Pisanosaurus revisited

Reconstructed skeleton of Pisanosaurus (Royal Ontario Museum)

Pisanosaurus mertii was originally described by Argentinian paleontologist Rodolfo Casamiquela in 1967, based on a poorly preserved but articulated skeleton from the upper levels of the Ischigualasto Formation (Late Triassic). The holotype and only known specimen (PVL 2577) is a fragmentary skeleton including partial upper and lower jaws, seven articulated dorsal vertebrae, four fragmentary vertebrae of uncertain position in the column, the impression of the central portion of the pelvis and sacrum, an articulated partial hind limb including the right tibia, fibula, proximal tarsals and pedal digits III and IV, the distal ends of the right and left femora, a left scapular blade (currently lost), a probable metacarpal III, and the impressions of some metacarpals (currently lost).

Pisanosaurus mertii holotype. Right lower mandible in medial (A) and lateral (B) views. Scale bar: 5 cm. From Agnolín and Rozadilla, 2017.

In the original description, Casamiquela considered that Pisanosaurus was a very distinct ornithischian, and even proposed a family: Pisanosauridae. The dentition and tooth-bearing bones of Pisanosaurus possess a large number of ornithischian traits, like its barricade-like dentition. But Pisanosaurus shows some features that strongly differ from those of ornithischians. For instance, vertebral centra are very elongated and transversely compressed, differing from the short and stout dorsal vertebrae of known ornithischians, including heterodontosaurids. The pelvis is another portion of the skeleton of Pisanosaurus strongly different from that of ornithischians.

Pisanosaurus mertii holotype. Dorsal vertebrae in left lateral (A) and right lateral (B) views. Scale bar: 5 cm. From Agnolín and Rozadilla, 2017.

On the other hand, Pisanosaurus shows some derived traits that resulted as unambiguous synapomorphies of the Silesauridae clade, and include: reduced to absent denticles on maxillary and dentary teeth; sacral ribs shared between two sacral vertebrae; lateral side of proximal tibia with a fibular flange (present also in heterodontosaurids and several saurischians); dorsoventrally flattened ungual phalanges; and ankylothecodonty, teeth partially fused to maxilla and dentary bone. The first and last characters are lacking in ornithischians. Of course, the inclusion of Pisanosaurus within Silesauridae implies that this taxon does not constitute the oldest ornithischian. This also suggests a significant gap between Pisanosaurus and the oldest unambiguous records of ornithischians: Laquintasaura and Lesothosaurus, which may be dated as Hettangian in age. This is consistent with previous interpretations proposing that ornithischian radiation occurred after the Triassic–Jurassic boundary.

References:

Federico L. Agnolín & Sebastián Rozadilla (2017): Phylogenetic reassessment of Pisanosaurus mertii Casamiquela, 1967, a basal dinosauriform from the Late Triassic of Argentina, Journal of Systematic Palaeontology DOI: 10.1080/14772019.2017.1352623

Baron, M. G., Norman, D. B. & Barrett, P. M. A new hypothesis of dinosaur relationships and early dinosaur evolution.  Nature 543, 501–506  (2017).  doi:10.1038/nature21700

Padian K. The problem of dinosaur origins: integrating three approaches to the rise of Dinosauria. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, Available on CJO 2013 doi:10.1017/S1755691013000431 (2013).

Terrestrial floras at the Triassic-Jurassic Boundary in Europe.

Proportions of range-through diversities of higher taxonomic categories of microfloral elements over the Middle Triassic–Early Jurassic interval (From Barbacka et al., 2017)

Over the last 3 decades, mass extinction events  have become the subject of increasingly detailed and multidisciplinary investigations. Most of those events are associated with global warming and proximal killers such as marine anoxia. Volcanogenic-atmospheric kill mechanisms include ocean acidification, toxic metal poisoning, acid rain, increased UV-B radiation, volcanic darkness, cooling and photosynthetic shutdown. The mass extinction at the Triassic-Jurassic Boundary (TJB) has been linked to the eruption of the Central Atlantic Magmatic Province (CAMP), a large igneous province emplaced during the initial rifting of Pangea. Another theory is that a huge impact was the trigger of the extinction event. At least two craters impact were reported by the end of the Triassic. The Manicouagan Impact crater in the Côte-Nord region of Québec, Canada was caused by the impact of a 5km diameter asteroid, and it was suggested that could be part of a multiple impact event which also formed the Rochechouart crater in France, Saint Martin crater in Canada, Obolon crater in Ukraine, and the Red Wing crater in USA (Spray et al., 1998).

