Friday, November 27, 2020

Your Friends The Titanosaurs, part 30 continued: the rest of Titanosaurus

I hope you've saved room for Round II! This time we visit three continents and most of the Cretaceous as we finish off the rest of the species attributed to Titanosaurus. Our guests are "Titanosaurus" falloti, "T." lydekkeri, "T." nanus, and "T." valdensis. Not all of them are still thought to be titanosaurs, and all four are mostly footnotes these days.

Sunday, November 22, 2020

Your Friends The Titanosaurs, part 30: Titanosaurus of India and Madagascar

My original plan was to cover Titanosaurus indicus plus the seven species that hadn't been otherwise included; overstuffed for Thanksgiving, if you like. After 4,500 words, not counting the 46 references, I thought that might be a little too overstuffed. So, I've decided to split the post in two. Today covers just T. indicus and the three other dubious species from India or Madagascar ("T." blanfordi, "T." madagascariensis, and "T." rahioliensis). Later this week, I'll put up the other half, which will take us to Argentina ("T." nanus), England ("T." lydekkeri and "T." valdensis), and Laos ("T." falloti). As usual, house rules apply: indeterminate species have gone back to their original genus and get quotation marks.

A few general remarks before we get started: Titanosaurus is notorious for the numerous far-flung species that have been attached to it (Wilson and Upchurch 2003). We've already seen the cream of the crop, the species which are still considered to be valid or potentially valid, albeit under different genera: Titanosaurus australis (Neuquensaurus), Titanosaurus colberti (Isisaurus), Titanosaurus dacus (Magyarosaurus), and Titanosaurus robustus (Neuquensaurus), plus Laplatasaurus araukanicus, which has sometimes been considered a species of Titanosaurus (e.g., Powell 2003). Let us not forget Titanosaurus montanus either, although technically it doesn't count because it was supposed to be a different genus; O. C. Marsh just came up with the idea of naming a sauropod Titanosaurus the same year that Richard Lydekker did. Anyway, this leaves us with the type species T. indicus and seven dubious species, not all of which are necessarily even titanosaurs. This doesn't get into another pool of fossils identified as Titanosaurus sp., but these are being left out here. If you'd like to delve into those, Wilson and Upchurch (2003) also address them. (One of the examples they address, also known as DGM Series C, later became Baurutitan britoi.)

Why were so many species attributed to Titanosaurus? My guess is there were several factors. First, paleontologists of the 19th century and early 20th century were more apt to add new species to existing genera than paleontologists today. Second, there wasn't much known about sauropods at the time, and almost all of what was known pertained to North American Jurassic sauropods. Third, Titanosaurus indicus is a blank canvas. Outside of characteristics that are now known to be widely distributed among titanosaurs, such as procoelous caudals, there wasn't much to define it. The great majority of the other Titanosaurus species are based on pretty scrappy material. Once the ball got rolling, the definition of Titanosaurus began to stretch and grow to accommodate the various new species, creating a positive feedback loop.

Sunday, November 15, 2020

How to complicate the Glenwood

Recently I've been doing some work with geological stratotypes in National Park Service areas. The naming of geological formations is not unlike naming a fossil species of organism (or living species for that matter), except the type locality is also the type "specimen", and you don't dig up the type locality and put it in a museum (although you could certainly take a core). Anyway, there are a few stratotypes located within Mississippi National River and Recreation Area. I've alluded a couple of times to the type locality of the St. Peter Sandstone being the bluff under Fort Snelling. This locality is no longer accessible due to protective measures, but the formation *is* exposed in the immediate vicinity. The type locality of the Hidden Falls Member of the Platteville is in Hidden Falls Park (Sloan 1956). Sloan didn't state specifically where, but I have a pretty good idea. Finally, there are two units named from the section exposed at Lock and Dam No. 1.

Beautiful exposure, even if you can't climb around on it.

