Your Friends The Titanosaurs, part 36.1: Coming Attractions (outside of Europe and South America)

Here we have a few of the many, many unnamed and/or undescribed titanosaurs. Some of their common habitats are abstracts, dissertations, press releases, and offhand comments in publications. Sometimes they're the subject of a paper describing some particular anatomical topic, or a geographic range extension. In other cases they're among five different morphotypes in a bonebed, and you can only hope that this pans out, because otherwise there's no negotiating with a von Huene puzzle. All the while you grit your teeth because you despair of ever seeing the really notable specimens published, especially so long as there's a theropod knuckle left to be described...

Anyway, this is not quite like "Coming Attractions in Dinosauria", which was based primarily on unnamed species with nicknames in long use. This is more like the original Addendum II in "The New Dinosaur Dictionary", except with specimen numbers and more references. It is certainly not exhaustive, and would take a lot more than a few weeks to make so. I am omitting embryonic forms (note that titanosaur nests have been found in several areas, including Argentina, India, and Spain), tracks, and named forms that need further description (Ampelosaurus, Futalognkosaurus, etc.). I've organized this by continent, and then by country; it turns out that if a country has terrestrial Cretaceous rocks, there is an excellent chance it has titanosaurs. Because of the sheer number of entries, I opted to split the post, with Europe and South America to come later.

I don't know if this will impress anyone, but I had a lot of fun writing this post.

Africa/Arabia

Egypt: The Quseir Formation has already produced Mansourasaurus shahinae, but there are other titanosaur specimens from the Dakhla and Kharga Oases, including a saltasaur-like caudal (Salem et al. 2021). One of the better specimens is MB.R.Vb-621–640, which has appeared mostly in abstracts (e.g., Díez Díaz et al. 2017; Lamanna et al. 2017; Gorscak et al. 2020). It was discovered in 1977 but, obviously, has not yet been formally described. The specimen includes five dorsal vertebrae and twelve bones of the girdles and limbs, most notably at least four of the five metatarsals of one of the feet (Díez Díaz et al. 2017), a portion of the skeleton poorly represented in titanosaurs. The abstracts agree in describing it as a notably gracile titanosaur. [Update, 2023/7/23: This is now Igai semkhu.] A second Quseir Formation titanosaur skeleton from Kharga Oasis is briefly described in Salem et al. (2020). This one includes a cervical, five dorsals, a caudal, and a tibia and astragalus, and has a more robust tibia than the first skeleton.

Jordan: Jordan is not exactly a hotbed of nonavian dinosaurs, but it has produced a titanosaur specimen. First mentioned in Wilson et al. (2006) as a strongly pneumatic form with similarities to Malawisaurus dixeyi, further details emerged in a pair of abstracts for the 2012 Society of Vertebrate Paleontology convention, D'Emic et al. (2012) and O'Connell et al. (2012). None of the abstracts have gone right out and given an inventory, but the material includes at least a posterior dorsal, sacrum, posterior caudals, and a femur belonging to a dwarf adult found in Maastrichtian-age rocks.

Kenya: Several titanosaurs have been described from Kenya, but none yet from the ?Maastrichtian Lapurr Sandstone (also spelled "Lapur", or reported as the Turkana Grits). They are there, though. The most readily accessible reference is probably the abstract Gorscak et al. (2019), which provides about a paragraph on KNM-WT 65086 and other titanosaurian specimens from the formation, indicating the presence of multiple species including giants. However, Gorscak's 2016 dissertation is available if you don't mind a 452 mb download, and includes a chapter on the specimen (as well as chapters on several other specimens that have since been official described, plus a re-analysis of the titanosaurs from Malawi. Hope you weren't too attached to Malawisaurus!). KNM-WT 65086 is an associated skeleton including fragments of presacrals, parts of nine caudals, partial right scapula and humerus, left ulna and distal radius, a partial femur, the left and part of the right fibula, an osteoderm, and other bits of limb bones. The caudals seem to lack pneumatic features. Several other kinds of sauropod caudals have been found in the formation, including amphiplatyan caudals suggesting but not guaranteeing non-titanosaurian titanosauriforms. The formation is also good for osteoderms, with large examples of "keeled", "ellipsoidal" and "bulb and root" forms. Gorscak (2016) postulated a minimum of four sauropod taxa.

