Kholumolumo ellenbergerorum and "Thotobolosaurus"

With the twin unofficial missions of Equatorial Minnesota to cover Elliot Formation "prosauropods" and find out what happened to unpublished dinosaurs of the past, it was imperative that Kholumolumo ellenbergerorum receive a post. Before we get into the actual science of K. ellenbergerorum, though, let's step back through the mists of time. First we'll stop off in the far-off world of 1985.

You've just gotten the Normanpedia (Norman 1985) and are busy committing it to memory. On page 97, in the "Dubious" section of the list of prosauropod genera, is Thotobolosaurus. It doesn't ring a bell, so you pull out "A Field Guide to Dinosaurs" (Lambert 1983). It shows up on page 104, as a "roccosaurid" with Riojasaurus and something called Roccosaurus, which wasn't in the Normanpedia. "Roccosaurids" apparently have "unusually sharp fangs and an especially strong joint linking hips with the backbone", but might actually be melanorosaurids. This last bit just ends up being confusing because (stepping outside of 1985 for a moment) "A Field Guide to Dinosaurs" was caught up in the contemporary fad for lumping all kinds of prosauropods, so Melanorosaurus is sunk into Euskelosaurus. Anyway, Thotobolosaurus is "a big prosauropod that lived in early Late Triassic Lesotho, southern Africa." Okay, sure. Does "The New Dinosaur Dictionary" (Glut 1982) have anything more, and can it explain what exactly is a Roccosaurus?

Yes, it can! Thotobolosaurus is described on page 243, in the main section of the book. It is listed as named by Ellenberger in 1970 and is a potential melanorosaurid. "From the Lower Stromberg Series of Lesotho and South Africa, this dinosaur is reportedly a large, possibly quadrupedal melanorosaurid very much like a true sauropod. The form corresponds to the ichnite Pseudotetrasauropus. A description of the genus has not yet been published." Wait, it's not actually described? How did it get into all of these books? (Same question, 1990s, just substitute Nurosaurus for Thotobolosaurus.) And what the heck is a Roccosaurus? It's not in here! (Actually, it is, on page 275, as "a new dinosaur genus based on good postcranial material... has short, high cervical vertebrae, four sacrals, and shortened forelimbs". Want to know where it is today? Just look for NM QR1551, one of the major catalog numbers assigned to Melanorosaurus readi.)

Flash ahead to 1990 and "The Dinosaur Data Book" (Lambert 1990). There's "Thotobolosaurus" again, as if it had been formally named and described. "Roccosaurus", meanwhile, is listed as Melanorosaurus. Neither are in "The Dinosauria", but both will get short entries in the "Nomina Nuda" section of Glut (1997). Following this, "Thotobolosaurus" disappeared from books, to linger as a taxonomic ghost on the Internet until April 2020, when it was resurrected as Kholumolumo (Peyre de Fabrègues and Allain 2020).

The story goes back much farther than the 1980s, or even Ellenberger (1970) for that matter. Peyre de Fabrègues and Allain (2020) include a history of this "prosauropod" and its many changes of alias, encompassing much more than just "Thotobolosaurus". We're going to go back to 1930, when Samuel Motsoane, principal of the Paris Evangelical Mission School at Bethesda, Lebanon, found some dinosaur fossils. He told Protestant missionary Paul Ellenberger about this in 1955. Ellenberger in turn began prospecting near the mission in Maphutseng in August 1955, bringing in his brother, geologist François Ellenberger. They found a rich bonebed at Thotobolo ea 'Ma-Beata' in September 1955. Later that year and in 1956, they excavated there with South African geologists A. W. Crompton of the National Museum of Bloemfontein and R. F. Ewer of Rhodes University in Grahamstown. These excavations produced 683 pieces (~450 complete bones), which were put in storage at the South African Museum in Cape Town (now the Iziko Museum). The Ellenberger brothers and the South African contingent fell out after this, and the material sent to Cape Town is currently at the University of Cape Town. The brothers instead began working with the Muséum national d'histoire naturelle (MNHN) in Paris, which received further collections of bones from 1959, 1963, and 1970. These included something like 400 pieces constituting 210 bones. Tracks collected at this time (remember the ichnites in Glut 1982) have gone missing.

