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Trackway evidence for a herd of juvenile sauropods from the Late Jurassic of Portugal

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Martin Lockley at University of Colorado
Martin Lockley
Christian A. Meyer at University of Basel
Christian A. Meyer
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... In the Cabo Espichel area, several Mesozoic dinosaurian tracksites have been described: at Praia do Guincho (Papo-Seco Formation, Barremian), a natural cast of an ornithopod dinosaur track was reported (Figueiredo et al., 2017); at Boca do Chapim and Praia do Areia do Mastro, tracks and natural casts were reported (Areia do Mastro Formation; lowermost Barremian) (Figueiredo et al., 2021;Figueiredo et al., 2022); Praia do Cavalo (Upper Jurassic) presents theropod tracks (Santos, 2003(Santos, , 2008; at Pedreira do Avelino Natural Monument (Upper Jurassic) a trackway of a large sauropod (Santos, 2003(Santos, , 2008) and a few pterosaur tracks (Mateus and Mil an, 2010) were reported; at Pedra da Mua (Upper Jurassic), trackways of sauropods and theropods have been reported, including evidence of gregarious behaviour based on seven parallel sauropod trackways (Lockley et al., 1994a;Santos, 2003Santos, , 2008; the Lagosteiros site (Hauterivian, Lower Cretaceous) shows trackways and tracks of theropods and ornithopods (Antunes, 1976(Antunes, , 1990Carvalho and Santos, 1993;Mateus and Antunes, 2003;Figueiredo, 2004Figueiredo, , 2008Figueiredo, , 2014. In 1997, Pedreira do Avelino, Pedra da Mua and Lagosteiros were protected as Natural Monuments. ...
... Sauropods could be segregated in small herds, according to age: the juveniles could live in groups, and adults may have chosen to live alone (i. e. Lockley et al., 1986Lockley et al., , 1994aLockley et al., , 2018Myers and Fiorillo, 2009;Xing et al., 2019). The set of tracks described in this work shows that most of the sauropod tracks belong to adult individuals. ...
... Dinosaur trackways have shown that some groups of dinosaurs travelled in herds. Such social behaviour has been described from the study of sauropod tracks: in Oxfordshire (England), from the Middle Jurassic (Day et al., 2004); at Pedra da Mua (Sesimbra region, Portugal), from the Upper Jurassic (Lockley et al., 1994a); and Las Cerradicas (Teruel province, Spain), from the Lower Cretaceous (Castanera et al., 2011); from ornithopod tracks at Tereñes (Asturias, Spain), from the Upper Jurassic (Piñuela et al., 2016); from hadrosaurid dinosaur tracks at Denali National Park (Alaska), from the Upper Cretaceous (Fiorillo et al., 2014). Some tracksites have also revealed that in some cases herds protected their young by keeping them in the centres of migrating groups (i.e., Thulborn and Wade, 1989;Malkani, 2007;Diedrich, 2010;Romano et al., 2018). ...
The dinosaur tracksite from the lower Barremian of Areia do Mastro Formation (Cabo Espichel, Portugal): implications for dinosaur behavior
Article
We described a dinosaur tracksite found in the uppermost part of the Areia do Mastro Formation (lowermost Barremian, Lower Cretaceous), located at 1.5 km north of Cabo Espichel (Sesimbra, Portugal). The studied tracks are distributed in a heavily trampled limestone bed which crops out alongside the rocky beach. The studied trampled surface is highly dinoturbated, 541 tracks assigned to sauropods, ornithopods and theropods were identified. The majority of footprints (336) were produced by the herbivores. The dinoturbated level is a micritic nodular limestone, deposited in a very shallow subtidal to intertidal lagoon environment. Due to the intense bioturbation and limited exposed area, it is difficult to clearly define trackways, but the tracks can provide information about the producers and their behaviours. It is inferred that dinosaurs crossed this area at different times; the herbivores (sauropods and ornithopods) may possibly used the coastal area as passage between feeding spots, while carnivores frequented the area to hunt in groups or individually.
