Lucas Fiorelli
Supervisors: Gerald Grellet-Tinner, Leonardo Salgado, and Diego Pol
Address: Entre Ríos y Mendoza s/n
CP: 5301
Anillaco, La Rioja, Argentina
Address: Entre Ríos y Mendoza s/n
CP: 5301
Anillaco, La Rioja, Argentina
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Papers by Lucas Fiorelli
Carpa Formation (Santonian, Late Creteceous) of northern Patagonia, Argentina, are decribed and reevaluated taxonomically. The type specimen is an articulated individual with hindlimb and axial remains. Microsuchus has several traits that allow to recognize it as a valid taxon (the presence of bulge lateral to the prezygapophyses of the second sacral vertebra; elongated posterior zeugopodia; proximal caudal centra with triangular cross section). Microsuchus was originally assigned as a goniopholidid based on the platycely of its vertebral centra, a character that is widely distributed among fossil crocodyliforms. A phylogenetical analysis using an expanded data matrix focusing on postcranial data was
performed in which Microsuchus and another long-limbed crocodyliform from the same location, Neuquensuchus,
were included. Microsuchus is depicted in a politomy as a basal mesoeucrocodylian, with its position as a neosuchian or notosuchian supported by a single synapomorphy on the different most parsimonious trees. Additionaly, Neuquensuchus is well nested among notosuchians, in contrast to previous hypothesis. These findings further increase the crocodyliform diversity of the Bajo de la Carpa
Formation and emphasize the relevance of its fauna in studies of the diversification of mesoeucrocodylians and notosuchians. Furthermore, the occurrence of both Microsuchus and Neuquensuchus gives evidence of a basal stock of mesoeucrocodylians with elongated limbs that was not previously reported for northern Patagonia.
developed in saltasaurines, a derived group of titanosaurs. Compared with other sauropods, some titanosaurs seem to have had very poor olfaction but would have been capable of capturing sounds in a relatively wide range of high frequencies, although not to the extent of living birds.
indicate a pristine preservation of the 50 mm thick calcium carbonate, which differs significantly from the soft shell of Chinese pterosaur eggs. Estimate of the eggshell conductance implies that the nest had a minimum moisture content of 75%. This moisture estimate, combined with geological and taphonomical data, suggests that P. guinazui may have adopted a nesting strategy similar to those of grebes and flamingos rather than being buried on land, as previously hypothesized. Moreover, our results demonstrate
that the nesting paleoenvironment of this pterosaur species was closely linked to a mesohaline lacustrine ecosystem in a basin governed by regional tectonic subsidence, a setting characteristic for the feeding and reproduction of modern flamingos.
association with kannemeyeriiform dicynodonts indicate that these large synapsids produced the communal latrines and had a gregarious behaviour comparable to that of extant megaherbivores. This is the first evidence of megaherbivore communal latrines in non-mammal vertebrates, indicating that this mammal-type behaviour was present in distant relatives of mammals, and predates its previous oldest
record by 220 Mya.
age. Recent fieldwork in the new locality of Brazo del Puma, in the lowermost levels of the Chan˜ares Formation, yielded three rhynchosaur tooth-bearing bones, which were collected five metres above the contact with the underlying
Tarjados Formation. The most complete specimen is the
posterior end of the alveolar region of a left dentary. The
dentary possesses densely packed tooth rows on the lingual
surface and medial half of the occlusal surface, showing
longitudinal Zahnreihen. The teeth of the occlusal surface
are worn flat and those of the lingual surface are organized
in multiple rows, supporting the referral of the specimen to
Rhynchosauridae. In addition, the dentary teeth are conical
to mesiodistally compressed, resembling the condition
observed in hyperodapedontines. The rhynchosaur remains
reported here are the oldest collected in Argentina and
among the oldest in South America, together with an unnamed form from Brazil. The new rhynchosaur specimens
come from levels in which dicynodonts are numerically
dominant, whereas cynodonts are considerably less abundant. Accordingly, the specimens reported here bolster
faunal differences within the Chan˜ares Formation and add
a new faunal component to this already diverse vertebrate
assemblage.