All this time, Paul Ellenberger was publishing on the Maphutseng dinosaur, beginning with Ellenberger (1955) and going up to 1970. As included in Glut (1982), it is indeed in Ellenberger (1970) that we get "Thotobolosaurus mabeatae". Before this time, it had been an unnamed form, an indeterminate melanorosaurid, or Euskelosaurus, because at one time or another every "prosauropod" from southern Africa has been either Euskelosaurus or Massospondylus. Sometimes a specimen actually is, although these days that's only true of Massospondylus. "Thotobolosaurus", as we saw above, became an unofficially official name for quite a while. Gauffre (1996) took another stab at unofficially naming it, this time as "Kholumolumosaurus ellenbergerorum", while other authors were content to use "Maphutseng dinosaur" or some comparable geographic appellation. After all of that, it was a bit of surprise to actually get this dinosaur described officially in print, now as Kholumolumo ellenbergerorum.

Genus and species: Kholumolumo ellenbergerorum. The genus name is from Sotho folklore, and is the name of a type of reptilian monster often used by Basotho people when describing dinosaurs. "Ellenbergerorum" honors the Ellenberger brothers (Peyre de Fabrègues and Allain 2020). Together, we get something like "the Ellenberger brothers' kholumolumo" or, less circularly, "the Ellenberger brothers' folkloric reptilian monster". This beats "Kholumolumosaurus", which is redundant ("folkloric reptilian monster lizard"), and "Thotobolosaurus mabeatae", meaning "Beata's mother's trash heap lizard", which seems kind of flippant about the whole thing.

In unrelated news, a specimen previously assigned to Chuanjiesaurus anaensis has been given its own genus and species. The name? Analong chuanjieensis, which basically flips the original name. I'm not sure whether that's brilliant, awful, or both.

Citation: Peyre de Fabrègues, C., and R. Allain. 2020. Kholumolumo ellenbergerorum, gen. et sp. nov., a new early sauropodomorph from the lower Elliot Formation (Upper Triassic) of Maphutseng, Lesotho. Journal of Vertebrate Paleontology in press: e1732996. doi:10.1080/02724634.2019.1732996.

Stratigraphy and geography: We've gone over some of this already. To recapitulate: K. ellenbergerorum comes from a bonebed in the Lower Elliot Formation, near a Protestant mission at Maphutseng, Mohale's Hoek District, southwestern Lesotho. Although the upper part of the Lower Elliot Formation has been suggested by some authors, Peyre de Fabrègues and Allain (2020) favor a provenance in the lower Lower Elliot.

Holotype: Out of the hundreds of bones known, a tibia catalogued as MNHN.F.LES381m was selected as the holotype, due to its diagnostic features. (When you think about it, it's got to be kind of difficult to select a holotype from a bonebed of disassociated bones, even if it's basically monospecific like this one.) The other bones were regarded as paratypes, except for two rauisuchid teeth (Peyre de Fabrègues and Allain 2020). If you recall the description of "roccosaurids" above, they were supposed to have "unusually sharp fangs". This gets into a long story about how it took a surprisingly long time for it to really sink in that carnivorous archosaur teeth can be found disassociated with non-carnivorous archosaurs.

Peyre de Fabrègues and Allain (2020) focused on the MNHN material. Of this, the presence of five left radii (as stated in the supplementary material; the text said "right", but that's a simple enough typo) indicates at least five individuals, and since the MNHN material is less than half of the total, the actual number of individuals could easily be ten, if not more. The bones are disarticulated, but the presence of small elements like phalanges and distal caudals indicates that although the carcasses clearly came undone, there was not enough dispersal going on to strip out the small bits. The two rauisuchian teeth show that there was some scavenging. The list of MNHN material includes a bit of everything, although skulls are only represented by two postorbitals, and overall the list seems relatively light on vertebrae and relatively heavy on limb bones and girdle elements. (The University of Cape Town material is not broken down like the MNHN material, so my impressions could be skewed.) In short, material-wise there is little lacking to make K. ellenbergerorum one of the best-known "prosauropods", even if actual proportions can't be determined due to the disassociation of the material.