Photographs of some Rhaetian–Hettangian spores and pollen from the Danish Basin (From Lindström, 2015)

Most mammal-like reptiles and large amphibians disappeared, as well as early dinosaur groups. In the oceans, this event eliminated conodonts and nearly annihilated corals, ammonites, brachiopods and bivalves. In the Southern Hemisphere, the vegetation turnover consisted in the replacement to Alisporites (corystosperm)-dominated assemblage to a Classopollis (cheirolepidiacean)-dominated one. But there was no mass extinction of European terrestrial plants during the TJB. The majority of genera and a high percentage of species still existed in its later stages, and replacement seems to have been local, explainable as a typical reaction to an environmental disturbance. In Greenland, for example, the replacement of Triassic wide-leaved forms with Jurassic narrow-leaved forms was linked to the reaction of plants to increased wildfire. In Sweden, wildfire in the late Rhaetian and early Hettangian caused large-scale burning of conifer forests and ferns, and the appearance of new swampy vegetation. In Austria and the United Kingdom, conifers and seed ferns were replaced by ferns, club mosses and liverworts. In Hungary, there was a high spike of ferns and conifers at the TJB, followed by a sudden decrease in the number of ferns along with an increasing share of swamp-inhabiting conifers.

Although certain taxa/families indeed became extinct by the end of the Triassic (e.g. Peltaspermales), the floral changes across Europe were rather a consequence of local changes in topography.

References:

Maria Barbacka, Grzegorz Pacyna, Ádam T. Kocsis, Agata Jarzynka, Jadwiga Ziaja, Emese Bodor , Changes in terrestrial floras at the TriassicJurassic Boundary in Europe, Palaeogeography, Palaeoclimatology, Palaeoecology (2017), doi: 10.1016/j.palaeo.2017.05.024

S. Lindström, Palynofloral patterns of terrestrial ecosystem change during the end-Triassic event — a review, Geol. Mag., 1–23 (2015) https://doi.org/10.1017/S0016756815000552

Van de Schootbrugge, B., Quan, T.M., Lindström, S., Püttmann, W., Heunisch, C., Pross, J., Fiebig, J., Petschick, R., Röhling, H.-G., Richoz, S., Rosenthal, Y., Falkowski, P. G., 2009. Floral changes across the Triassic/Jurassic boundary linked to flood basalt volcanism. Nat. Geosci. 2, 589–594. doi: 10.1038/NGEO577.

N.R. Bonis, W.M. Kürschner, Vegetation history, diversity patterns, and climate change across the Triassic/Jurassic boundary, Paleobiology, 8 (2) (2012), pp. 240–264 https://doi.org/10.1666/09071.1

Palynology of the Ischigualasto Formation.

Ischigualasto-perfil-gusano-montañas

Image from Ischigualasto Park (http://www.ischigualasto.gob.ar/)

Ischigualasto is an arid, sculpted valley, in northwest Argentina (San Juan Province), limiting to the north with the Talampaya National Park, in La Rioja Province. Both areas belong to the same geological formation: the Ischigualasto-Villa Unión Basin which is centered on a rift zone that accumulated thick terrestrial deposits during the Triassic. This basin preserves a complete and continuous fossiliferous succession of continental Triassic rocks.

The Ischigualasto Formation is known worldwide for its tetrapod assemblage, which included the oldest known record of dinosaurs. Adolf Stelzner in 1889 published the first data on the geology of Ischigualasto, but it was not until 1911, that Bondenbender briefly refers to the fossils of the site. Several thin volcanic ash horizons, indicates that the deposition of the Ischigualasto Formation began at the Carnian Stage (approximately 228 mya), and consists of four lithostratigraphic members which in ascending order include the La Peña Member, the Cancha de Bochas Member, the Valle de la Luna Member, and the Quebrada de la Sal Member.