and an opportunistic environment-dependent-reproduction relationship with their paleoenvironment. The taphonomy of this monotypic nesting site is described in order to assess its biostratinomy and fossil diagenesis. The 3D spatial egg-clutch arrangement is well preserved and suggests the presence of autochthonous associations with in-situ ovipositions, without nesting trace. The egg-bearing level presents a matrix-supported packed biofabric and an “intrinsic” biogenic accumulation that implies a gregarious behavior through their preferential colonization of a specific paleohydrothermal nesting site. The biocenosis of biogenically deposited eggs, with eggshell mineralization
is altered by hydrothermal activity, and extremophile microorganisms. The thanatocenosis and time-averaging were virtually null because the eggs buried in the substrate immediately after being laid. In contrast, the taphocenosis represents the most important taphonomic feature. The eggshells display several synchronous taphonomic alterations produced by acidic hot spring solutions, while re-calcification and silicification are the most common eggshell recrystallizations. The presence of microchert, chalcedony, ~10 μm-diamteer opal-A microspheres, epidote
aggregates, several macro-structures, and fibrous clay minerals replacing parts of the eggshell and egg membranes, indicates an extreme hydrothermal alteration under exceptional conditions. The presence of fossil stromatolites, cyanobacterians, and diatoms, coupled with sedimentary structures (microfacies and hydrothermal microfabrics), and with eggs taphonomy imply that the Sanagasta nesting site represents a particular instance of taphocenosis and a Lagerstätte. The Sanagasta dinosaur nesting site is the first documented case of environment-dependent reproduction amongst neosauropods.
(Lagostomopsis) Kraglievich, and Octodontidae indet. (Rodentia, Caviomorpha). This new mammalian assemblage, together with one previously recorded, has several taxa in common with mammalian associations from Central Argentina (i.e., La Pampa Province). Among those coming from Northwestern Argentina, the major affinity is with the El Jarillal Member (Chiquimil Formation) and then the Andalhuala Formation (both in Catamarca Province). The Salicas Formation fauna is considered as latest Miocene in age until new fossil discoveries
and radioisotopic dates allow a better calibration. The fossil biota and geological data suggests that the Salicas Formation was deposited in flatlands, under warm and dry conditions. This environment was dominated by open grasslands, with forested areas near rivers and lagoons."
determine the factors that underpinned the choice of these specifi c sites. Here, we report the fi rst defi nitive evidence of a group of sauropods that nested repetitively and purposely at a Cretaceous hydrothermal site at Sanagasta, La Rioja Province, Argentina. The discovery of this new colonial nesting locality shows nest fi delity over a long time, and a symbiotic relationship between egg clutches and a peculiar hydrothermal environment that favoured their incubation. Sedimentary and geochemical analyses of 80 clutches and their large eggs with thick eggshells substantiate that the Sanagasta sauropods were specifi cally using the soil moisture and thermoradiance to incubate their eggs, similar to a few extant species, namely, the megapode, Megapodius pritchardii , which is known to lay its egg clutches in burrows at volcanically heated nesting grounds.
with prominent ventral keel and well developed anteroventral parapophyses. As in basal crocodylomorphs, it possesses two sacral vertebrae. Also, a much enlarged scapular blade, with well developed acromial ridge and the posterior edge similar to Sichuanosuchus. The pronounced deltopectoral crest in the complete humerus is equivalent to Sichuanosuchus and as this, a circular, elongated and thin shaft with the medial condyle longer than the lateral one. Also, the complete ulna and radius is similar in their proportions to Sichuanosuchus. As this, the pubis is lengthened, very thin in the half section and not very expanded distally. The femur, tibia and fibula are elongated and similar to other non-derivated crocodyliforms. Besides
representing the first Cretaceous “protosuchian” of Gondwana, the occurrence of these outside of Asia and Europe during the Cretaceous offers new evidence of pre-Albian dispersion between Gondwana and Central Asia through Europe.
