A New Dicynodont (Therapsida: Anomodontia) from the Permian of Southern Brazil and Its Implications for Bidentalian Origins
Authors:
Boos et al
Abstract:
Dicynodonts were a highly successful group of herbivorous therapsids that inhabited terrestrial ecosystems from the Middle Permian through the end of the Triassic periods. Permian dicynodonts are extremely abundant in African deposits, but are comparatively poorly known from the other regions of Gondwana. Here we describe a new South American dicynodont, Rastodon procurvidens gen. et sp. nov., from the Boqueirão farm site of the Rio do Rasto Formation, Paraná Basin, Guadalupian/Lopingian of Brazil. Diagnostic features of R. procurvidens include uniquely anteriorly-curved maxillary tusks, well-developed ridges extending from the crista oesophagea anteriorly along the pterygoid rami, strong posterior angulation of the posterior pterygoid rami, and a bulbous, well-developed retroarticular process of the articular. Phylogenetic analysis indicates that R. procurvidens is the earliest and most basal member of Bidentalia, a cosmopolitan clade that includes Permian and Triassic dicynodonts whose dentition is usually reduced to a pair of maxillary tusks.
Submarine palaeoenvironments during Emeishan flood basalt volcanism, SW China: Implications for plume–lithosphere interaction during the Capitanian, Middle Permian (‘end Guadalupian’) extinction event
Authors:
Jerram et al
Abstract:
Plume-induced lithospheric uplift and erosion are widely regarded as key features of large igneous province (LIP) emplacement, as is the coincidence of LIP eruption with major extinction and oceanic anoxic events (OAE). The Emeishan LIP, which erupted during the Capitanian (previously termed ‘end Guadalupian’) extinction event, has provided the most widely discussed example of axisymmetric doming above a rising mantle ‘plume’; advocates have argued that in excess of 500 m of uplift occurred over greater than 30 000 km2 causing extensive radially distributed erosion and alluvial fan formation. However, the recognition of submarine hydromagmatic and phreatomagmatic-style volcanism, as well as syn-volcanic marine sediments interbedded in the eruptive succession, now requires further examination to this simple plume–uplift model.
Here we present data from newly discovered sections from the center of the putative uplifted area (around Lake Er Hai, SW Yunnan Province,) that provide a more complete history of the Emeishan volcanism. These reveal that platform carbonate deposition was terminated by rapid subsidence, followed quickly by the onset of volcanism. Importantly, these eruptions also coincide with widespread losses amongst fusulinacean foraminifera and calcareous algae. For at least the lower two thirds of the 4–5 km thick lava pile, eruptions continued at or below sea level, as testified by the presence of voluminous mafic volcaniclastic deposits, pillow lavas, and development of syn-volcanic reefal limestones in the Emeishan inner zone. Only in the later stages of eruption did terrestrial lava flows become widely developed. This onset of volcanism in a submarine setting and the consequent violent, phreatomagmatic-style eruptions would have had a profound effect on marine fauna and exacerbated any volcanically induced climate effects during the Capitanian. The late Permian of SW China at the time of the Emeishan was an extended area of thinned lithosphere with epeiric seas, which appear to have been sustained through the onset of LIP emplacement. Therefore, while there remains substantial geochemical support of a plume origin for Emeishan volcanism, LIP emplacement cannot be ubiquitously associated with regional pre-eruption uplift, particularly where complex lithospheric structure exists above a plume.
Widespread shoaling of sulfidic waters linked to the end-Guadalupian (Permian) mass extinction
Authors:
Zhang et al
Abstract:
Multiple sulfur isotopes (32S, 33S, 34S, and 36S) measured on pyrites from the Penglaitan section, the Global Stratotype Section and Point (GSSP) for the Guadalupian-Lopingian Series boundary, and from the auxiliary Tieqiao section in South China show a sulfur isotope signal of negative and positive δ34S with negative Δ33S. We suggest that these data indicate mixing of 34S-enriched and 34S-depleted sulfur in the sediments, which may have been driven by shoaling of sulfidic waters. Similar isotopic data of negative δ34S with negative Δ33S were also observed from the EF section in the Delaware Basin of west Texas (USA). The consistency of the minor sulfur isotopic anomalies from both South China and west Texas suggests a causal link between widespread shoaling of sulfidic waters and the end-Guadalupian mass extinction.