... Second, some anatomical aspects were analyzed by Thulborn (1989Thulborn ( , 1990, who points out that the relationship between the height at the hip joint and the footprint length (h/L) varies in systematic fashion among dinosaur taxa, and that this h/L ratio certainly changes during ontogeny, on account of the allometric growth that prevails in terrestrial vertebrates. However, Alexander's method continues being valid in trackways speed estimations (e.g., Moratalla et al. 1988;Lockley et al. 1994b;Castanera et al. 2011). In an attempt to make it more precise, several authors have made modifications to the original equation (Thulborn 1982(Thulborn , 1989(Thulborn , 1990Thulborn and Wade 1984;Lockley 1986;Alexander 1991;Christiansen 1998;Henderson 2003;. ...
... The gregarious behavior has been analyzed through taphonomy studies in bone accumulations (Coria 1994;Winkler et al. 2000;Bandyopadhyay et al. 2002;González Riga and Astini 2007;Carballido et al. 2017;González Riga et al. 2018), in nests and nesting sites (Chiappe et al. 2004;Chiappe and Coria 2013), and in tracks and trackway associations. The latter constitutes one of the best evidences of herding behavior in sauropods (e.g., Bird 1944;Ostrom 1985;Lockley et al. 1986Lockley et al. , 1994bPittman and Gillette 1989;Barnes and Lockley 1994;Pittman and Lockley 1994;Schulp and Brokx 1999;Lockley and Meyer 2000;Day et al. 2004;Myers and Fiorillo 2009;Castanera et al. 2011Castanera et al. , 2012Castanera et al. , 2014Plotnick 2012). The association of different trackways can provide information about herd speed, direction, size and structure (relative position of juveniles and adults during their movement), and their composition (juvenile-adult relationship) (Lockley 1986;Myers and Fiorillo 2009). ...
Sauropod Ichnology: Overview and New Research Lines from a South American Perspective
Chapter
  • Apr 2022
Forty years ago, L. Branisa and G. Leonardi discovered the first sauropodomorph tracks in South America during expeditions to Toro Toro (Bolivia). Since then, numerous findings, mainly in Argentina, Bolivia, and Brazil, have increased the record. The first research lines mainly covered morphological description, ichnotaxonomic identification, and behavioral analyzes (e.g., gregariousness and speed). Some Cretaceous tracksites allowed the description of three new ichnotaxa: Sauropodichnus giganteus (Calvo in Ameghiniana 28:241–258, 1991), Titanopodus mendozensis (González Riga and Calvo in Palaeontology 52:631–640, 2009), and Calorckosauripus lazari (Meyer et al. in Ann Soc Geol Pol 88:223–241, 2018), the former two corresponding to Argentina, and the latter to Bolivia. A new research line named ‘ichnology and comparative anatomy’ has become relevant in the last years linking the skeletal information to the ichnological record, thus providing an integral interpretation of the set. This kind of approach allowed making more accurate inferences about paleoecological aspects, including limb posture, gauge, gait, speed, and size diversity recorded. In sum, this chapter aims to provide an overview of the South American sauropodomorph ichnotaxonomy, the history of the discoveries, and the results of new research lines in development.
... Second, some anatomical aspects were analyzed by Thulborn (1989Thulborn ( , 1990, who points out that the relationship between the height at the hip joint and the footprint length (h/L) varies in systematic fashion among dinosaur taxa, and that this h/L ratio certainly changes during ontogeny, on account of the allometric growth that prevails in terrestrial vertebrates. However, Alexander's method continues being valid in trackways speed estimations (e.g., Moratalla et al. 1988;Lockley et al. 1994b;Castanera et al. 2011). In an attempt to make it more precise, several authors have made modifications to the original equation (Thulborn 1982(Thulborn , 1989(Thulborn , 1990Thulborn and Wade 1984;Lockley 1986;Alexander 1991;Christiansen 1998;Henderson 2003;. ...
... The gregarious behavior has been analyzed through taphonomy studies in bone accumulations Winkler et al. 2000;Bandyopadhyay et al. 2002;, in nests and nesting sites Chiappe and Coria 2013), and in tracks and trackway associations. The latter constitutes one of the best evidences of herding behavior in sauropods (e.g., Bird 1944;Ostrom 1985;Lockley et al. 1986Lockley et al. , 1994bPittman and Gillette 1989;Barnes and Lockley 1994;Pittman and Lockley 1994;Schulp and Brokx 1999;Lockley and Meyer 2000;Day et al. 2004;Castanera et al. 2011Castanera et al. , 2012Castanera et al. , 2014Plotnick 2012). The association of different trackways can provide information about herd speed, direction, size and structure (relative position of juveniles and adults during their movement), and their composition (juvenile-adult relationship) (Lockley 1986;). ...