Carpa Formation (Santonian, Late Creteceous) of northern Patagonia, Argentina, are decribed and reevaluated taxonomically. The type specimen is an articulated individual with hindlimb and axial remains. Microsuchus has several traits that allow to recognize it as a valid taxon (the presence of bulge lateral to the prezygapophyses of the second sacral vertebra; elongated posterior zeugopodia; proximal caudal centra with triangular cross section). Microsuchus was originally assigned as a goniopholidid based on the platycely of its vertebral centra, a character that is widely distributed among fossil crocodyliforms. A phylogenetical analysis using an expanded data matrix focusing on postcranial data was
performed in which Microsuchus and another long-limbed crocodyliform from the same location, Neuquensuchus,
were included. Microsuchus is depicted in a politomy as a basal mesoeucrocodylian, with its position as a neosuchian or notosuchian supported by a single synapomorphy on the different most parsimonious trees. Additionaly, Neuquensuchus is well nested among notosuchians, in contrast to previous hypothesis. These findings further increase the crocodyliform diversity of the Bajo de la Carpa
Formation and emphasize the relevance of its fauna in studies of the diversification of mesoeucrocodylians and notosuchians. Furthermore, the occurrence of both Microsuchus and Neuquensuchus gives evidence of a basal stock of mesoeucrocodylians with elongated limbs that was not previously reported for northern Patagonia.
developed in saltasaurines, a derived group of titanosaurs. Compared with other sauropods, some titanosaurs seem to have had very poor olfaction but would have been capable of capturing sounds in a relatively wide range of high frequencies, although not to the extent of living birds.
indicate a pristine preservation of the 50 mm thick calcium carbonate, which differs significantly from the soft shell of Chinese pterosaur eggs. Estimate of the eggshell conductance implies that the nest had a minimum moisture content of 75%. This moisture estimate, combined with geological and taphonomical data, suggests that P. guinazui may have adopted a nesting strategy similar to those of grebes and flamingos rather than being buried on land, as previously hypothesized. Moreover, our results demonstrate
that the nesting paleoenvironment of this pterosaur species was closely linked to a mesohaline lacustrine ecosystem in a basin governed by regional tectonic subsidence, a setting characteristic for the feeding and reproduction of modern flamingos.
association with kannemeyeriiform dicynodonts indicate that these large synapsids produced the communal latrines and had a gregarious behaviour comparable to that of extant megaherbivores. This is the first evidence of megaherbivore communal latrines in non-mammal vertebrates, indicating that this mammal-type behaviour was present in distant relatives of mammals, and predates its previous oldest
record by 220 Mya.
age. Recent fieldwork in the new locality of Brazo del Puma, in the lowermost levels of the Chan˜ares Formation, yielded three rhynchosaur tooth-bearing bones, which were collected five metres above the contact with the underlying
Tarjados Formation. The most complete specimen is the
posterior end of the alveolar region of a left dentary. The
dentary possesses densely packed tooth rows on the lingual
surface and medial half of the occlusal surface, showing
longitudinal Zahnreihen. The teeth of the occlusal surface
are worn flat and those of the lingual surface are organized
in multiple rows, supporting the referral of the specimen to
Rhynchosauridae. In addition, the dentary teeth are conical
to mesiodistally compressed, resembling the condition
observed in hyperodapedontines. The rhynchosaur remains
reported here are the oldest collected in Argentina and
among the oldest in South America, together with an unnamed form from Brazil. The new rhynchosaur specimens
come from levels in which dicynodonts are numerically
dominant, whereas cynodonts are considerably less abundant. Accordingly, the specimens reported here bolster
faunal differences within the Chan˜ares Formation and add
a new faunal component to this already diverse vertebrate
assemblage.