Mali: O'Leary et al. (2004) described a group of specimens including three caudals, an osteoderm, and a possible fibula from a bone bed in the "Continental Intercalaire" of northern Mali. It shows up from time to time in the titanosaur osteoderm literature and in discussions of large vertebrate biogeography.

Morocco: At least one titanosaur species is represented in the Cenomanian Kem Kem Group of Morocco, where titanosaur fossils including narrow-crowned teeth, caudal vertebrae, a humerus, and a partial ulna have been found (Ibrahim et al. 2020). Of note, the partial ulna is quite large; the report is confusing at this point, because the text discusses UCRC PV9 from the Douira Formation as 51 cm wide (20 in), while Table 10 discussed UCRC PV362 from the Gara Sbaa Formation, "only" 44.0 cm (17.3 in) in maximum width. Either way, there was a large-bodied titanosaur in the Kem Kem, potentially in the vicinity of Paralititan stromeri (of roughly the same age, from the other end of north Africa). The much younger Maastrichtian-age Couche III phosphates of Sidi Dauoi have produced sauropod remains including a partial hindlimb (OCP DEK/GE 31) originally described as titanosauriform (Suberbiola et al. 2004), more recently interpreted as titanosaurian based on additional fossils (Longrich et al. 2017).

Who'd like a giant titanosaur? This is Figure 109 in Ibrahim et al. (2020). The caption reads "Proximal section of a right ulna of a giant sauropod. UCRC PV9 in (A) anterior, (B) posterior
and (C) distal view. Scale bar equals 20 cm. Abbreviations: alp anterolateral process amp anteromedial
process opr olecranon process."  CC BY 4.0. (Incidentally, there aren't a lot of photos available for this post; for one thing, abstracts are rarely illustrated.)

Niger: Titanosaurs are reportedly present in both the Elrhaz Formation (late Aptian–early Albian) and Echkar Formation (late Albian–Cenomanian) of Niger (Ibrahim et al. 2020), although I have a slight suspicion that the Echkar report is the same as the Elrhaz report. Very little detail has escaped on the Nigerien titanosaur record; about the best I've found is in a footnote to Sereno et al. (1999) (#13), which cites a tall, narrow astragalus and procoelous caudals with spongy bone and anteriorly placed neural arches.

Saudi Arabia: A group of seven titanosaur caudals has been found in the Adaffa Formation (probably Campanian–Maastrichtian) of extreme northwestern Saudi Arabia. They are poorly preserved; only one has a good neural arch and neural spine, which is relatively long and strongly backswept (Kear et al. 2013).

The best caudal from the Adaffa titanosaur, cropped from Figure 2 of Kear et al. (2013) to remove the theropod tooth. CC BY 4.0.

South Africa: I don't have much on this one, but a couple of vertebrae considered possibly titanosaurian have been found in an unnamed unit of Santonian age near the coast in eastern South Africa (Kennedy et al. 1987).

Sudan: Although the Wadi Milk dromaeosaur is by far the most famous nonavian dinosaur of Sudan, at least one titanosaur taxon is also present in the Wadi Milk Formation of northern Sudan. In fact, sauropod remains are regarded as the most common dinosaur fossils of this unit (Rauhut 1999). Titanosaurs are represented by teeth, sixteen vertebrae, and a partial fibula. Differences among the vertebrae suggest more than one species was present (Rauhut 1999).

Zimbabwe: Isolated and fragmentary dinosaurian fossils are known from the Gokwe Formation, central Zimbabwe, which has a poorly constrained age but is thought to be Cretaceous. Titanosaur caudals from the Gokwe Formation include two morphologies. There are also titanosaurian-type teeth that are rod-like and have high-angle wear facets, and a titanosaurian osteoderm (Woolley et al. 2015).