Last Titans: Titanosaurs From the Campanian–Maastrichtian Age
Chapter
  • Apr 2022
After the extinction of rebbachisaurids during the Cenomanian–Turonian interval, titanosaurs were the only group of sauropods to face the K–Pg event. This same global pattern also holds for the end-Cretaceous (Campanian–Maastrichtian) titanosaur record in South America, where their remains can be found from southern Argentina to Ecuador, with more frequent findings in Argentina and Brazil. In this chapter, we review these fossil findings and the main aspects of the taxonomy, systematics, and paleogeographic implications of this record and briefly discuss the importance of these occurrences for the understanding of titanosaur evolution. The diversity and abundance of end-Cretaceous titanosaur taxa in South America represent about 25% of the known Titanosauria species in the world, which makes them the most common group of large terrestrial herbivores of that time. Cretaceous titanosaurs from South America also vary highly in morphology and size, comprising small to large-sized taxa, for example. Their record mainly consists of appendicular and axial remains, including rare skull material, but also comprises eggs, nests, footprints, and coprolites. In South America, by the end of the Late Cretaceous, titanosaurs were generally represented by more derived titanosaurians that are mainly taxonomically assigned to more derived species within Aeolosaurini and Saltasaurinae.
... The multiple Mussaurus aggregations in the Early Jurassic breeding ground of the Laguna Colorada Formation are interpreted as the oldest skeletal evidence of structured age-segregated gregariousness amongst dinosaurs, pre-dating by over 40 million years reports from Late Jurassic and Cretaceous neosauropods [24][25][26] . Our new findings on Mussaurus adds important insights on the social behaviour of early sauropodomorphs. ...
Earliest evidence of herd-living and age segregation amongst dinosaurs
Article
Full-text available
  • Oct 2021
Sauropodomorph dinosaurs dominated the herbivorous niches during the first 40 million years of dinosaur history (Late Triassic–Early Jurassic), yet palaeobiological factors that influenced their evolutionary success are not fully understood. For instance, knowledge on their behaviour is limited, although herding in sauropodomorphs has been well documented in derived sauropods from the Late Jurassic and Cretaceous. Here we report an exceptional fossil occurrence from Patagonia that includes over 100 eggs and skeletal specimens of 80 individuals of the early sauropodomorph Mussaurus patagonicus, ranging from embryos to fully-grown adults, with an Early Jurassic age as determined by high-precision U–Pb zircon geochronology. Most specimens were found in a restricted area and stratigraphic interval, with some articulated skeletons grouped in clusters of individuals of approximately the same age. Our new discoveries indicate the presence of social cohesion throughout life and age-segregation within a herd structure, in addition to colonial nesting behaviour. These findings provide the earliest evidence of complex social behaviour in Dinosauria, predating previous records by at least 40 My. The presence of sociality in different sauropodomorph lineages suggests a possible Triassic origin of this behaviour, which may have influenced their early success as large terrestrial herbivores.
... A chapel (Fig. 2B), which quickly became a national pilgrimage site, was built at the top of the cliff in 1410 (Antunes and Mateus, 2003). The cliff has at least 38 sauropod trackways and two theropod trackways on its face (Lockley et al., 1994;Santos et al., 2008). In the eighteenth century, a series of tile panels was placed in the chapel to illustrate the legend of Our Lady of the Mule Stone (Santos et al., 2008;Baucon et al., 2012). ...
The illustration of dinosaur tracks through time
Chapter
  • Sep 2021
Fossils have stirred the imagination globally for thousands of years, starting well before they were recognized as the remains of once-living organisms and proxies of former worlds. This volume samples the history of art about fossils and the visual conceptualization of their significance starting with biblical and mythological depictions, extending to renditions of ancient life as it flourished in long-vanished habitats, and on to a modern understanding that fossil art conveys lessons for the betterment of the human condition. The 29 papers and accompanying artwork illustrate how art about fossils has come to be a significant teaching tool not only about evolution of past life, but also about conservation of our planet for the benefit of future generations.