and an opportunistic environment-dependent-reproduction relationship with their paleoenvironment. The taphonomy of this monotypic nesting site is described in order to assess its biostratinomy and fossil diagenesis. The 3D spatial egg-clutch arrangement is well preserved and suggests the presence of autochthonous associations with in-situ ovipositions, without nesting trace. The egg-bearing level presents a matrix-supported packed biofabric and an “intrinsic” biogenic accumulation that implies a gregarious behavior through their preferential colonization of a specific paleohydrothermal nesting site. The biocenosis of biogenically deposited eggs, with eggshell mineralization
is altered by hydrothermal activity, and extremophile microorganisms. The thanatocenosis and time-averaging were virtually null because the eggs buried in the substrate immediately after being laid. In contrast, the taphocenosis represents the most important taphonomic feature. The eggshells display several synchronous taphonomic alterations produced by acidic hot spring solutions, while re-calcification and silicification are the most common eggshell recrystallizations. The presence of microchert, chalcedony, ~10 μm-diamteer opal-A microspheres, epidote
aggregates, several macro-structures, and fibrous clay minerals replacing parts of the eggshell and egg membranes, indicates an extreme hydrothermal alteration under exceptional conditions. The presence of fossil stromatolites, cyanobacterians, and diatoms, coupled with sedimentary structures (microfacies and hydrothermal microfabrics), and with eggs taphonomy imply that the Sanagasta nesting site represents a particular instance of taphocenosis and a Lagerstätte. The Sanagasta dinosaur nesting site is the first documented case of environment-dependent reproduction amongst neosauropods.
(Lagostomopsis) Kraglievich, and Octodontidae indet. (Rodentia, Caviomorpha). This new mammalian assemblage, together with one previously recorded, has several taxa in common with mammalian associations from Central Argentina (i.e., La Pampa Province). Among those coming from Northwestern Argentina, the major affinity is with the El Jarillal Member (Chiquimil Formation) and then the Andalhuala Formation (both in Catamarca Province). The Salicas Formation fauna is considered as latest Miocene in age until new fossil discoveries
and radioisotopic dates allow a better calibration. The fossil biota and geological data suggests that the Salicas Formation was deposited in flatlands, under warm and dry conditions. This environment was dominated by open grasslands, with forested areas near rivers and lagoons."
determine the factors that underpinned the choice of these specifi c sites. Here, we report the fi rst defi nitive evidence of a group of sauropods that nested repetitively and purposely at a Cretaceous hydrothermal site at Sanagasta, La Rioja Province, Argentina. The discovery of this new colonial nesting locality shows nest fi delity over a long time, and a symbiotic relationship between egg clutches and a peculiar hydrothermal environment that favoured their incubation. Sedimentary and geochemical analyses of 80 clutches and their large eggs with thick eggshells substantiate that the Sanagasta sauropods were specifi cally using the soil moisture and thermoradiance to incubate their eggs, similar to a few extant species, namely, the megapode, Megapodius pritchardii , which is known to lay its egg clutches in burrows at volcanically heated nesting grounds.
with prominent ventral keel and well developed anteroventral parapophyses. As in basal crocodylomorphs, it possesses two sacral vertebrae. Also, a much enlarged scapular blade, with well developed acromial ridge and the posterior edge similar to Sichuanosuchus. The pronounced deltopectoral crest in the complete humerus is equivalent to Sichuanosuchus and as this, a circular, elongated and thin shaft with the medial condyle longer than the lateral one. Also, the complete ulna and radius is similar in their proportions to Sichuanosuchus. As this, the pubis is lengthened, very thin in the half section and not very expanded distally. The femur, tibia and fibula are elongated and similar to other non-derivated crocodyliforms. Besides
representing the first Cretaceous “protosuchian” of Gondwana, the occurrence of these outside of Asia and Europe during the Cretaceous offers new evidence of pre-Albian dispersion between Gondwana and Central Asia through Europe.