A tapinocephalid dinocephalian (Synapsida, Therapsida) from the Rio do Rasto Formation (Paraná Basin, Brazil): Taxonomic, ontogenetic and biostratigraphic considerations
Authors:
Boos et al
Abstract:
Permian tetrapod fossils have been recovered from the Rio do Rasto Formation of Brazil since the 1970s. Previous studies of this fauna indicated strong affinities with the Guadalupian-Lopingian vertebrates of South Africa and Eastern Europe, suggesting biostratigraphic correlations between these areas. Here, a new dinocephalian specimen from the Rio do Rasto Formation in the Serra do Cadeado area (Paraná State, Brazil) is described based on fragmentary skull remains and an associated left lower jaw ramus. Despite the fragmentary nature of these remains, they represent the most complete tapinocephalid specimen known from South America. Comparison with other tapinocephalids indicates that the material described herein represents a juvenile or sub-adult specimen. Although it is not possible to identify this material to the genus level, it most closely resembles the ‘moschopines’ Moschops and Moschognathus from the Tapinocephalus Assemblage Zone of South Africa. As dinocephalians are known to be restricted to the Guadalupian, they are one of the best tetrapod biostratigraphic markers for the Rio do Rasto Formation, indicating that at least some of the strata in the areas where they occur [Serra do Cadeado (Paraná State), Fagundes farm and Boqueirão farm (Rio Grande do Sul State)] are Guadalupian. Vertebrate fossils from Rio do Rasto Formation occur in disperse, isolated and discontinuous outcrops, so that they have been grouped in ‘local faunas’. However, most of the specimens lack precise stratigraphic provenance data and even occurring in locations near each other they are not necessarily contemporary. Thus, until a more robust stratigraphic framework is developed, we suggest discontinuing use of ‘local faunas’ to this stratigraphic unit.
Extending the database of Permian palaeo-wildfire on Gondwana: Charcoal remains from the Rio do Rasto Formation (Paraná Basin), Middle Permian, Rio Grande do Sul State, Brazil
Authors:
Manfroi et al
Abstract:
Macroscopic charcoal has been recovered from a bonebed from the Barro Alto outcrop, Middle Permian (Guadalupian), Rio do Rasto Formation, in the southernmost part of the Paraná Basin, Brazil. During this time, a global biotic crisis was taking place and changes in the palaeofloristic composition can be observed. Records of palaeowildfires from the Guadalupian Period are scarce on a global scale. In the Paraná Basin low plant diversity, compared to the entirety of Gondwana, persists and just a few plant bearing localities have so far been described from the northern part of the basin. In the studied locality, macroscopic charcoal could be detected in an outcropping bonebed and the occurrence of palaeo-wildfires was confirmed. Besides representing the first palaeobotanical data for the Guadalupian of the southern part of the Paraná Basin, the charcoal remains analyzed here are also a significant record of the local environmental conditions during the increase in aridity observed during the Guadalupian and Lopingian in Gondwana. This contributes to the understanding of regional changes that took place during this interval.