The Twin Cities Basin is just a small part of the area where the St. Peter–Glenwood–Platteville–Decorah sequence is exposed, and across this area, the rocks differ and the names differ. Reconciling nomenclature across states is part of what Mossler (2008) was about. An example of outstate attention on the Twin Cities Basin is Templeton and Willman (1963), from the Illinois Geological Survey. This publication brings the Illinois nomenclature into the Twin Cities, via a section at Lock and Dam No. 1 (p. 226–227). If you're used to the local names, the Lock and Dam section requires a lot of translation. For example, the Platteville is considered a group, divided into three formations and nine members. The Templeton and Willman nomenclature has not taken off in Minnesota, which has preferred a simpler system, and I can't say I disagree. It seems like overkill to pack that many formal divisions into about 9 vertical meters (30 ft), many of which are not visually distinguishable at a distance of a few meters. Their divisions of the transition from the St. Peter to the Platteville are another matter.

Beginning with our old friend the Pecatonica and going down, Templeton and Willman (1963) divided the Lock and Dam rocks into the Chana Member and Hennepin Member of the Pecatonica Formation (Platteville Group), the Harmony Hill Member and Nokomis Member of the Glenwood Formation, and the St. Peter Sandstone. Two of these members were named from this section: the Hennepin Member and Nokomis Member. In turn:

My initial thoughts. Arrows point to the approximate center of the named unit, so I don't have to commit to contacts like the coward and cad I am. The scale boxes are *very* rough; I was able to put a scale bar on the outer wall, but this is some distance back, so prefer a larger number and don't take it too seriously.

...Or is this what Templeton and Willman intended? Might be easier if I could get closer.

The Chana Member, 28 cm thick (11 in), is basically what we recognize as the Pecatonica;

The Hennepin Member is 69 cm (27 in) thick, divided into three parts: 46 cm (18 in) of clay-rich greenish-gray limestone, and green shale, over 15 cm (6 in) of clay-rich greenish-gray slightly sandy limestone, over 8 cm (3 in) of brown dolomitic sandstone (Templeton and Willman 1963). This unit, or at least the upper part of it, can be easily distinguished visually beneath the blocky overlying rocks (pers. obs.; see also the photos at the end of this post—areas that look white are weathered). The other two parts I'm not as sure about. The Hennepin Member should be thicker than the Chana and the underlying Harmony Hill put together, but from my photos and observations, I see four distinct units over whitish sandstone. Three of them seem fairly similar in thickness: the classic Minnesota Pecatonica, a light-colored recessive unit, and a greenish-brown recessive unit. Below them is a thinner, even more recessive yellow-brown unit. My first thought was that the greenish recessive unit was the Harmony Hill Member, but if I put it in the Hennepin Member and assume that the 15 cm and 8 cm beds of Templeton and Willman (1963) are both included in it, we get something much closer to the overall relative thicknesses T&W found for the Hennepin versus the Chana. Plus, the underlying thin yellow-brown interval looks to be closer to their description of the Harmony Hill Member. The internal proportions are still out of whack, though (three roughly equal units in photos versus 28 cm over 46 cm over 15+8 cm in the publication). (In case you were wondering, there is a grand total of zero [0] photos of outcrops in T&W '63, which is regrettable. A single decent photo with a couple of arrows would have helped immensely.) At any rate, the Hennepin Member is more shaley than the description in T&W '63 implies, or, rather, is less carbonate-rich (Mossler 2008). The Minnesota Geological Survey regards it as the upper part of the Glenwood Formation, rather than the basal part of the Platteville (Mossler 2008);

The Harmony Hill Member is 23 cm (9 in) of yellow-green shale (Templeton and Willman 1963);

The Nokomis Member is also tricky. Templeton and Willman (1963) described it as 330 cm (110 in) thick, divided into 10 to 18 cm (4 to 7 in) of silty white sandstone, over 58 cm (23 in) of very silty, greenish-buff to red-brown, thin-bedded, partly ferruginous sandstone, over 224 cm (78 in) of very silty white to yellow-buff sandstone. Mossler (2008) noted that an interval of silty sand is commonly found in Minnesota between standard Glenwood and standard St. Peter, and this interval has frequently been included in the Glenwood. (The Glenwood is a bigger deal outside of Minnesota.) However, this interval is difficult to distinguish from standard St. Peter in well logs and natural gamma logs, so for practical purposes Mossler (2008) recommended including it in the St. Peter. It is also difficult to identify a difference just by looking at the outcrop from the vantage point of the landing. The best I can do is identify a couple of slope breaks that may indicate changes in mineralogy which are not otherwise apparent from color or bedding at that distance. A contact in the vicinity of the lower slope break would give a thickness in the vicinity of the quoted figure.

