarge). They appear to be guessing. Apparently they are not sure how ichthyosaurs are related to other reptiles.

Here 
proximal outgroup taxa for ichthyosaurs include Wumengosaurus, Thaisaurus and Xinminosaurus (in ascending order) not Thadeosaurus. These large reptile tree taxa demonstrate a gradual accumulation of basal ichthyosaur traits. The Ji et al taxa, HovasaurusClaudiosaurus and Thadeosaurus do not. In the large reptile tree these three are basal younginiformes, related, yes, but much more distantly related to ichthyosaurs.

Figure 1. Ichthyosaur family trees compared. Left: subset of the large reptile tree. Right: from Ji et al. 2015. Note the lack of correct outgroups in the Ji et al study. They have no idea which taxa are proximal ancestors.

Figure 1. Click to enlarge. Ichthyosaur family trees compared. Left: subset of the large reptile tree. Right: from Ji et al. 2015. Note the lack of correct outgroups in the Ji et al study. They have no idea which taxa are proximal ancestors. Yellow are taxa found in both trees.

Figure 1. Subset of the LRT focusing on the clade Ichthyosauria.

Figure 1. Subset of the LRT focusing on the clade Ichthyosauria updated November 4, 2018 with a shift of the Hupehsuchidae closer to the base of the Ichthyosauria.

So, as an experiment, 
we’ll delete the large reptile tree proximal outgroup taxa in order to match more closely the Ji et al taxon list. What is recovered now?

  1. Hovasaurus, Claudiosaurus and Thadeosaurus now nest together in an outgroup clade.
  2. Hupehsuchus + Xinminosaurus, Grippia and (Utatsusaurus + (Shastasaurus  pacificus + Shastasaurus alexandrae) now form clades at the base of the Ichythyosauria.
  3. Then Chaohusaurus nests at the base of the rest of the Ichthyosauria with the same topology as the subset of the large reptile tree.

A few differences between the two topologies without deletions…
Note the morphological mismatches in the Ji et al. topology not found in the large reptile tree.

  1. In the large reptile tree Chaohusaurus nests between two similar taxa, Parvinatator and Besanosaurus. In the Ji et al. tree Chaohusaurus nests between the mismatched and odd Hupehsuchus and a clade of basal ichthyosaurs as the basalmost ichthyosaur, even though it has a derived ichthyosaur shape and traits.
  2. In the large reptile tree the derived, but still Triassic, Cymbospondylus petrinus nests between its contemporary, Mixosaurus and several other giant serpentine ichthyosaurs. All have a depressed cranium with a central ridge. The unrelated flat-headed C. buchseri nests elsewhere with similar deep-bodied, high-crested Shonisaurus popularis. By contrast, in the Ji et al. tree C. piscosus (= petrinus) and C. buchseri nest together with the very primitive, very small, Xinminosaurus, which does not have such a depressed cranium with a central crest.
  3. Ji et al. have a clade of Shastasauridae that includes only shastasaurs. In the large reptile tree, that clade also includes the odd little hupehsuchids and demonstrates how these little toothless enigmas evolved from larger forbearers. Ji et al. provided several skull reconstructions. Perhaps a few more would help to resolve the distinct topologies.

Those are the major issues.
The rest can be swept up later. I’d like to see the authors either expand their own taxon list or work off the large reptile tree to confidently establish a series of outgroup taxa for the Ichthyosauria that actually demonstrate a gradual accumulation of character traits, instead of doing what they did. Then we might have closer correspondence in tree topology. And we’re going to have to figure out Cymbospondylus… is it derived? or primitive?

References
Ji C, Jiang D-Y,  Motani R, Rieppel O, Hao -C & Sun Z-Y 2015. Phylogeny of the Ichthyopterygia incorporating recent discoveries from South China, Journal of Vertebrate Paleontology, DOI: 10.1080/02724634.2015.1025956

Unexpected centralia in the ichthyosaur, Chaohusaurus

A recent paper
by Montani et al. 2015 purported to indicate the presence of two centralia in the wrist of a juvenile basal ichthyosaur, Chaohusaurus (AGM CH-6628-22). It was not present in the adult. That, on its face of it is odd. The specimen, despite appearing to be undisturbed, lacked an ulna. That is also odd. Finally, no sister taxa have centralia. So the appearance here (Fig. 1) is triply odd.

Figure 1. Reinterpretation of Motani et al. 2015 showing how the purported radius could be a radius and ulna overlapping, the lateral centrale (lc) could be distal tarsals 3 and 4, and the medial centrale (mc) could be m4.2. Metacarpal "0" could be a part of the ulna. No sister taxa have centralia.

