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Patagotitan
Patagotitan
Patagotitan
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ARTICLE IN PRESS
1
2 A new giant titanosaur sheds light
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4
5
rspb.royalsocietypublishing.org on body mass evolution among
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sauropod dinosaurs
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9 Jose L. Carballido1, Diego Pol1, Alejandro Otero2, Ignacio A. Cerda3,
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Research Leonardo Salgado3, Alberto C. Garrido4, Jahan Ramezzani5, Nestor R. Cuneo1
12 Cite this article: Carballido JL, Pol D, Otero A, and Marcelo J. Krause1
13
Cerda IA, Salgado L, Garrido AC, Ramezzani J, 1
14 CONICET, Museo Paleontologico Egidio Feruglio, Trelew U9100GYO, Argentina
Cuneo NR, Krause MJ. 2017 A new giant 2
CONICET, Division Paleontologa de Vertebrados, Museo de La Plata, La Plata B1900FWA, Argentina
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titanosaur sheds light on body mass evolution 3
16 CONICET, Instituto de Investigacion en Paleobiologa y Geologa, Universidad Nacional de Ro Negro,
17 among sauropod dinosaurs. Proc. R. Soc. B General Roca 8332, Argentina
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Museo Provincial de Ciencias Naturales Juan Olsacher, Zapala 8340, Argentina
18 20171219. 5
Department of Earth, Atmospheric and Planetary Sciences, Massachusetts Institute of Technology,
19 http://dx.doi.org/10.1098/rspb.2017.1219 Cambridge, MA 02139, USA
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21 JLC, 0000-0003-3227-8034
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23 Titanosauria was the most diverse and successful lineage of sauropod dino-
Received: 31 May 2017
24 saurs. This clade had its major radiation during the middle Early Cretaceous
Accepted: 6 July 2017 and survived up to the end of that period. Among sauropods, this lineage
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26 has the most disparate values of body mass, including the smallest and
27 largest sauropods known. Despite recent findings have improved our knowl-
28 edge on giant titanosaur anatomy, there are still many unknown aspects about
29 Subject Category: their evolution, especially for the most gigantic forms and the evolution of
30 body mass in this clade. Here we describe a new giant titanosaur, which
Palaeobiology
31 represent the largest species described so far and one of the most complete
32 titanosaurs. Its inclusion in an extended phylogenetic analysis and the optim-
Subject Areas: ization of body mass reveals the presence of an endemic clade of giant
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34
palaeontology, evolution, taxonomy titanosaurs inhabited Patagonia between the Albian and the Santonian. This
35 and systematics clade includes most of the giant species of titanosaurs and represents the
36 major increase in body mass in the history of Titanosauria.
37 Keywords:
38 Titanosauria, Cretaceous, Patagonia,
39 taphonomy, Gondwana
40 1. Introduction
41 The origin of titanosaur dinosaurs origin was inferred to date back to the latest
42 Jurassic [1] (ca 150 Ma) and recent analyses place their time of diversification
43 Author for correspondence: during the Early Cretaceous (BarriasianBarremian ca 140125 Ma) [27].
44 Jose L. Carballido Titanosauria is a diverse clade (approx. 90 genera) of sauropods and also the
45 e-mail: jcarballido@mef.org.ar only one that survived until the end-Cretaceous. In addition to their taxonomic
46 diversity (approx. 30% of valid sauropod genera) this clade has a remarkable vari-
47 ation on body size among its members, including the largest known sauropods
48 (e.g. Argentinosaurus, Puertasaurus; with rough estimates of body mass above 60
49 tons [8]) and the smallest sauropods known to date (e.g. Rinconsaurus, Saltasaurus;
50 with estimated body masses of approximately 6 tons [8]). Understanding the
51 evolution of body mass in this clade is key for understanding many aspects of
52 sauropod evolution, including changes in their paleoecology and growth strat-
53 egies. Studies on dinosaur body mass evolution have noted an increase in
54 variation of body size among titanosaurs during the Cretaceous in comparison
55 with other sauropod groups, and some previous studies noted an overall trend
56 to body mass decrease [810]. Nonetheless, recent studies failed to find any
57 exceptional body mass shifts in sauropod evolution during the Cretaceous, a
58 result interpreted as product of niche saturation achieved during the Triassic
59 Jurassic [8]. A common problem in studies on the evolution of body size in
Electronic supplementary material is available
60 Titanosauria has been the lack of well sampled phylogenetic studies of the
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online at rs.figshare.com. group [11,12], especially considering large bodied titanosaurs, whose record
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& 2017 The Author(s) Published by the Royal Society. All rights reserved.
rspb.royalsocietypublishing.org
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holotype (MPEF-PV 3400)
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paratype (MPEF-PV 3399)
69 additional paratypes
missing elements 5m
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Figure 1. Reconstructed skeleton and body silhouette of Patagotitan mayorum showing preserved elements from the holotype and paratypic specimens.