... Because dinosaurs hatched from eggs, thereby limiting the maximum body size of a newly hatched individual, dinosaur species characterized by large adult body sizes could span a substantial ontogenetic size range (Carpenter, 1999). Footprints thought to have been made by small (or at least immature) dinosaurs are known for several trackmaker clades (Currie and Sarjeant, 1979;Lockley et al., 1994Lockley et al., , 2006Lockley et al., , 2012Pascual-Arribas and Hern andez-Medrano, 2011;Dalman, 2012;Kim et al., 2012Kim et al., , 2018Kim et al., , 2019Fiorillo et al., 2014;Xing and Lockley, 2014;Díaz-Martínez et al., 2015a;Fiorillo and Tykoski, 2016;Castanera et al., 2020;Enriquez et al., 2021), but distinguishing footprints of a particular morphotype made by juveniles of large-bodied species from those of adults of small-bodied species remains challenging. ...
Pedal Proportions of Small and Large Hadrosaurs and Other Potentially Bipedal Ornithischian Dinosaurs
Article
Foot skeletons of small (young) hadrosaurid dinosaurs were compared with those of large (adult) hadrosaurids to assess the extent of pedal shape change during ontogeny. Foot skeletons of juvenile hadrosaurids were also compared with those of similar-sized adult, bipedal, non-hadrosaurian ornithischians to which the juvenile hadrosaurids were closer in size, to investigate the possibility that pedal shape change during hadrosaurid ontogeny would have been great enough for feet (and therefore footprints) of young hadrosaurids to have been more similar to those of small-bodied ornithischians than those of large adult hadrosaurids. Although possible allometric shape changes in hadrosaurid pedal proportions are detected, these are so subtle that the feet of young hadrosaurids are far more similar to those of adult hadrosaurs than those of small-bodied, non-hadrosaurid,ornithischians. Footprints made by conspecific hadrosaurids of different size and age are therefore likely to have been similar in shape, and footprints made by juvenile hadrosaurs are unlikely be misidentified as prints made by adults of smaller-bodied, more gracile, bipedal ornithischians.
New ornithopod footprints from the Areia do Mastro Formation (Lower Cretaceous), Espichel Cape (Portugal, Western Iberia) and their context in the Iberian ichnological ornithopod record
Article
  • Mar 2022
Two new ornithopod natural casts are reported from the Praia do Areia do Mastro site, at Espichel Cape (near Sesimbra), western central mainland Portugal (western Iberia). In this Tracksite two geological formations occur: Areia do Mastro Formation and Papo-Seco Formation (lower Barremian). This locality (Praia do Areia do Mastro) occurs in a sedimentary succession comprising limestones, marls, sands and conglomerates, interpreted as marine, lagoonal and estuarine deposits. In the Papo-Seco Formationhave yielded fossil remains of dinosaurs (teeth and bones) and other vertebrates (Figueiredo et al, 2015, 2016)... The ichnological record is composed by two footprints, one of them incomplete, preserved as natural casts in convex hiporelief. They are tridactyl, mesaxonic and wider than long. The digit impressions are short, wide and have a blunt edge. The footprints present a round and wide posterior surface and have one pad impression in each digit and a wide and subrounded metatarsophalangeal pad impression. Based on the commented features, they are assigned herein to Caririchnium isp. This ichnogenus is present in other Lower Cretaceous sites in the Iberian Peninsula and is also present in other Barremian tracksites of Portugal. In Spain, Caririchnium tracks are typical of continental settings, while in Portugal they were mainly found in estuarine paleoenvironments. This contribution enhances the knowledge of the Early Cretaceous ornithopod diversity and distribution in the Iberian Peninsula.