Tiarajudens eccentricus and Anomocephalus africanus, two bizarre anomodonts (Synapsida, Therapsida) with dental occlusion from the Permian of Gondwana
Authors:
Cisneros et al
Abstract:
Anomodontia was a highly successful tetrapod clade during the Permian and the Triassic. New morphological information regarding two bizarre basal anomodonts is provided and their palaeoecological significance is explored. The osteology of the recently discovered Tiarajudens eccentricus Cisneros et al. 2011, from the Brazilian Permian, is described in detail. The taxon exhibits unusual postcranial features, including the presence of gastralia. Additional preparation and computed tomography scans of the holotype of Anomocephalus africanus Modesto et al. 1999 discovered in the Karoo Basin of South Africa allow a reappraisal of this genus. Anomocephalus is similar to Tiarajudens with regard to several traits, including a battery of large, transversally expanded, palatal teeth. Molariform teeth are present in the mandible of the African taxon, providing additional insight into the function of the earliest tooth-occlusion mechanism known in therapsids. At least two waves of tooth replacement can be recognized in the palate of Anomocephalus. The outsized, blade-like caniniforms of the herbivorous Tiarajudens allow several non-exclusive ecological interpretations, among which we favour intraspecific display or combat. This behaviour was an alternative to the head-butting practised by the contemporary dinocephalians. Combat specializations that are considered typical of Cenozoic herbivores likely evolved during the Middle Permian, at the time the first communities with diverse, abundant tetrapod herbivores were being assembled.
When and how did the terrestrial mid-Permian mass extinction occur? Evidence from the tetrapod record of the Karoo Basin, South Africa
Authors:
Day et al
Abstract:
A mid-Permian (Guadalupian epoch) extinction event at approximately 260 Ma has been mooted for two decades. This is based primarily on invertebrate biostratigraphy of Guadalupian–Lopingian marine carbonate platforms in southern China, which are temporally constrained by correlation to the associated Emeishan Large Igneous Province (LIP). Despite attempts to identify a similar biodiversity crisis in the terrestrial realm, the low resolution of mid-Permian tetrapod biostratigraphy and a lack of robust geochronological constraints have until now hampered both the correlation and quantification of terrestrial extinctions. Here we present an extensive compilation of tetrapod-stratigraphic data analysed by the constrained optimization (CONOP) algorithm that reveals a significant extinction event among tetrapods within the lower Beaufort Group of the Karoo Basin, South Africa, in the latest Capitanian. Our fossil dataset reveals a 74–80% loss of generic richness between the upper Tapinocephalus Assemblage Zone (AZ) and the mid-Pristerognathus AZ that is temporally constrained by a U–Pb zircon date (CA-TIMS method) of 260.259 ± 0.081 Ma from a tuff near the top of the Tapinocephalus AZ. This strengthens the biochronology of the Permian Beaufort Group and supports the existence of a mid-Permian mass extinction event on land near the end of the Guadalupian. Our results permit a temporal association between the extinction of dinocephalian therapsids and the LIP volcanism at Emeishan, as well as the marine end-Guadalupian extinctions.
A high-latitude Gondwanan lagerstätte: The Permian permineralised peat biota of the Prince Charles Mountains, Antarctica
Authors:
Slater et al
Abstract:
The Toploje Member chert is a Roadian to Wordian autochthonous–parautochthonous silicified peat preserved within the Lambert Graben, East Antarctica. It preserves a remarkable sample of terrestrial life from high-latitude central Gondwana prior to the Capitanian mass extinction event from both mega- and microfossil evidence that includes cryptic components rarely seen in other fossil assemblages. The peat layer is dominated by glossopterid and cordaitalean gymnosperms and contains moderately common herbaceous lycophytes, together with a broad array of dispersed organs of ferns and other gymnosperms. Rare arthropod-plant and fungal-plant interactions are preserved in detail together with a plethora of fungal morphotypes, Peronosporomycetes, arthropod remains and a diverse coprolite assemblage. Comparisons to other Palaeozoic ecosystems show that the macroflora is of low diversity. The fungal and invertebrate-plant associations demonstrate that a multitude of ecological interactions were well developed by the Middle Permian in high-latitude forest mires that contributed to the dominant coal deposits of the Southern Hemisphere. Quantitative analysis of the constituents of the silicified peat and of macerals within adjacent coal seams reveals that whilst silicified peats provide an unparalleled sample of the organisms forming Permian coals, they do not necessarily reflect the volumetric proportions of constituents within the derived coal. The Toploje Member chert Lagerstätte provides a snapshot of a rapidly entombed mire climax ecosystem in the closing stages of the Palaeozoic, but prior to the onset of the protracted crisis that engulfed and overthrew these ecosystems at the close of the Permian.