One of the most outstanding features of K. ellenbergerorum is its size. Granted, it's not a patch on our titanosaurs, but a "prosauropod" estimated to be on the order of 9 m long (30 ft) and 3 metric tons (3.3 short tons) (Peyre de Fabrègues and Allain 2020) is perfectly respectable for the Late Triassic. The limb and girdle bones tend to the stocky, robust side of the spectrum. The known sacral vertebra is also notably robust. (One gets the impression, between this and some of the other large "prosauropods", that sauropodomorphs were still figuring out how to be big at this point, and so weren't yet doing it "gracefully".) Phylogenetically, K. ellenbergerorum plops down at the base of a clade called Massopoda, between the earlier-diverging plateosaurs and the later-diverging massospondyls, making its own little sub-clade with North American Sarahsaurus and East Asian Xingxiulong (Peyre de Fabrègues and Allain 2020). Geographically that's kind of eclectic, but you could get away with that kind of thing back then thanks to Pangaea.

Peyre de Fabrègues and Allain (2020) also compared K. ellenbergerorum to the other "prosauropods" of the Lower Elliot. The most similar seemed to be Plateosauravus, although as noted K. ellenbergerorum did not closely clade with any Lower Elliot taxa. McPhee et al (2017), referring to it by the "Maphutseng dinosaur" alias, considered it to potentially fill an ecological role between the robust oddball Blikanasaurus and the more generically "prosauropodan" Eucnemesaurus and Plateosauravus, and suggested that yet another ex-Euskelosaurus from the Lower Elliot, the hindlimb and tail designated SAM-PK 382, may be something similar.

Figure 9 in McPhee et al. (2017), depicting different morphotypes. You may now substitute Kholumolumo for the "Maphutseng dinosaur". CC-BY-4.0.

References

Ellenberger, P. 1955. Note préliminaire sur les pistes et restes osseux de vertébrés du Basutoland (Afrique du Sud). Comptes Rendus de l’Académie des Sciences 240:889–891.

Ellenberger, P. 1970. Les niveaux paléontologiques de première apparition des mammifères primordiaux en Afrique du Sud et leur ichnologie: établissement de zones stratigraphiques détaillées dans le Stormberg du Lesotho (Afrique du Sud) (Trias supérieur à Jurassique). Pages 343–370 in S. H. Haughton, editor. IUGS Commission on Stratigraphy: Proceedings and Papers of the 2nd Gondwana Symposium, Pretoria, July to August 1970. Council for Scientific and Industrial Research, Pretoria, South Africa.

Gauffre, F.-X. 1996. Phylogénie des dinosaures prosauropodes et étude d’un nouveau prosauropode du Trias supérieur d’Afrique australe. Dissertation. Muséum National d’Histoire Naturelle, Paris, France.

Glut, D. F. 1982. The new dinosaur dictionary. Citadel Press, Secaucus, New Jersey.

Glut, D. F. 1997. Dinosaurs: the encyclopedia. McFarland & Co., Jefferson, North Carolina.

Lambert, D. 1983. A field guide to dinosaurs. Avon Books, New York, New York.

Lambert, D. 1990. The dinosaur data book. Avon Books, New York, New York.

McPhee, B. W., E. M. Bordy, L. Sciscio, and J. N. Choiniere. 2017. The sauropodomorph biostratigraphy of the Elliot Formation of southern Africa: tracking the evolution of Sauropodomorpha across the Triassic–Jurassic boundary. Acta Palaeontologica Polonica 62:441–465. doi:10.4202/app.00377.2017.

Norman, D. 1985. The illustrated encyclopedia of dinosaurs. Crescent Books, New York, New York.

Peyre de Fabrègues, C., and R. Allain. 2020. Kholumolumo ellenbergerorum, gen. et sp. nov., a new early sauropodomorph from the lower Elliot Formation (Upper Triassic) of Maphutseng, Lesotho. Journal of Vertebrate Paleontology in press: e1732996. doi:10.1080/02724634.2019.1732996.