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Lapparentosaurus madagascariensis skeletal

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Lapparentosaurus madagascariensis

Etymology: "Lapparent's lizard from Madagascar"

Time horizon: Middle Jurassic, Bathonian epoch (~167 mya)

Length: ~24m (~79 ft.), based on adult caudal remains and scaling up proportions from immature specimens. *Updated with correct scaling for Lydekker's giant vertebrae*

Probable mass: 30+ tons

The earliest "normal" brachiosaur that we have at least semi-decent material for, Lapparentosaurus madagascariensis. Of course, exactly what is "normal" for a brachiosaur is still very much open to debate, but at least it's the most basal one that has the long thumb metacarpals typical of later species in the family. This animal is surprisingly well-represented in the fossil record, yes considering that, it's one of the most poorly researched brachiosaurs in existence. The first remains were found in the late 1800s, as British and French paleontologists rushed worldwide to discover new dinosaurs, often fragments of sauropods that were inevitably thrown without much thought into Titanosaurus, Cetiosaurus, or Bothriospondylus.

Such was the case with the material which would later become known as Lapparentosaurus. The history of "Bothriospondylus" is a long and insanely confusing one. But in a nutshell it was one of the original Dinosauria described by Owen in the heady times of the Victorian age, and the type species appears to have been a brachiosaur, or at least a basal macronarian of some sort. Other remains from England and Portugal were attributed as various species of Bothriospondylus, and over the years, due to various rivalries between Owen, Seeley, Mantell, Lydekker, Von Huene, Nopcsa and many others, the panoply of ascribed remains filtered through countless reclassifications as new forms of Cetiosaurus, Oplosaurus, Ornithopsis, Pelorosaurus, and Bothriospondylus. Most of the bones that made up this confusing tangle had no relation to each other, and could be said to represent entire Jurassic and Early Cretaceous faunas jumbled together. Some were unmistakable brachiosaurids, others were basal "cetiosaurid"-grade eusauropods, some were rebbachisaurs referred from Argentina (e.g. Nopcsaspondylus), and yet others were random bits of Iguanodon thrown in by Sir Owen for good measure and the sake of obligatory Crystal Palace imperial nationalism. Various remains, whether in England, Madagascar, or South America, were hastily referred to this genus by later paleontologists, all before modern taxonomy and cladistics had come into being. Bothriospondylus madagascariensis formed a chimera of nearly every large non-titanosaur sauropod found in Madagascar (the titanosaur remains were, not surprisingly, all thrown into Titanosaurus without much analysis either).

In Madagascar, Lydekker and his French counterpart Armand Thevenin collected most of the bones from the larger B. madagascariensis specimens currently included in Lapparentosaurus. Many of these are centra, missing most of the neural arch, as well as limb bones from a few large specimens. They do not appear to be juveniles due to remaining small parts of the neural arch being fused to the centrum. In 1943, Albert-Felix de Lapparent, a French paleontologist and Catholic priest who would later gain fame for dinosaur discoveries all over Africa and southern Europe, unearthed a far more complete specimen in France and also referred it to Bothriospondylus. This animal is a distinct brachiosaur species, and not related to the Madagascar material. In 1955, Lavocat referred a specimen which is now known to be very different from Lapparentosaurus, with no less than five carpal bones and a much more primitive thumb unit that had the short metacarpal and huge swiveling scythe-claw common in more basal macronarians, in addition to having well-developed secondary phalanges, another primitive trait. More discoveries by Collignon (1953) and de Ricqles (1968) filled in the gaps, though with material which may actually belong to yet another genus. In 1975, Annie Ogier, a PhD student in the University of Paris, both increased and frustrated our understanding of this animal when she described a plethora of juvenile remains and assigned MAA 91-92, a pair of posterior dorsal neural arches, as the holotype of B. madagascariensis (despite the fact that Lydekker had already assigned the fragmentary mid-dorsal vertebra BMNH R2598 as the holotype in 1895). Ogier's findings were only documented in her PhD thesis, which was never published and exists in only a few copies in Paris libraries. She seems to have since disappeared from the paleontological profession since then, and never published further papers, thus depriving many researchers of the photographic evidence for most of these specimens.