Figure 1. Reinterpretation of Motani et al. 2015 showing how the purported radius could be a radius and ulna overlapping, the lateral centrale (lc) could be distal tarsals 3 and 4, and the medial centrale (mc) could be m4.2. Metacarpal “0” could be a part of the ulna. No sister taxa have centralia. Click to enlarge.

Montani et al. report. 
“no amphibious sister taxa to ichthyopterygians have been discovered so far.”

Not so.
For the last four years the large reptile tree lists many sister taxa of increasing distance to the Ichthyopterygia, beginning with Wumengosaurus, basal mesosaurs and the several pachypleurosaurs that led to these taxa. The centralia is absent over several nodes prior to their appearance.

Centralia last appear
in Claudiosaurus, Adelosaurus, Sinosaurosphargis and Largocephalosaurus, but not thereafter. As the Enaliosauria becomes more aquatic, carpals are lost, beginning with the two centralia. Pachypleurosaurs do not have centralia. Neither do mesosaurs.

The (AGM CH-6628-22) specimen
of a juvenile Chaohusaurus that Motani et al. believe to have centralia (Fig. 1) has widely spaced and largely cartilaginous (poorly ossified) elements, some of which, like the ulnare and radiale are clearly disturbed from their in vivo placements. There is a long bone they label the lateral centrale and a short bone they label a medial centrale where such bones belong and this is the basis for their claim. There is even a medial metacarpal “0”, which anchors a sixth medial digit in the related Hupehsuchus, but is not known in Chaohusaurus, which is not a basal ichthyosaur.

We saw a similar reappearance
of digit “0” in Limusaurus, a theropod with embryonic hands retained into adulthood. Claudiosaurus, Adelosaurus and Sinosaurosphargis have a pisiform lateral to the ulnare, but it is similar in size and shape to the ulnare. More derived enaliosaurs lack a pisiform along with the centralia.

The problem is,
the lateral centrale in the above named enaliosaurs is not elongated (double wide), as it is purported to be in the juvenile Chaohusaurus, but rounded and similar in size and shape to the medial centrale.

A solution:
In my experience with missing bones alongside extra bones the answer might be to reinterpret the extra bones as the missing bones, only displaced. Perhaps the ulna is not missing from the juvenile Chaohusaurus, but instead is resting partly atop the radius (Fig. 1) as it appears to do so with that white line dividing the pair. If so the ulna can be restored, but the medial part is damaged. Here the purported metacarpal “0” might be part of the ulna. That’s a guess. The lateral centrale might be distal tarsals 3 and 4, as in the adult “D” specimen (Fig. 1), double wide. The purported distal tarsal 4 then is reinterpreted as distal tarsal 2 here. The medial centrale is reinterpreted as m4.2, which is also missing. The new restoration more closely matches adult specimens and sister taxa. It would also be nice if somehow we could determine that more radius was hidden beneath that portion of the displaced ulna.

This is parsimony and phylogenetic bracketing at its best.
If correct, this scenario just requires you to accept that a certain amount of displacement occurred during taphonomy, as in our old friend, Sordes, the pterosaur. I have not seen the specimen. So, if correct, this is another example of DGS, digital graphic segregation, and an example of pulling more data out of a photograph than was pulled out with the specimen in hand.

References:
Motani R et al. 2015. New evidence of centralia in Ichthyopterygia reiterating bias from paedomorphic characters on marine reptile phylogenetic reconstruction. Journal of Vertebrate Paleontology. 6 pp.

Hauffiopteryx (BRLSI M1399): a CT-scanned Jurassic ichthyosaur skull

Figure 1. BRLSI M1399 is a new ichthyosaur that has been subjected to CT scanning and colorizing. It had huge eyeballs evidently not spherical in shape (there was no room in the skull). The original paper did not put the palate together. That is remedied here. Click to enlarge.

Figure 1. Hauffiopteryx, BRLSI M1399, is a new ichthyosaur that has been subjected to CT scanning and colorizing. It had huge eyeballs evidently not spherical in shape (there was no room in the skull). The original paper did not put the palate together or separate the posterior mandibles. Those are remedied here. At lower left are hypothetical eyeballs. A short F-stop is ideal for light gathering. Click to enlarge.

A new ichthyosaur, Hauffiopteryx, has been CT scanned.
You can see a rotating image of that Marek et al. (2015) scan here.