Antarctica

A fragmentary procoelous middle caudal from the Campanian Santa Maria Formation of James Ross Island provides evidence for the presence of titanosaurs in Antarctica. This is also the first evidence of sauropods in general from the continent (Cerda et al. 2011, 2012).

Asia

China: We have a few actual informal names for China, but as we saw in Part 35, the situation with titanosaur-like sauropods from China is pretty hazy; committing to an identification for an undescribed specimen is brave or foolish. For example, there is "Yunxianosaurus hubeinensis", briefly reported in Li (2001). It is from the Majiacun Group of Hubei and is represented by partial remains including cervicals, dorsals, sacrals, caudals, ribs, scapula, ilium, pubis, humerus, radius, ulna, femur, and tibia (tip of the hat to The Theropod Database!). The informally named "Megacervixosaurus tibetensis" (one of several longstanding informal names introduced by Zhao in the mid-1980s) from the Zonggo Formation of Tibet is sometimes regarded as a titanosaur on biostratigraphic grounds. Presumably some large cervical vertebrae are involved, given the name. It would be interesting to see how it compares to various Cretaceous Chinese sauropods with long necks and uncertain positions. (See also: "Xinghesaurus")

Mongolia: The most notorious unnamed titanosaur from Mongolia is the Bor Guvé form, a.k.a. IGM 100/3005. It is represented by fifteen caudals and one posterior dorsal that probably belong to the same individual, found in what is thought to be the lower Upper Cretaceous Bayan Shireh Formation (best known for its therizinosaur infestation). The Bor Guvé form gets several pages in Kspeka and Norell (2010), in part to show that it isn't the same as Erketu ellisoni, also known from the Bayan Shireh Formation of Bor Guvé (about 2 km/1 mi from IGM 100/3005).

In addition, there is plenty of undescribed titanosaur material in the younger Barun Goyot and Nemegt Formations, but most of it is pieces-parts or footprints (Currie et al. 2018). The latter include some impressively large specimens from the Nemegt, representing individuals "64%–120% larger" than the largest skeletal remains (Stettner et al. 2018).

Siberian Russia: Averianov et al. (2020) distinguished three indeterminate lithostrotian taxa from three middle caudals found in the Barremian–Aptian Ilek Formation of western Siberia, which has also produced the aptly named titanosauriform Sibirotitan.

Uzbekistan: I could have split this between Kazakhstan, Kyrgyzstan, Tajikistan, and Uzbekistan, which have all produced fragmentary sauropod remains, but most of the specimens are skinny titanosaur-type teeth (Averianov and Sues 2017). At the moment the most significant specimen is a titanosaurian braincase (CCMGE 628/12457) from the Turonian Bissekty Formation of Dzharakuduk, Uzbekistan, which was the major subject of Sues et al. (2015) after an inauspicious start as a braincase of Turanoceratops (Nesov 1995). We just covered another major subject of Sues et al. (2015), caudal vertebra USNM 538127, now the holotype of Dzharatitanis kingi. For my money the fairly trad titanosaurian braincase and the distinctive caudal indicate two different taxa.

Australia/New Zealand

Australia: Poropat et al. (2019) briefly reported the 2018 discovery of a partial skull and hindlimb of a titanosauriform from the Cenomanian–early Turonian Winton Formation of Queensland, Australia. Although this is not necessarily a titanosaurian, the longstanding tendency of Wintonotitan wattsi to slip into Titanosauria makes a mention here seem appropriate.

There are a number of Winton Formation titanosaur/iform specimens that have not been described or allocated to an existing species. They go under nicknames such as "Cooper", "George", "Sid", "Tom", and "Zac". "Cooper" and "George" seem to be the best-publicized specimens, and are currently listed on a Wikipedia article as pertaining to the informal "Bananabendersaurus", which does not seem like the kind of name that is going to survive to publication. Anyway, this is reportedly the largest known Australian dinosaur. [Update, 2021/06/07: And as mentioned in the comments, we have our first contestant to come off the board, with "Cooper" becoming the holotype of Australotitan cooperensis.]