In 1986 José Bonaparte analyzed the B. madagascariensis specimens and redescribed most of the material, including Ogier's juvenile bones, as Lapparentosaurus, in honor of Albert-Felix de Lapparent, although Lapparent had never handled the Madagascar material. To add a further irony, he chose Ogier's MAA 91-92 as the holotype instead of Lydekker's original material (which though fragmentary, is nevertheless good enough to be diagnostic of a basal Europasaurus-grade brachiosaur).

After Lavocat's material and the French and South American specimens were thrown out in recent years, what remains is a more or less unified Lapparentosaurus at the basal end of Brachiosauridae, closest to Europasaurus, and more derived than Atlasaurus, Volkheimeria, the Klamelisaurids, and whatever Lavocat's bizarre 5-carpal bone species turns out to be. The material currently placed in Lapparentosaurus represents three main individuals and as many as seven less complete ones from multiple growth stages. What is shown here is only the material which is known from available photographs, published or otherwise, or could otherwise be reasonably figured from the specimens still referred to this taxon. This may be only a small sample of the valid Lapparentosaurus material lying in museum vaults in London, Paris, or elsewhere. Lapparentosaurus is also one of the first dinosaurs to be subjected to histological age analysis, and in 1983 it was determined that one mature individual was 45 years old, and still had not stopped growing. Sexual maturity in big sauropods thus took place long before "full size range" was reached. It was also shown in 1995 to have been relatively fast-growing given the presence of a large amount of fibrolamellar and harversian bone, which firmly established Bakker's theory that sauropods were warm-blooded.

References:

R. Lydekker, 1895, "On bones of a sauropodous dinosaur from Madagascar", Quarterly Journal of the Geological Society of London 51: 329-336

J.F. Bonaparte, 1986, "Les dinosaures (Carnosaures, Allosauridés, Sauropodes, Cétosauridés) du Jurassique Moyen de Cerro Cóndor (Chubut, Argentina)", Annales de Paléontologie (Vert.-Invert.) 72(3): 325-386

de Lapparent, A.F., 1943, "Les dinosauriens jurassiques de Damparis (Jura)", Mémoires de la Société Géologique de France 47: 5–20

Mannion, P.D., 2010 "A revision of the sauropod dinosaur genus 'Bothriospondylus' with a redescription of the type material of the middle Jurassic form 'B. madagascariensis'", Palaeontology, 53(2): 277–296

Ogier, A., 1975. Etude de nouveaux ossements de Bothriospondylus (Sauropode) d’un gisement du Bathonien de Madagascar. Unpublished PhD Thesis, Universite´ de Paris, 102 pp.

de Ricqlès, A., 1983, "Cyclical growth in the long bones of a sauropod dinosaur", Acta Palaeontologia Polonica 28: 225-232

Rimblot-Baly, F., A. de Ricqlès, & L. Zylberberg, 1995, "Analyse paléohistologique d'une série de croissance partielle chez Lapparentosaurus madagascariensis (Jurassique Moyen): essai sur la dynamique de croissance d'un dinosaure sauropode", Annales de Paléontologie 81: 49–86

Thevenin, A. 1907. Paleontologie de Madagascar. Annales de Paleontologie, 2, 121–136.
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ForbiddenParadise64's avatar
So was Madagascar an island at the time. That leaves interesting implications for Island fauna. 
Paleo-King's avatar
Actually in the Jurassic it appears to have been attached to Africa. Which makes sense, as Africa had no shortage of brachiosaurs. As mainland megafauna, Lapparentosaurus doesn't seem all that unusual. Lavocat's sauropod is much weirder, but this is probably due to being an early primitive radiation rather than being island fauna.
ForbiddenParadise64's avatar
Ah right. So what would be the largest Madagascan fauna from when it was an island? Thanks for answering :)
Paleo-King's avatar
Probably Rapetosaurus? It's hard to tell since only a few species of titanosaurs from Cretaceous Madagascar have been described.
ForbiddenParadise64's avatar
Ah ok fair enough. I was just wondering how large island animals can really get. 
On the topic of very big animals, do you have any definite view on the largest dinosaur, or is it less clear (eg largest brachiosaur, titanosaur, diplodocid etc)? I just wanted to know your opinion and if any new or obscure candidates have reared their head. Thanks for the reply bye. 
Paleo-King's avatar
The largest dinosaur period is probably some sort of titanosaur. As of now.....