From the abstract: “New information on the braincase, palate and occiput are provided from three-dimensional scans of an exceptionally preserved ichthyosaur (‘Hauffiopteryx’ typicus) skull from the Toarcian (183–174 Ma, Lower Jurassic) of Strawberry Bank, England. This ichthyosaur has unusual, hollow, tubular hyoid bars. The occipital and braincase region is fully reconstructed, creating the first digital cranial endocast of an ichthyosaur. Enlarged optic lobes and an enlarged cerebellum suggest neuroanatomical adaptations that allowed it to be a highly mobile, visual predator. The olfactory region also appears to be enlarged, suggesting that olfaction was more important for ichthyosaurs than has been assumed. Phylogenetic analysis suggests this ichthyosaur is closely related to, but distinct from, Hauffiopteryx, and positioned within Thunnosauria, a more derived position than previously recovered. These results further our knowledge of ichthyosaur cranial anatomy in three dimensions and provide a platform in which to study the anatomical adaptations that allowed ichthyosaurs to dominate the marine realm during the Mesozoic.”

Figure 2. From Marek et al. (2015), a cladogram of the higher ichthyosaurs. Pink arrow points to Eurhinosaurus and Leptonectes where Hauffiopteryx nests when the more derived taxa are not included on the large reptile tree.

Figure 2. From Marek et al. (2015), a cladogram of the higher ichthyosaurs. Pink arrow points to Eurhinosaurus and Leptonectes where Hauffiopteryx nests when the more derived taxa are not included on the large reptile tree.

The authors report, “Most post-Triassic ichthyosaurs belong to the clade Thunnosauria, with Hauffiopteryx typicus recovered as the immediate out-group to this clade (Fischer et al. 2013). Therefore, this species is an important marker in the transition to the great majority of advanced ichthyosaurs.”

Figure 2. Subset of the large reptile tree focusing on ichthyosaurs. Note most of the more derived ichthyosaurs from Marek et al. 2015, are not listed here. So we're not comparing apples to apples here.

Figure 3. Subset of the large reptile tree focusing on ichthyosaurs. Note most of the more derived ichthyosaurs from Marek et al. 2015 (Fig. 2), are not listed here. So we’re not comparing apples to apples here.

The authors further report, “Most Lower Jurassic ichthyosaur specimens are preserved in flattened and compressed form. This is especially true of exceptionally preserved specimens from Holzmaden, southern Germany (Toarcian, Lower Jurassic), which may show soft tissues and body outlines, but the skeletons are flattened and conceal details, especially within the skull. Other ichthyosaurs may be three dimensional, but disarticulated.”

Figure 4. A more complete but crushed specimen of Hauffiopteryx along with tracings and reconstructions of key parts.

Figure 4. A more complete but crushed specimen of Hauffiopteryx along with tracings and reconstructions of key parts. Click to enlarge. Black hand bones are metacarpals. Note the differences in maxilla length. The 3D specimen appears to have a shorter maxilla no further forward than the naris, unlike the crushed specimen or Eurhinosaurus. Two species of Ophthalmosaurus show the same sort of variation.

Both specimens
of Hauffiopteryx have a box-like cranium housing huge eyes along with a small, sharp rostrum. Ophthalmosaurus, Leptonectes and Eurhinosaurus (Fig. 6) more or less share these traits and, give the taxon list of the large reptile tree, they all nest together. This may change with the addition of more taxa, as shown in figure 2.

The lacrimal question
In the CT scanned specimen (Fig.1) a slender bone extends along the ventral naris and extends slightly outside of it. In the crushed specimen (Fig. 2) the area ventral to the naris is crushed and broken. In sister taxa the lacrimal extends along the lower rim of the naris, but it was not colorized that way in figure 1. So I wonder about it.

The maxilla question
In the 3D specimen (Fig. 1) the yellow maxilla does not extend anteriorly beyond the large narrow naris. That’s not the case in the crushed specimen or Eurhinosaurus. Similarly in various species of Ophthalmosaurus the maxilla may be long or short. In the 3D specimen (Figs. 1, 5) there is a depression aligned with what would have been the pmx/mx suture. So I wonder if part of the maxilla in the 3D specimen was improperly colorized originally?The tiny teeth at the anterior of the possible maxilla suggest that may be the actual maxilla Marek et al. may have misidentified a splintered break as a suture.

Figure 5. The disputed maxilla in BRLSI M1399. Marek et al. colorized the maxilla only to the anterior naris, but that might be a break. Some sister taxa extend the maxilla beyond the the naris and the tiny teeth at the thin anterior of the new maxilla both indicate a possible error was made, mistaking a break for a suture. If valid, this is what DGS can do. Click to enlarge.