The breaks are easier to see in oblique view than straight on. Note also how plants like to colonize the Glenwood–upper Nokomis interval.


Mossler, J. H. 2008. Paleozoic stratigraphic nomenclature for Minnesota. Minnesota Geological Survey, St. Paul, Minnesota. Report of Investigations 65.

Sloan, R. E. 1956. Hidden Falls Member of Platteville Formation, Minnesota. Bulletin of the American Association of Petroleum Geologists 40(12):2955–2956.

Templeton, J. S., and H. B. Willman. 1963. Champlainian Series (Middle Ordovician) in Illinois. Illinois Geological Survey, Champaign, Illinois. Bulletin 89. [large file]

Sunday, November 1, 2020

Your Friends The Titanosaurs, part 29.5: Bravasaurus and Punatitan

No sooner do I announce that Titanosaurus is up next for the titanosaurs than a couple of gate-crashers show up to butt in line: Bravasaurus arrierosorum and Punatitan coughlini. These two taxa were described in the same paper (Hechenleitner et al. 2020) and from the same locality (Quebrada de Santo Domingo) and stratigraphic unit (Ciénaga del Río Huaco Formation), albeit from different levels. They are from Argentina, but not Patagonia; instead, they are from the western, Andean part of the northwestern province of La Rioja, becoming the first named titanosaurs from this province. They are also the first titanosaurs named from the Ciénaga del Río Huaco Formation. The age of this formation has not yet been tightly constrained but is regarded as Campanian–Maastrichtian. In addition to the titanosaur bones, at least three laterally extensive horizons of eggs attributed to titanosaurs were found, between the two bone-bearing levels.

Part of Figure 1 from Hechenleitner et al. (2020), featuring reconstructions of Punatitan coughlini (c) and Bravasaurus arrierosorum (d), with red highlights showing known fossils (scale bar = 1 m). CC-BY-4.0.

Sunday, October 25, 2020

Hyoliths V: The Hyolith Frontier

Hyolith relationships and paleobiology continue to be active research topics, which isn't bad for animals that had their heyday more than 500 million years ago and are primarily known from diminutive shells. Let's get right down to the issue that has haunted study of Hyolitha, as posed in the title of Smith (2020): "Finding a home for hyoliths". For a couple of years recently, they'd been drifting into the welcoming arms of brachiopods and other lophophores (Moysiuk et al. 2017; Sun et al. 2018), as discussed in previous entries. However, cracks quickly began to emerge. A group of authors has published several papers challenging a close relationship with lophophores, instead emphasizing shell structural similarities with mollusks (Li et al. 2019, 2020) or being a bit more conservative and placing them nearer the base of Lophotrochozoa (lophophores, annelid worms, mollusks, etc.; Liu et al. 2020a).

Liu et al. (2020a) questioned the evidence presented by Moysiuk et al. (2017) and Sun et al. (2018). Liu et al. interpreted the putative pedicles of Sun et al. (2018) as crushing of the tip of the shell, with no evidence for a pedicle or an opening for one in intact hyolith shells. They concurred with Moysiuk et al. (2017) that hyoliths had a tentaculate feeding structure, but they did not regard this as homologous with a true lophophore. Smith (2020), in a commentary on Liu et al (2020a), illustrated two potential options: that hyoliths are closer to the mollusks and annelids, in which case the tentaculate feeding apparatuses of hyoliths and lophophores are not related but shells are an ancestral trait of lophotrochozoans (which also means that the Cambrian record of these groups is more complete than otherwise thought); or that hyoliths are closer to brachiopods, making the shell something that appears multiple times while limiting the tentaculate apparatus to the brachiopod line. (I suppose there could be an option where shells are ancestral *and* the hyolith tentacles are related to the lophophore feeding structure, but this wasn't explored.)

Dueling options for hyolith relationships, from Smith (2020). CC-BY-4.0.