Here's how the biggest titanosaurs rank out in first-last place:

1. Tie between Alamosaurus (referred Mexican fibula + Fowler & Sullivan's neck centrum) and Puertasaurus (1 cervical, 1 dorsal, 2 unpublished caudals).  Both of these animals were around 120+ ft. long and probably 100 tons.
2. Tie between Argentinosaurus and the Chubut Monster paleoking.blogspot.com/2014/05… both of these animals were probably pushing 110+ ft. long and 80-90 tons
3. Tie between Ruyangosaurus (cervical rib, anterior and posteiror dorsals, additional unpublished dorsals, dorsal rib, upper femur, tibia) and "Argyrosaurus" (referred femur FMNH 13018) - probably between 75-90 tons. Ruyangosaurus may have gotten longer than 100 ft., Argyrosaurus was probably not as long but still huge at 90+ ft.
4. Tie between Dreadnoughtus (majority of skeleton) and Paralititan (partial sacrals, 2 caudals, humerus, and shoulder and arm fragments) - both are around 80-90 ft. long and 65-70 tons
5. Tie between Futalognkosaurus (much of skeleton) and "Antarctosaurus" giganteus (2 femurs, pubis fragments, other random bits) - both at around 90ft. long and 65 tons. Either one of these animals could switch places with 4th place, it's a very close call. There was a bombed-to-dust centrum from "Aegyptosaurus sp" which may also have been a possible contender.

As for the biggest non-titanosaur Somphospondyli:

1. "Huanghetitan" ruyangensis (ribs, sacrum, caudals, unpublished cervicals) - 90+ ft. and 70-80 tons?
2. The French Monster (femur, caudals, toe bones and other assorted parts) - 100+ ft. and 65-75 tons? Gracile Paluxysaurus-morph Chubutisaurid by the looks of things. Possibly tied with Sauroposeidon at 110ft. and 65 tons.
2. Yongjinglong datangi (large part of the skeleton) - 100+ ft. and 60 tons?
3. Daxiatitan binglingi (nearly complete cervical and dorsal series, sacrum and ilia, femur, scapula, isolated caudal, etc.) - 100 ft. and 50-55 tons?
4. Huanghetitan liujiaxiaensis (scapula, coracoid, sacrum, caudals etc. possibly some unpublished cervicals and dorsals) - 80 ft. and 50 tons? (hard to tell since the museum mounts appear to be largely speculative)

Largest brachiosaurs:

1. Tie between "Brachiosaurus" nougaredi and Breviparopus (if either of them are indeed brachiosaurs). Both around 120ft. long and 75+ tons?
2. Fusuisaurus zhaoi (fragments of ribs, hips, femur and first 3 caudals) - 100ft. long and 70 tons?
2. Brachiosaurus sp. (Potter Creek specimen) - 95ft. long and 60+ tons?
3. Abydosaurus macintoshi (referred adult ribs) - 95ft. long and 55+ tons?
3. Giraffatitan (HMN XV2) - 85-90ft. long and 50-55 tons?
4. Tie between "Ultrasauros" (BYU scapulacoracoid, referred partial scapula and dorsal) and "The Archbishop" (majority of skeleton) - 80-85ft. long and 40 tons?
5. Lapparentosaurus (Lydekker's giant caudals) - 79ft. long and 35 tons?
6. Lusotitan (femur, tibia, fragments of hips, dorsals, ribs, caudals, fibula, humeri, scapula) - 72ft. long and 30+ tons?