New Zealand: Molnar and Wiffen (2007) reported a partial centrum (CD.586) interpreted as a titanosaurian's procoelous middle caudal (it's not always obvious which end is the anterior and which is the posterior when you don't have the neural arch). It was found in the Maungataniwha Sandstone (probably about the Campanian–Maastrichtian boundary) near Hawke's Bay on the North Island of New Zealand.

India

Among the various challenges facing study of titanosaurs from India and Pakistan are the inevitable specimens that either do not fit or cannot be compared to the named taxa. For example, there is the "Nand axis" (196/CRP/GSI/05) of the Lameta Formation of Nand, India, which is much more pneumatic than the known cervicals of Isisaurus colberti and can't be compared to Jainosaurus septentrionalis (Wilson and Mohabey 2006). Wilson et al. (2019) described three titanosaur vertebrae. One is an anterior caudal neural arch (338/GSI/PAL/CR/2017) from Bara Simla Hill and the other two are dorsals from Rahioli (GSI/GC/OGF107 and GSI/GC/2905), likely from one individual. The dorsals may pertain to Isisaurus but this is unclear in the absence of comparative material for Jainosaurus. If they *are* from Isisaurus, they may indicate a relationship to lognkosaurians because they also resemble dorsals of Mendozasaurus. Coincidentally enough, the caudal neural arch also has similarities to anterior caudal neural arches of Mendozasaurus (and Futalognkosaurus), but it is even more similar to that of Tengrisaurus starkovi.

North America

Canada: Now here's a headscratcher: Ryan and Evans (2018) reported the discovery of a titanosaur caudal centrum from the late Santonian Deadhorse Coulee Member of the Milk River Formation, southern Alberta. This definitely qualifies for "where did that come from?" Were titanosaurs all over North America in the Late Cretaceous, but they just avoided depositional basins until Alamosaurus? Was this an early failed colonization, perhaps from a northern route? Did some bright joker plant a titanosaur caudal?

References

Averianov, A., and H.-D. Sues. 2017. Review of Cretaceous sauropod dinosaurs from central Asia. Cretaceous Research 69:184–197. doi:10.1016/j.cretres.2016.09.006.

Averianov, A. O., S. V. Ivantsov, and P. P. Skutschas. 2020. Caudal vertebrae of titanosaurian sauropod dinosaurs from the Lower Cretaceous Ilek Formation in western Siberia, Russia. Cretaceous Research 107:104309. doi:10.1016/j.cretres.2019.104309.

Cerda, I., A. Paulina Carabajal, L. Salgado, R. Coria, and J. J. Moly. 2011. The first record of sauropod dinosaurs from Antarctica. Journal of Vertebrate Paleontology, Program and Abstracts, 2011:86.

Cerda, I. A., A. Paulina Carabajal, L. Salgado, R. A. Coria, M. A. Reguero, C. P. Tambussi, and J. J. Moly. 2012. The first record of a sauropod dinosaur from Antarctica. Naturwissenschaften 99:83–87.

Currie, P. J., J. A. Wilson, F. Fanti, B. Mainbayar, and K. Tsogtbaatar. 2018. Rediscovery of the type localities of the Late Cretaceous Mongolian sauropods Nemegtosaurus mongoliensis and Opisthocoelicaudia skarzynskii: stratigraphic and taxonomic implications. Palaeogeography, Palaeoclimatology, Palaeoecology 494:5–13. doi:10.1016/j.palaeo.2017.10.035.

D'Emic, M. D., and J. A. Wilson. 2012. Bone histology of a dwarf sauropod dinosaur from the latest Cretaceous of Jordan and a possible biomechanical explanation for "titanosaur-type" bone histology. Journal of Vertebrate Paleontology, Program and Abstracts, 2012:83.

Díez Díaz, V., E. Gorscak, M. C. Lamanna, D. Schwarz, and I. El-Dawoudi. 2017. The metatarsus of a Late Cretaceous titanosaur (Dinosauria: Sauropoda) from the Kharga Oasis of Egypt. Zitteliana 91:31–32.