Dinoturbation in Upper Jurassic siliciclastic levels at Cabo Mondego (Lusitanian Basin, Portugal): evidences in a fluvial-dominated deltaic succession
Article
At Cabo Mondego (western central Portugal), the Upper Jurassic marine to coastal succession contains several stratigraphic levels preserving dinosaur footprints on the surface bedding plane, as well as convolute bedding and soft sediment injection structures interpreted as dinoturbation structures. At least nineteen new three-dimensional structures observed in cross-sections are interpreted as produced by dinosaur trampling. The identification of three-dimensional structures of dinosaur footprints provides an important complement to the information obtained from footprints preserved on single bedding surfaces, such as the substrate consistency, potential trackmaker identification, and the possibility to enhance the distinction of sauropods and tridactyl dinosaurs, and paleoenvironmental interpretations. In the lower part of the Arenitos da Boa Viagem Formation, eight levels of probable lowermost Kimmeridgian age (ca. 157–156 Ma), displaying the above-mentioned deformational structures, were analyzed in detail. They support interpretations concerning the relationship between the footprints and the substrate consistency at the time of their formation. Three distinct cohesiveness patterns, defined by the penetration of the feet from the paleosurface, are the result of different degrees of substrate cohesiveness. Identifying the trackmakers of levels belonging to the middle Oxfordian–lower Kimmeridgian has important implications for Late Jurassic ecosystem reconstructions, as the footprints observed in Cabo Mondego indicate a change in the morphotypes throughout the Upper Jurassic succession.
A reevaluation of the Late Jurassic dinosaur tracksite Barkhausen (Wiehengebirge, Northern Germany)
Article
Full-text available
  • Jul 2021
A restudy of the Barkhausen dinosaur tracksite shows that the track-bearing surface reveals considerably more detail than previously indicated, and a new map is presented, showing the trackways of nine sauropods, traveling north, possibly as a group. These are among the smallest sauropod tracks recorded in Europe. There is also evidence of two large theropods crossing the area, one moving to the south and the other to the west. Evidence of at least three other sauropods is registered in the form of isolated manus traces that represent larger individuals. Previous interpretations inferred that sauropod trackways trended south, and therefore suggested a predator chasing its prey as in the purported but controversial attack scenario claimed for the famous Paluxy River site in Texas. Based on the present study, this scenario is no longer tenable for the Barkhausen tracksite. The description of Elephantopoides barkhausensis (Kaever and Lapparent, 1974) shows that it represents a moderately wide gauge, but small manus sauropod and can be assigned under the ichnofamily label Parabrontopodidae. E. barkhausensis as originally defined was a nomen dubium , but it has since been re-described semi-formally, without renaming, we emend the description and assigned them to the ichnotaxon Parabrontopodus barkhausensis comb. nov. These tracks could have been produced by the small sauropod dinosaur taxon Europasaurus . The problematic ichnotaxon Megalosauropus teutonicus (Kaever and Lapparent, 1974), which represents a large three-toed theropod, is assigned to the recently described ichnogenus Jurabrontes from the Late Kimmeridgian of the Swiss Jura mountains as Jurabrontes teutonicus comb. nov. Furthermore, we attribute the theropod tracks from the time equivalent Langenberg quarry to the same ichnotaxon.
A diverse Late Cretaceous vertebrate tracksite from the Winton Formation of Queensland, Australia
Article
Full-text available
  • Jun 2021
The Upper Cretaceous 'upper' Winton Formation of Queensland, Australia is world famous for hosting Dinosaur Stampede National Monument at Lark Quarry Conservation Park, a somewhat controversial tracksite that preserves thousands of tridactyl dinosaur tracks attributed to ornithopods and theropods. Herein, we describe the Snake Creek Tracksite, a new vertebrate ichnoassemblage from the 'upper' Winton Formation, originally situated on Karoola Station but now relocated to the Australian Age of Dinosaurs Museum of Natural History. This site preserves the first sauropod tracks reported from eastern Australia, a small number of theropod and ornithopod tracks, the first fossilised crocodyliform and ?turtle tracks reported from Australia, and possible lungfish and actinopterygian feeding traces. The sauropod trackways are wide-gauge, with manus tracks bearing an ungual impression on digit I, and anteriorly tapered pes tracks with straight or concave forward posterior margins. These tracks support the hypothesis that at least one sauropod taxon from the 'upper' Winton Formation retained a pollex claw (previously hypothesised for Diamantinasaurus matildae based on body fossils). Many of the crocodyliform trackways indicate underwater walking. The Snake Creek Tracksite reconciles the sauropod-, crocodyliform-, turtle-, and lungfish-dominated body fossil record of the 'upper' Winton Formation with its heretofore ornithopod-and theropod-dominated ichnofossil record.
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