Largest mamenchisaurs:

1. M. sinocanadorum (116ft. long and 60-70 tons?)
2. Chuangjiesaurus (90ft. long and 50 tons?)
3. M. jingyanensis (85-90ft. long and 50 tons?)
4. Hudiesaurus sinojapanorum (80+ft. long and 35-40 tons?)
5. M. anyuensis?????
6. "Omeisaurus" jiaoi (80ft. long, 20 tons?)

Largest Diplodocoids:

1. "Amphicoelias" fragillimus (as per Zach Armstrong's latest reconstruction, most likely a basal rebbachisaur-like diplodocoid at 95ft. long and 55+ tons - a far cry from previous estimates using Diplodocus-like proportions) - may be similar in size to Parabrontopodus distercii.
2. Tie between Supersaurus (majority of skeleton from multiple specimens) - 120ft. long and 50 tons
3. Apatosaurus ajax (Oklahoma specimen) - 95ft. long and 45 tons
4. Probably some sort of African barosaurine, Tornieria sp. or other.... who knows lol.


And then of course with basal sauropods you have the famed Turiasaurus which lags in at 80ft. and MAYBE 30 tons. Its size has been greatly exaggerated and basically it just looks like an oversized Camarasaurus with a crazy-long tail.
AcrocanthosaurusA's avatar
I mostly see 30 meters and 50 tonnes for turiasaurus.
Paleo-King's avatar
Those are outdated figures. It wasn't that big. The neck was rather short in Turiasaurids, and most of the creature's length was tail. Plus we need to consider how much slimmer the rib cage was compared to titanosauriformes.
ForbiddenParadise64's avatar
Hmmm. Sounds very interestinf to me. Doesn't it say on the blog that the chubut beast is larger than argentinosaurus? Still, that's a pretty mean feat to accomplish. Also, I'm interested in the idea of sauroposeidon not being a brachiosaur after all, which makes it a lot more mysterious. Wasn't it 17-18m tall and 60 tonnes approximately? And your estimate of Breviparopis at 22m tall is ALS pretty remarkable- I admire your work. 
Whats this news about amphicoelias, I've seen some significant debate about the creature and the new theoretical design. There are still many who think it was a huge diplodocid. Finally, in still convinced the  P. Disterci is a diplodocid due to be sheer size of the footprints- the largest being 1.65m in diameter. I calculated that based on a conservative diplodocus, the creature would be even larger than Carpenters estimate of amphicoelias. Ie 61m long and 143 tonnes, comparing footprints using Wedel's study on footprints. That's about as light as I can imagine a sauropod with that big feet being. I wanted to know a good scientific opinion on the matters that's all. 
Paleo-King's avatar
Diplodocoids, including diplodocids (especially apatosaurines) had oversized feet relative to other sauropods of similar size. So perhaps instead of thinking of P. distercii as a scaled-up Diplodocus, the most parsimonious approach would be to think of it like a giant Apatosaurus or Supersaurus. (or maybe even a basal rebbachisaur-like animal) This way it could have huge feet without being to preposterously big. Judging by the shape of "Amphicoalias" fragillimus' neural arch, it's not the same genus as true Amphicoelias, and it's probably not even in the same family. A. fragillimus looks to be a basal diplodocoid, something like the rebbachisaurids. It's definitely much bigger than the other basal forms, but in that configuration it is nowhere near the 175-foot diplodocus-on-steroids dimensions that Ken Carpenter uses.

Not to say that A. fragillimus and P. distercii couldn't be that big, but I tend to go the most conservative approach when all you're dealing with is lost vertebrae fragments and footprints (and not very sharp ones at that).