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74 has been extremely fragmentary for many years [13,14]. Recent (c) Paratypes
75 discoveries of more complete remains of giant titanosaurs The repeated elements recovered at this site include both axial
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129 with vertical neural spine (reversal to the condition of non- compressed. The presence of laterally expanded neural
130 titanosauriforms, convergently acquired in some lithostrotian spines in posterior cervical vertebrae (as in Futalognkosaurus
131 titanosaurs); first caudal vertebra with flat anterior and convex [15], Mendozasaurus [17] and Ligabuesaurus [21]) cannot be
132 posterior articular surfaces; anterior caudal vertebrae with determined in Patagotitan due to poor preservation. Posterior
133 neural spines four to six times mediolaterally wider than antero- cervical vertebrae and all dorsal vertebrae bear deep pleuro-
134 posteriorly long; anterior caudal neural spines incipiently bifid coels that open into the internal camellate bone tissue. Based
135 with anteriorly directed tips; marked bulge on the posterolateral on the morphological differences among the dorsal vertebrae
136 surface of the humerus (convergently acquired in some derived of Patagotitan, we identify that nine distinct vertebral pos-
137 lithostrotians); straight lateral edge on distal femur. itions are represented. Based on comparisons with dorsal
138 series of other titanosaurs (and assuming a total number of
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192 SPRL
PODL SPOL
193
SPOL
194 SPDL SPOL
SPRL PRDL
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196
197
CPOL
198 dp
199 PCDL
CPRL
200
pp
pp
201 PCDL
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255 caudal vertebrae, but its presence cannot be determined in straight lateral margin along its proximal third, and flat distal
256 between these two regions due to the lack of preserved condyles. One distinctive character observed in the three recov-
257 elements. The transverse process of Patagotitan is somewhat ered humeri is the presence of a well-developed bulge on the
258 similar to the wing-like process of diplodocoids but differ posterolateral surface (figure 2r), resembling that of some
259 from those in certain aspects. The dorsal margin of the trans- derived lithostrotians (e.g. Neuquensaurus, Opisthocoelicaudia
260 verse process is much shorter than the ventral margin [31]). The triradiate ulna has a deep radial fossa, and the olecra-
261 in Patagotitan (figure 2hk), whereas diplodocoids have a non process is weakly developed. The distal condyles of the
262 remarkably long dorsal margin, that is similar in length to radius are perpendicular to its long axis.
263 the ventral margin and is dorsolaterally oriented [27]. The pubis is larger than the ischium, and the puboischial
264 The morphology of the dorsoventrally high and antero- articulation is almost half the pubic length. The ischium has a
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318 equation (e.g. Giraffatitan, Apatosaurus; [8]). Uncertainties in supplementary material).
319 large dinosaur body mass estimations from scaling equations The analysis resolved Patagotitan as the titanosaur sauro-
320 are notably large and errors bars in the cases of Patagotitan pod most closely related to Argentinosaurus, a clade that is
321 and Dreadnoughtus overlap between 59 tons (minimal body supported by a single unambiguous synapomorphic character,
322 mass for Patagotitan) and 74 tons (maximal body mass for the presence of anterior dorsal vertebrae with elongated neural
323 Dreadnoughtus). The completeness of Patagotitan allowed the spines (neural spine is more than twice as dorsoventrally
324 reconstruction of volumetric models (following the method high as mediolaterally wide; character (ch.) 174). These two
325 described by [38]) and yielded even larger differences between giant titanosaurs are nested within Lognkosauria, a diverse
326 Patagotitan and other sauropods (including Dreadnoughtus). clade that includes most of the largest titanosaurs known.