Carbon and sulfur isotopic fluctuations associated with the end-Guadalupian mass extinction in South China
Authors:
1. Yan Detian (a, b)
2. Zhang Liqin (c)
3. Qiu Zhen (b)
Affiliations:
a. Key Laboratory of Tectonics and Petroleum Resources of Ministry of Education, China University of Geosciences, Wuhan 430074, China
b. Institute of Geology and Geophysics, Chinese Academy of Sciences, Beijing 100029, China
c. Key Laboratory of Geospace Environment and Geodesy of Ministry of Education, Wuhan University, Wuhan 430079, China
Abstract:
Concentrations of total organic matter (TOC), carbon isotopic compositions of carbonate and organic matter (δ13Ccarb, δ13Corg), and sulfur isotopic compositions of carbonate associated sulfate (δ34Ssulfate) across the Guadalupian–Lopingian (G–L) boundary were analyzed from identical samples of Tieqiao section, Laibin, Guangxi province, South China. The δ13Ccarb values show a positive excursion from −0.45‰ to the peak of 3.80‰ in the Laibin limestone member of the Maokou Formation, followed by a drastic drop to −2.60‰ in the lowest Heshan formation, then returned to about 1.58‰. Similar to the trends of the δ13Ccarb values, Δ13Ccarb–org values also show a positive excursion followed by a sharp negative shift. The onset of a major negative carbon isotope excursion postdates the end Guadalupian extinction that indicates subsequent severe disturbance of the ocean–atmosphere carbon cycle. The first biostratigraphic δ34Ssulfate values during the G–L transition exhibit a remarkable fluctuation: a dramatic negative shift followed by a rapid positive shift, ranging from 36.88‰ to −37.41‰. These sulfate isotopic records suggest that the ocean during the G–L transition was strongly stratified, forming an unstable chemocline separating oxic shallow water from anoxic/euxinic deep water. Chemocline excursions, together with subsequent rapid transgression and oceanic anoxia, were likely responsible for the massive diversity decline of the G–L biotic crisis.
Through careful study of an ancient ancestor of modern turtles, researchers now have a clearer picture of how the turtles' most unusual shell came to be. The findings, reported on May 30 in Current Biology, a Cell Press publication, help to fill a 30- to 55-million-year gap in the turtle fossil record through study of an extinct South African reptile known as Eunotosaurus.
"The turtle shell is a complex structure whose initial transformations started over 260 million years ago in the Permian period," says Tyler Lyson of Yale University and the Smithsonian. "Like other complex structures, the shell evolved over millions of years and was gradually modified into its present-day shape."
The turtle shell isn't really just one thing—it is made up of approximately 50 bones. Turtles are the only animals that form a shell through the fusion of ribs and vertebrae. In all other animals, shells are formed from bony scales on the surface; they don't stick their bones on the outsides of their bodies.
"The reason, I think, that more animals don't form a shell via the broadening and eventually suturing together of the ribs is that the ribs of mammals and lizards are used to help ventilate the lungs," Lyson says. "If you incorporate your ribs into a protective shell, then you have to find a new way to breathe!" Turtles have done just that, with the help of a muscular sling.
Until recently, the oldest known fossil turtles, dating back about 215 million years, had fully developed shells, making it hard to see the sequence of evolutionary events that produced them. That changed in 2008 with the discovery of Chinese Odontochelys semitestacea, a reptile about 220 million years old, which had a fully developed plastron—the belly side of the shell—but only a partial carapace on its back.
Eunotosaurus takes the turtle and its shell back another 40 million years or so. It had nine broadened ribs found only in turtles. And like turtles, it lacked the intercostal muscles running between its ribs. But Eunotosaurus didn't have other features common to Odontochelys and turtles, including broad spines on their vertebrae.
Lyson says he and his colleagues now plan to investigate various other aspects of turtles' respiratory systems, which allow them to manage with their ribs locked up into a protective outer shell. "It is clear that this novel lung ventilation mechanism evolved in tandem with the origin of the turtle shell," he says.