The Chubut beast MAY be larger than Argentinosaurus ... the largest femur seen in the photos is definitely larger than the femur shaft recovered from Argentinosaurus (that one is missing both ends but even its likely reconstructed size isn't as big as the biggest Chubut femur.) However there's no guarantee that the Argentinosaurus femur was from a full grown individual (you get the same problem with the Chubut remains of course). The body is too incomplete to tell and individuals of both animals could have gotten substantially larger.
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Paleo-King's avatar
This just in: I re-checked the scaling of some of Lydekker's original finds, and it turns out the largest specimens of Lapparentosaurus actually belong to an individual that rivaled or even exceeded the Berlin Giraffatitan in size. So we're not talking about 17 tons but more like 30 tones or more for the adult mass of this animal. Whew, this "midsize" brachiosaur actually turned out to be a real whopper.
SanderKools's avatar
now I'm thinking of it, you say that the 5 carpal sauropod of Lavocat  is a more archaic sauropod, could this be Archaeodontosaurus descouensi? The sauropod of which only a jaw and teeth are known, from which it can be seen that it is very archaic (it had serrations on it's teeth).

Paleo-King's avatar
It's possible, but the long cylinder-clustered metacarpals are very brachiosaur-like so unless Archaeodontosaurus is a very primitive brachiosaur (the main specimens do not appear to be brachiosaur teeth) then I doubt it. Also the carpals, if further fused into the usual two bones, would resemble those of Atlasaurus.

We may also be dealing with an immature growth stage in the carpals, it's possible that the immature animals exhibited a primitive throwback to more basal sauropod ancestors. If this is correct, then as Lavocat's sauropod grew, the 5 carpals fused into the 2 carpals typical of brachiosaurs.

If it is a brachiosaur (its metacarpals don't look like much else), Lavocat's creature would be the most primitive one ever found, with the possible exception of Volkheimeria.
SanderKools's avatar
so it is possible that there are at least three and possible four sauropod species from Jurassic Madagascar? 
- Lapparentosaurus
- Archaeodontosaurus
- Lavocat's 5 carpal sauropod (what is there known of this specimen except a front leg?)
? The creature with teeth that have small serrations ( bsgf.geoscienceworld.org/conte… ) C&D if they don't belong to Lavocat's creature
Paleo-King's avatar
Yes this sounds right... but I suspect it's possible that last specimen does belong to Lavocat's creature. Aside from the serrations the teeth do look like a brachiosaur.
SanderKools's avatar
I really wish someone would set up an expedition to fill in the gaps a bit, I get the idea that no one thinks this is interesting enough to go digging. while I have the impression that there must be very much remains lying down there, looking at the fossil market where relatively much jurassic Madagascar material can be found (even I have a few teeth and a vertebra, and I bought them in the Netherlands, which is quite a distance from Madagascar)
SpinoInWonderland's avatar
Now I know what dinosaur I would restore next....may I use this as reference for a life restoration, non-profit?

Anyway, ~20+ tonnes seems too massive judging by it's body size, I think around ~10-15 tonnes is more likely.
Paleo-King's avatar
That's fine. I'd like to see the results :)

I think some of the larger specimens may have topped 20 tons, there are some caudals that I didn't include here which may be from a 60-foot or longer individual, but scaling them is tricky because the distal tail proportions in mature individuals are not well known.

The adult reconstruction here has a similar-sized body to Camarasaurus lentus, which is 15-20 tons according to most experts. It may be on the lower end at 15 tons, though I suspect 17-18 tons is more likely.
SpinoInWonderland's avatar
You want to see the results and here they are: fav.me/d6mizij
SanderKools's avatar
Is the skull based on Europasaurus?
Paleo-King's avatar
It's actually a mix of Europasaurus and Atlasaurus with additional traces of features from all brachiosaur skulls.
SanderKools's avatar
thank you for the explanation
Kazuma27's avatar
Too bad we don't have a skull...
Paleo-King's avatar
There is a jaw fragment, I know that much. There may be many more skull pieces that haven't been published. A lot of Lapparentosaurus material is apparently being sold over the internet and in other ways. And unfortunately it's never very well-documented. I try to restore every fossil I can, even if it's commercially excavated and lacks a museum number - but photos themselves are scarce.
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