Gorscak, E. 2016. Descriptive and comparative morphology of African titanosaurian sauropods: new information on the evolution of Cretaceous African continental faunas [file is 452 mb]. Dissertation. Ohio University, Athens, Ohio.

Gorscak, E., J. Sertich, and F. K. Manthi. 2019. Titanosaurian sauropod dinosaur fossils from the Upper Cretaceous Lapur Sandstone (Turkana Grits), Turkana Basin, northwestern Kenya. Journal of Vertebrate Paleontology, Program and Abstracts, 2019:108.

Gorscak, E., M. C. Lamanna, V. Díez Díaz, D. Schwarz, B. S. Salem, G. Abu El-Kheir, and H. Sallam. 2020. A titanosaurian sauropod from the Campanian Quseir Formation of the Kharga Oasis, Egypt, supports Afro-Eurasian dinosaur faunal connectivity during the Late Cretaceous. Journal of Vertebrate Paleontology, Program and Abstracts, 2020:no page number because in the future we just know where things are.

Ibrahim, N., P. C. Sereno, D. J. Varricchio, D. M. Martill, D. B. Dutheil, D. M. Unwin, L. Baidder, H. C. E. Larsson, S. Zouhri, and A. Kaoukaya. 2020. Geology and paleontology of the Upper Cretaceous Kem Kem Group of eastern Morocco. ZooKeys 928:1–216. doi:10.3897/zookeys.928.47517.

Kear, B. P., T. H. Rich, P. Vickers-Rich, M. A. Ali, Y. A. Al-Mufarreh, A. H. Matari, A. M. Al-Massari, A. H. Nasser, Y. Attia, and M. A. Halawani. 2013. First dinosaurs from Saudi Arabia. PLoS ONE 8(12):e84041. doi:10.1371/journal.pone.0084041.

Kennedy, W. J., H. C. Klinger, and N. J. Mateer. 1987. First record of an Upper Cretaceous sauropod dinosaur from Zululand, South Africa. South African Journal of Science 83:173–174.

Ksepka, D. T., and M. A. Norell. 2010. The illusory evidence for Asian Brachiosauridae: new material of Erketu ellisoni and a phylogenetic reappraisal of basal Titanosauriformes. American Museum Novitates 3700.

Lamanna, M. C., E. Gorscak, V. Díez Díaz, D. Schwarz, and I. El-Dawoudi. 2017. Reassessment of a partial titanosaurian sauropod dinosaur skeleton from the Upper Cretaceous (Campanian) Quseir Formation of the Kharga Oasis, Egypt. Zitteliana 91:50–51.

Li, Z. 2001. Distribution, burying and classification of dinosaur fossils in Upper Cretaceous strata at Meipu Town, Yunxian County of Hubei Province. Hubei Geology & Mineral Resources 15(4):25–31.

Longrich, N. R., X. Pereda-Suberbiola, N.-E. Jalil, F. Khaldoune, and E. Jourani. 2017. An abelisaurid from the latest Cretaceous (late Maastrichtian) of Morocco, North Africa. Cretaceous Research 76:40–52. doi:10.1016/j.cretres.2017.03.021.

Molnar, R. E., and J. Wiffen. 2007. A presumed titanosaurian vertebra from the Late Cretaceous of North Island, New Zealand. Arquivos do Museu Nacional, Rio de Janeiro 65(4):505–510.

Nesov, L. A. 1995. [Dinosaurs of northern Eurasia: new data about assemblages, ecology and paleobiogeography]. Izdatelstvo Sankt-Peterburgskogo Universiteta, St. Petersburg, Russia. [in Russian]

O'Connell, T. L., J. A. Wilson, and I. S. Zalmout. 2012. Air space proportion in a dorsal vertebra of a new titanosaur (Dinosauria: Sauropoda) from Jordan. Journal of Vertebrate Paleontology, Program and Abstracts, 2012:151.

O'Leary, M. A., E. M. Roberts, J. A. Head, F. Sissoko, and M. L. Bouaré. 2004. Titanosaurian (Dinosauria: Sauropoda) remains from the "Continental Intercalaire" of Mali. Journal of Vertebrate Paleontology 24(4):923–930.