1–3. Retusotriletes herbstii sp. nov; 4–5. Rogalskaisporites cicatricosus; 6. Rugulatisporites

1–3. Retusotriletes herbstii sp. nov; 4–5. Rogalskaisporites cicatricosus; 6. Rugulatisporites

During the Late Triassic two distinct microfloras have been recognised in the southern hemisphere: the Ipswich microflora and the Onslow microflora. The Ipswich province, characterized by the abundance of bisaccate pollen, monosulcate pollen and trilete spores, evolved in southern and eastern Australia, Transantarctic Mountains region, South Africa and Argentina. The Onslow province is a mixture of Gondwanan and European taxa recognized in of north-western Australia, Madagascar, East Africa, Indian, and East Antarctic (Cesari and Colombi; 2013).

The recognition of Carnian European species in the Valle de la Luna Member of the Ishchigualasto Formation expands the distribution of the Onslow-type palynofloras. This assemblage was recovered from the site known as “El Hongo” in the Provincial Park, and contain the diagnostic “Onslow” species: Samaropollenites speciosus, Enzonalasporites vigens, Patinasporites densus, Vallatisporites ignacii, Ovalipollis pseudoalatus and Cycadopites stonei. This assemblage indicates that the Valle de la Luna Member was likely deposited under more humid conditions. It also implies the existence of a latitudinal floral belt from Timor (through the Circum-Mediterranean area) to western Argentina.

 

References:

Cesari, Silvia N., Colombi, Carina, Palynology of the late Triassic ischigualasto formation, Argentina: Paleoecological and paleogeographic implications, Palaeogeography, Palaeoclimatology, Palaeoecology (2016), doi: 10.1016/j.palaeo.2016.02.023

Césari, S. N., Colombi, C. E., 2013. A new Late Triassic phytogeographical scenario in westernmost Gondwana. Nature communications, 4.

Spalletti, L. A. Artabe, A. E. & Morel, E. M. Geological factors and evolution of southwestern Gondwana Triassic plants. Gondwana Res. 6, 119–134 (2003).

 

Ecosystem instability in the Late Triassic and the early evolution of dinosaurs.

The Late Triassic Petrified Forest Member of the Chinle Formation (Photo from AASG)

The Late Triassic Petrified Forest Member of the Chinle Formation (Photo from AASG)

Dinosaurs likely originated in the Middle Triassic and the first unequivocal dinosaur fossils are known from the late Carnian, but much about the geological and temporal backdrop of early dinosaur history remains poorly understood. A key question is why early dinosaurs were rare and species-poor at low paleolatitudes throughout the Late Triassic Period, for at least 30 million years after their origin.

The oldest well-dated identified dinosaurs are from the late Carnian (approx. 230 Ma) of the lower Ischigualasto Formation in northwestern Argentina. Similarly, the Santa Maria and Caturrita formations in southern Brazil preserve basal dinosauromorphs, basal saurischians, and early sauropodomorphs. In North America, the oldest dated occurrences of vertebrate assemblages with dinosaurs are from the Chinle Formation, but are less abundant and species rich compared to those from South America. The fact that those assemblages were at moderately high paleolatitudes during the Late Triassic, and the North American assemblages were near the paleoequator supports the hypotheses for a diachronous rise of dinosaurs across paleolatitudes (Irmis et al., 2011).

A reconstructed scene from the Late Triassic (Norian) of central Pangea. (Credit: image from Brusatte, S. L. 2008)

A reconstructed scene from the Late Triassic (Norian) of central Pangea. (Image from Brusatte, S. L. 2008, Dinosaurs, Quercus Publishing, London).

The Late Triassic is marked by a return to the “hothouse” condition of the Early Triassic, with two greenhouse crisis that may also have played a role in mass extinctions and long-term evolutionary trends (Retallack, 2013). The paleoclimate was a very arid with intense evaporation rate. Although there was at least one time of significant increase in rainfall known as the “Carnian Pluvial Event”, possibly related to the rifting of Pangea. Now, a multiproxy study  suggests  that fluctuating aridity in tropical and subtropical Pangea could explain why Triassic dinosaur faunas at low latitudes are restricted to small, slower growing carnivorous forms, whereas large-bodied herbivores, including sauropodomorph dinosaurs, are absent at low paleolatitudes during the Late Triassic “hothouse.” The palynomorphs recovered from sediments of the Chinle Formation indicate a major change from a seed fern-dominated (Alisporites) assemblage with accessory gymnosperms to one dominated by conifers and seed ferns in the lower portion of the Petrified Forest Member. In addition, the extensive charcoal record in the Petrified Forest Member provides evidence of paleo-environmental variability and aridity. 