Figure 5. The disputed maxilla in BRLSI M1399. Marek et al. colorized the maxilla only to the anterior naris, but that might be a break. Some sister taxa extend the maxilla beyond the the naris and the tiny teeth at the thin anterior of the new maxilla both indicate a possible error was made, mistaking a break for a suture. If valid, this is what DGS can do. Click to enlarge.

If the traits identified here are valid, Hauffiopteryx and its new sister are closer to Eurhinosaurus (Fig. 6) than Marek et al. nested them. Though relatively smaller, the crescent-shaped tail of the crushed Hauffiopteryx (Fig. 4) is also similar to that of Eurhinosaurus (Fig. 6).

Figure 1. Eurhinosaurus, a derived ichthyosaur, in several views.

Figure 6. Eurhinosaurus, a derived ichthyosaur, in several views.

 

References
Marek RD, Moon BC, Wiliams M and Benton MJ 2015. The skull and endocranium of a Lower Jurassic ichthyosaur base on digital reconstructions. Palaeontology 2015: 1-20.

 

 

Flipper size in Mesozoic Enaliosaurs.

Earlier we looked at three tylosaurs distinguished by their widely varying flipper size. Today we’ll do the same with a few closely related pliosaurs (Fig. 1) and ichthyosaurs (Fig. 2).

Figure 1. Three pliosaurs, Trinacromerum, Brachauchenius and Kronosaurus, to scale, all Late Cretaceous.

Figure 1. Three pliosaurs, Trinacromerum, Brachauchenius and Kronosaurus, to scale, all Late Cretaceous. Variations in flipper size, rib count and girdle size mark the major differences here. The girdles anchored large ventral swimming muscles. Scale bar = 1 meter. How flipper size affected speed and agility is a topic that has not been brought up yet, as far as I know.

Pliosaurs were some of the largest marine reptiles of all time.
They reached an acme in the Late Cretaceous with a variety of morphologies derived from smaller ancestors. Shown here are Trinacromerum, Brachauchenius and Kronosaurus, three taxa rarely shown together to scale. The big difference is in flipper size. It’s also worth noting the size of the girdles anchoring each flipper set. In whales, flipper size also varies greatly and not always for swimming advantage.

Ichthyosaurs,
even closely related ones (Fig. 2), also had widely varying flipper sizes. Here we see ‘Cymbospondylus’ buchseri and the related Guizhouichthyosaurus. Both were probably sinuoous swimmers, rather than tail or flipper swimmers.

Figure 2. Two closely related ichthyosaurs, Guizhouichthyosaurus tangae and "Cymbospondylus" buchseri, one with large flippers, one with small.

Figure 2. Two closely related ichthyosaurs, Guizhouichthyosaurus tangae and “Cymbospondylus” buchseri, one with large flippers, one with small.

References
Carpenter K 1996. A review of short-necked plesiosaurs of the Western Interior, North America. Neues Jahrbuch fur Geologie und Palaontologie, Abhandlungen 201(2):259-287.
Hampe O 2005. Considerations on a Brachauchenius skeleton (Pliosauroidea) from the lower Paja Formation (late Barremian) of Villa de Leyva area (Colombia). Fossil Record – Mitteilungen aus dem Museum für Naturkunde in Berlin 8 (1): 37-51.
Longman HA 1924. A new gigantic marine reptile from the Queensland Cretaceous, Kronosaurus queenslandicus new genus and species. Memoirs of the Queensland Museum 8: 26–28.
O’Keefe FR 2001. A cladistic analysis and taxonomic revision of the Plesiosauria (Reptilia: Sauropterygia). Acta Zoologica Fennica 213:1-63.
Owen R 1840. British Fossil Reptiles. Chapter V Order—Sauropterygia, Owen. Genus—Pliosaurus, Owen. 152-165.
Romer AS and Lewis AD 1959. A mounted skeleton of the giant plesiosaur Kronosaurus. Breviora 112: 1-15.
Williston SW 1903. North American plesiosaurs. Field Columbian Museum, Pub. 73, Geological Series 2 (1):1-79.
Williston SW 1907. The skull of Brachauchenius, with special observations on the relationships of the plesiosaurs. United States National Museum Proceedings 32: 477-489.
Williston SW 1908. North American Plesiosaurs: Trinacromerum. Journal of Geology 16(8): http://www.jstor.org/stable/30068152

wiki/Brachauchenius
wiki/Pliosaurus

wiki/Kronosaurus
wiki/Trinacromerum