Poropat, S. F., P. D. Mannion, P. Upchurch, and D. A. Elliott. 2019. New sauropod dinosaur discoveries in the lower Upper Cretaceous Winton Formation (Cenomanian–lower Turonian) of Queensland, Australia: implications for titanosaurian evolution. Journal of Vertebrate Paleontology, Program and Abstracts, 2019:171.

Rauhut, O. W. M. 1999. A dinosaur fauna from the Late Cretaceous (Cenomanian) of northern Sudan. Palaeontologia Africana 35:61–84.

Ryan, M. J., and D. C. Evans. 2018. The first occurrence of a sauropod body fossil in Canada, with implications for the "sauropod hiatus" in North America. Journal of Vertebrate Paleontology, Program and Abstracts, 2018:207.

Salem, B. S., G. Abu El-Kheir, M. C. Lamanna, E. Gorscak, S. El-Sayed, and H. Sallam. 2020. A new titanosaurian sauropod dinosaur partial skeleton from the Late Cretaceous (Campanian) of the Kharga Oasis, Western Desert of Egypt. Journal of Vertebrate Paleontology, Program and Abstracts, 2020:no page number because in the future we just know where things are.

Salem, B. S., P. M. O'Connor, E. Gorscak, S. El-Sayed, J. J. Sertich, E. Seiffert, and H. M. Sallam. 2021. Dinosaur remains from the Upper Cretaceous (Campanian) of the Western Desert, Egypt. Cretaceous Research 123:article 104783. doi:10.1016/j.cretres.2021.104783.

Sereno, P. C., A. L. Beck, D. B. Dutheil, H. C. Larsson, G. H. Lyon, B. Moussa, R. W. Sadleir, C. A. Sidor, D. J. Varricchio, G. P. Wilson, and J. A. Wilson. 1999. Cretaceous sauropods from the Sahara and the uneven rate of skeletal evolution among dinosaurs. Science 286(5443):1342–1347.

Stettner, B., W. S. Persons IV, and P. J. Currie. 2018. A giant sauropod footprint from the Nemegt Formation (Upper Cretaceous) of Mongolia. Palaeogeography, Palaeoclimatology, Palaeoecology 494:168−172. doi:10.1016/j.palaeo.2017.10.027.

Suberbiola, X. P., N. Bardet, M. Iarochène, B. Bouya, and M. Amaghzaz. 2004. The first record of a sauropod dinosaur from the Late Cretaceous phosphates of Morocco. Journal of African Earth Sciences 40:81–88.

Sues, H.-D., A. Averianov, R. C. Ridgely, and L. M. Witmer. 2015. Titanosauria (Dinosauria, Sauropoda) from the Upper Cretaceous (Turonian) Bissekty Formation of Uzbekistan. Journal of Vertebrate Paleontology 35(1):e889145. doi:10.1080/02724634.2014.889145.

Wilson, J. A., and D. M. Mohabey. 2006. A titanosauriform (Dinosauria: Sauropoda) axis from the Lameta Formation (Upper Cretaceous: Maastrichtian) of Nand, central India, Journal of Vertebrate Paleontology 26(2):471–479.

Wilson, J. A., H. Mustafa, and I. Zalmout. 2006. Latest Cretaceous reptiles from the Hashemite Kingdom of Jordan. Journal of Vertebrate Paleontology 26(supplement to 3):140A.

Wilson, J. A., D. M. Mohabey, P. Lakra, and A. Bhadran. 2019. Titanosaur (Dinosauria: Sauropoda) vertebrae from the Upper Cretaceous Lameta Formation of western and central India. Contributions of the Museum of Paleontology, University of Michigan 33(1):1–27.

Woolley, H., J. Sertich, C. A. Forster, D. Munyikwa, S. D. Sampson, K. Curry Rogers, and R. R. Rogers. 2015. Titanosaurian and other vertebrate remains from the Cretaceous Gokwe Formation, central Zimbabwe. Journal of Vertebrate Paleontology, Program and Abstracts, 2015:241.