 

References:

Jessica H. Whiteside, Sofie Lindström, Randall B. Irmis, Ian J. Glasspool, Morgan F. Schaller, Maria Dunlavey, Sterling J. Nesbitt, Nathan D. Smith, and Alan H. Turner. 2015. Extreme ecosystem instability suppressed tropical dinosaur dominance for 30 million years. PNAS: doi:10.1073/pnas.1505252112

Brusatte, S. L., Nesbitt, S. J., Irmis, R. B., Butler, R. J., Benton, M. J., and Norell, M. A. 2010. The origin and early radiation of dinosaurs. Earth-Science Reviews, 101, 68-100

Holz, M., Mesozoic paleogeography and paleoclimates – a discussion of the diverse greenhouse and hothouse conditions of an alien world, Journal of South American Earth Sciences (2015), doi: 10.1016/j.jsames.2015.01.001

Nesbitt,  S. J., Irmis,  R. B, Parker,  W. G. (2007) A critical re-evaluation of the Late Triassic di-nosaur taxa of North America. J Syst Palaeontology 5(2):209243

Sellwood, B.W. & Valdes, P.J. 2006. Mesozoic climates: General circulation models and the rock Record. Sedimentary Geology 190:269–287.

 

Brief paleontological history of planktonic foraminifera.

neoglobo

Neogloboquadrina dutertrei. (Credit: Dr Kate Darling).

Planktonic foraminifera made their first appearance in the Late Triassic. Although, identifying the first occurrence of planktonic foraminifera is complex, with many suggested planktonic forms later being reinterpreted as benthic. They are present in different types of marine sediments, such as carbonates or limestones, and are excellent biostratigraphic markers.

Their test are made of  globular chambers composed of secrete calcite or aragonite, with no internal structures and  different patterns of chamber disposition: trochospiral, involute trochospiral and planispiral growth. During the Cenozoic, some forms exhibited supplementary apertures or areal apertures. The tests also show perforations and a variety of surface ornamentations like cones, short ridges or spines.

The phylogenetic evolution of planktonic foraminifera are closely associated with global and regional changes in climate and oceanography.

planktonic foraminifera evolution

The evolution of early planktonic foraminifera (From Boudagher-Fadel, 2013)

All species of Late Triassic and Jurassic planktonic foraminifera are members of the superfamily Favuselloidea. They present a test composed by aragonite, with microperforations, and sub-globular adult chambers. After the major End Triassic event, the Jurassic period saw warm tropical greenhouse conditions worldwide. The surviving planktonic foraminifera were usually dominated by small globular forms.

It was suggested  that a second transition from a benthic to a planktonic mode of life took place at the Jurassic, which occurred under conditions similar to those that triggered planktonic speciation in the Late Triassic (hot and dry global climate, and low sea levels).

During the Cretaceous,  the favusellids must have made the transition from being aragonitic to calcitic.  Also, in the Late Aptian there was a significant number of planktonic foraminiferal extinctions, but these were compensated by the establishment of a large number of new genera at the Aptian–Albian boundary.

Planktonic foraminifera from the Sargasso Sea in the North Atlantic Ocean. (Photograph courtesy Colomban de Vargas, EPPO/SBRoscoff.)

Planktonic foraminifera from the Sargasso Sea in the North Atlantic Ocean. (Photograph courtesy Colomban de Vargas, EPPO/SBRoscoff.)

The Paleogene assemblage of planktonic foraminifera was derived from the few species that survive the mass extinction event at the end of the Cretaceous.

In the Early Miocene, the planktonic foraminifera were most abundant and diverse in the tropics and subtropics, and after the Mid-Miocene Climatic Optimum, many species were adapted to populate temperate and sub-polar oceans.

During the Middle and Late Pliocene, the final closure of the Central American seaway, changed oceanic circulation and drove a significant number of species extinctions. Most modern, living species originated in the Pliocene and Pleistocene.

References:

Armstrong, H. A., Brasier, M. D., 2005. Microfossils (2nd Ed). Blackwell, Oxford.

Boudagher-Fadel, MK; (2013) Biostratigraphic and Geological Significance of Planktonic Foraminifera. (2nd ed.)