327 Volumetric body mass estimates also have uncertainties related The successive sister taxa of Lognkosauria are Notocolossus
Opisthocoelicaudia
7
380 body mass
Neuquensaurus
Rapetosaurus
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Alamosaurus
381 6
Saltasaurus
382
lsisaurus
383
384 70 Rinconsauria
385 60 unknown
Baurutitan
Rinconsaurus
Aeolosaurus
386
Overosaurus
387
Futalognkosaurus
70
Mendozasaurus
388
Malarguesaurus
Bonitasaura
Muyelensaurus
Notocolossus
389
Epachthosaurus
Quetecsaurus
390
Argentinosaurus
Dreadnoughtus
Drusilasaura
Puertasaurus
392
time (millons years)
393
Erketu
Patagotitan
394 90
Phuwiangosaurus
395 late
Wintonotitan
396 cret
397
Chubutisaurus
100
early
Sauroposeidon
398
399 cret
Malawisaurus
Lognkosauria
Tapuiasaurus
400 110
401
402
403 120
404
405
406 Lithostrotia
Eutitanosauria
407
Titanosauria
408 length: 1328 CI: 0.37
Somphospondyli MPTs: 5500 RI: 0.72
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410
Figure 3. Phylogenetic relationships of Patagotitan mayorum and body mass evolution in Somphospondyli. Reduced strict consensus tree (excluding Sauroposeidon,
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Ligabuesaurus and Trigonosaurus) calibrated against the age. Branch colours indicates body mass inferred through scaling equation [37] and continuous character
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optimization in TNT (ancestral branches). See electronic supplementary material for complete phylogenetic analysis, body mass optimization and taxon definitions
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followed here.
414
415
416 well supported, requiring 12 extra steps to place this taxon In the present analysis, Alamosaurus, from the Maastrichtian of
417 as the sister lineage to Lognkosauria, and 22 extra steps if North America, is recovered as a derived lithostrotian. This
418 Patagotitan is forced to be a lithostrotian titanosaur. Despite taxon was recently recovered as closely related to Lognkosauria
419 the amount of missing data in Argentinosaurus (87%), six [46]. Nevertheless, in our data set (which includes character
420 extra steps are needed to place this taxon as the sister scorings based on [46]) this position is highly suboptimal,
421 lineage to Lognkosauria, and eight extra steps if it is forced to requiring 36 extra steps to place Alamosaurus as the sister
422 be inside Lithostrotia. Puertasaurus and Quetecsaurus (approx. taxon to Lognkosauria.
423 90% missing data) are the most unstable lognkosaurian Rinconsauria, the sister clade of the lineage formed by
424 taxa, and with one extra step they can be placed as sister taxa Longkosauria Notocolossus Bonitasaura is supported by
425 to Lognkosauria. Six extra steps are needed if these taxa are three unambiguous synapomorphies: (1) anterior caudal
426 forced into Lithostrotia. Drusilasaura (94% missing data) vertebrae with the anterior face of the centrum anteriorly
427 was also found to be unstable in suboptimal trees, as it requi- directed (ch. 254); (2) posterior caudal vertebrae with flat-
428 res three extra steps to place it as sister taxa to Lognkosauria tened centra (ch. 262) and (3) gracile humerus (ch. 304).
429 and eight extra steps if it is forced as the sister lineage to The original definition of Aeolosaurinae (as titanosaurs
430 Lithostrotia. A large number of extra steps are needed to force more closely related to Aeolosaurus and Gonwanatitan than
431 Futalognkosaurus (73% missing data) or Mendozasaurus (6 7% to Saltasaurus and Opisthocoelicaudia [50]; see the electronic
432 missing data) inside Lithostrotia, with 10 extra steps needed supplementary material), points to the clade formed by
433 to move Mendozasaurus and 18 for Futalognkosaurus. Moving Rinconsauria plus Lognkosauria. Nevertheless, this clade
434 Notocolossus (80% missing data) inside Lithostrotia requires was traditionally used to include a small clade of derived tita-
435 only two extra steps. This analysis reveals that Lognkosauria nosaurs closely related to the genus Aeolosaurus and a new
436 is relatively well supported except for Quetecsaurus, Drusilasaura definition (node based) of Aeolosaurinae is here proposed
437 and Puertasaurus, an expected instability given the large amount in order to keep the original aim of the definition (see the
438 of missing data in these taxa (more than 90%). Finally, placing electronic supplementary material).
439 Lognkosauria inside Lithostrotia (as in previous analyses Dreadnoughtus was here recovered as the sister taxon to
440 [15,16,49]) requires nine extra steps, indicating that the more Lithostrotia [12], and therefore not related to the other giant tita-
441 basal position here obtained for this clade is well supported. nosaurs from Patagonia. The clade formed by Dreadnoughtus
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444 (ch. 132); (2) first caudal vertebra with convex anterior articular of titanosaur body masses also includes events of marked
445 surface (ch. 224); (3) the absence of ventral bulge in anterior body mass decrease in Rinconsauria and three lineages
446 caudal vertebrae (ch. 227) and (4) the absence of a deep infragle- within Lithostrotia (including probably dwarfism events
447 noid groove in the coracoid (ch. 290). Ten extra steps are [34,36,53]: figure 3).