Another area of interest in recent years has been the feeding methods and internal anatomy, bolstered by Cambrian fossils with soft tissue traces. Liu et al. (2020a, 2020b) have documented a tentaculate feeding apparatus in a second hyolith genus, the orthothecid Triplicatella. This hyolith had a more "tuft"-like array of tentacles than the broader "gull-wing" spread of the hyolithid Haplophrentis, and like other orthothecids did not have helens (long curved rods; see Skovsted et al. 2020 for the latest on helens). Liu et al. proposed that Triplicatella was not a filter/suspension feeder, as proposed for Haplophrentis, but was instead a stationary deposit feeder, indicating distinct lifestyles for the orthothecid and hyolithid orders of hyoliths.

Interpretation of the internal anatomy of Triplicatella, an orthothecid (so no helens; compare to Figure 7 in Liu et al. 2020c). Figure 8 in Liu et al. (2020b). CC-BY-4.0.


Li, L., X. Zhang, C. B. Skovsted, H. Yun, B. Pan, and G. Li. 2019. Homologous shell microstructures in Cambrian hyoliths and molluscs. Palaeontology 62(4):515–532. doi:10.1111/pala.12406.

Li, L., C. B. Skovsted, H. Yun, M. J. Betts, and X. Zhang. 2020. New insight into the soft anatomy and shell microstructures of early Cambrian orthothecids (Hyolitha). Proceedings of the Royal Society B 287(1933):20201467. doi:10.1098/rspb.2020.1467.

Liu, F., C. B. Skovsted, T. P. Topper, Z. Zhang, and D. Shu. 2020a. Are hyoliths Palaeozoic lophophorates? National Science Review 7(2):453–469. doi:10.1093/nsr/nwz161.

Liu, F., C. B. Skovsted, T. P. Topper, and Z. Zhang. 2020b. Revision of Triplicatella (Orthothecida, Hyolitha) with preserved digestive tracts from the early Cambrian Chengjiang Lagerstätte, South China. Historical Biology. doi:10.1080/08912963.2020.1747059.

Liu, F., C. B. Skovsted, T. P. Topper, and Z. Zhang. 2020c. Soft part preservation in hyolithids from the lower Cambrian (Stage 4) Guanshan Biota of south China and its implications. Palaeogeography, Palaeoclimatology, Palaeoecology. doi:10.1016/j.palaeo.2020.110079.

Moysiuk, J., M. R. Smith, and J.-B. Caron. 2017. Hyoliths are Palaeozoic lophophorates. Nature 541:394–397. doi:10.1038/nature20804.

Skovsted, C. B., M. Martí Mus, Z. Zhang, B. Pan. L. Li, F. Liu, G. Li, and Z. Zhang. 2020. On the origin of hyolith helens. Palaeogeography, Palaeoclimatology, Palaeoecology 555:109848. doi:10.1016/j.palaeo.2020.109848.

Smith, M. R. 2020. Finding a home for hyoliths. National Science Review 7(2):470–471. doi:10.1093/nsr/nwz194.

Sun, H., M. R. Smith, H. Zeng, F. Zhao, G. Li, and M. Zhu. 2018. Hyoliths with pedicles illuminate the origin of the brachiopod body plan. Proceedings of the Royal Society B: Biological Sciences 285(1887). doi:10.1098/rspb.2018.1780.

Sunday, October 18, 2020

Your Friends The Titanosaurs, part 29: Tapuiasaurus, Tengrisaurus, and Traukutitan

We start off with two of the geologically oldest described titanosaurs before jumping back to the Late Cretaceous of Patagonia. If you're keeping track, you may notice that Titanosaurus seemingly should have been included. It's being held over for next month, which will be all Titanosaurus, including various species originally named to Titanosaurus that don't belong there and haven't otherwise been covered.

Sunday, October 11, 2020

Compact Thescelosaurus Year Five

It's time for the annual review of The Compact Thescelosaurus. As is traditional, there's a new sheet, which this year covers mosasaurs, as you might have guessed from the image. The classification diagram page has been updated as well. (Don't forget, Wednesday the 14th is also National Fossil Day!)

Mosasaurus conodon, hanging out at the Science Museum.