448 needed to place Dreadnoughtus (46% missing data) within The age of Patagotitan indicates that the radiation of lognko-
449 Lognkosauria, indicating a markedly suboptimal hypothesis. saurians (and related forms) started in the late Early Cretaceous,
450 Characters present in Lognkosauria that increase their length temporally coincident with the diversification of lithostrotians
451 when Dreadnoughtus is forced in this clade include: (1) the [4,54,55], the other major clade of titanosaurs. The phylogenetic
452 more rounded cervical centra (ch 132); (2) the vertically oriented diversification event of the major clades of eutitanosaurs in the
453 neural spine in anterior dorsal vertebrae (ch. 172); (3) the non- AptianAlbian involved major changes in body mass indica-
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507 P Dodson, H Osmolska), pp. 259322. Berkeley, CA: Titanosauria) del Cretacico Tardo de Mendoza, journal.pone.0025763)
508 University of California Press. Argentina. Ameghiniana 42, 535 548. 31. Otero A. 2010 The appendicular skeleton of
509 4. Zaher H et al. 2011 A complete skull of an early 18. Conybeare A, Haynes G. 1984 Observations on Neuquensaurus, a Late Cretaceous saltasaurine
510 cretaceous sauropod and the evolution of advanced elephant mortality and bones in water holes. Quat. sauropod from Patagonia, Argentina. Acta Paleontol.
511 titanosaurians. PLoS ONE 6, e16663. (doi:10.1371/ Res. 22, 189200. (doi:10.1016/0033- Pol. 55, 399426. (doi:10.4202/app.2009.0099)
512 journal.pone.0016663) 5894(84)90039-5) 32. Klein N, Sander M. 2008 Ontogenetic stages in
513 5. DEmic MD. 2012 The early evolution of 19. Upchurch P. 1998 The phylogenetic relationships of the long bone histology of sauropod dinosaurs.
514 titanosauriform sauropod dinosaurs. Zool. J. Linn. sauropod dinosaurs. Zool. J. Linn. Soc. 124, 43 Paleobiology 34, 247 263. (doi:10.1666/0094-
515 Soc. 166, 624671. (doi:10.1111/j.1096-3642.2012. 103. (doi:10.1111/j.1096-3642.1998.tb00569.x) 8373(2008)034[0247:OSITLB]2.0.CO;2)
516 00853.x) 20. Carballido JL, Sander PM. 2014 Postcranial axial 33. Chinsamy-Turan A. 2005 The microstructure of
rspb.royalsocietypublishing.org
570 45. Navarrete C, Casal G, Martnez R. 2011 Drusilasaura 51. Goloboff PA, Mattoni CI, Quinteros AS. 2006 58. Huber BT, Norris RD, MacLeod KG. 2002 Deep-sea
571 deseadensis gen. et sp. nov., un nuevo titanosaurio Continuous characters analyzed as such. Cladistics 22, paleotemperature record of extreme warmth during
572 (Dinosauria-Sauropoda), de la Formacion Bajo 589601. (doi:10.1111/j.1096-0031.2006.00122.x) the Cretaceous. Geology 30, 123126. (doi:10.
573 Barreal, Cretacico Superior del norte de Santa Cruz, 52. Carrano MT. 2006 Body-size evolution in the 1130/0091-7613(2002)030,0123:DSPROE.2.0.
574 Argentina. Rev. Brasilera Paleontol. 14, 1 14. Dinosauria. In Amniote paleobiology: perspectives on CO;2)
575 (doi:10.4072/rbp.2011.1.01) the evolution of mammals (eds MT Carrano, RW 59. Upchurch Jr GR, Dilcher DL. 1990 Cenomanian
576Q2 46. Tykoski RS, Fiorillo AR. 2016 An articulated cervical Blob, TJ Gaudin, JR Wible). angiosperm leaf megafossils, Dakota Formation,
577 series of Alamosaurus sanjuanensis Gilmore, 1922 53. Stein K, Csiki Z, Curry Rogers K, Weishampel DB, Rose Creek locality, Jefferson County, southeastern
578Q3 (Dinosauria, Sauropoda) from Texas: new Redelstorff R, Carballido JL, Sander PM, Stanley SM. Nebraska.
579 perspective on the relationships of North Americas 2010 Small body size and extreme cortical bone 60. Sereno PC, Wilson JA, Conrad JL. 2004 New