Jörg Fröbisch

The Triassic dicynodont Kombuisia (Synapsida, Anomodontia) from Antarctica, a refuge from the terrestrial Permian-Triassic mass extinction

Fossils from the central Transantarctic Mountains in Antarctica are referred to a new species of the Triassic genus Kombuisia, one of four dicynodont lineages known to survive the end-Permian mass extinction. The specimens show a unique combination of characters only present in this genus, but the new species can be distinguished from the type species of the genus, Kombuisia frerensis, by the presence of a reduced but slit-like pineal foramen and the lack of contact between the postorbitals. Although incomplete, the Antarctic specimens are significant because Kombuisia was previously known only from the South African Karoo Basin and the new specimens extend the taxon’s biogeographic range to a wider portion of southern Pangaea. In addition, the new finds extend the known stratigraphic range of Kombuisia from the Middle Triassic subzone B of the Cynognathus Assemblage Zone into rocks that are equivalent in age to the Lower Triassic Lystrosaurus Assemblage Zone, shortening the proposed ghost lineage of this taxon. Most importantly, the occurrence of Kombuisia and Lystrosaurus mccaigi in the Lower Triassic of Antarctica suggests that this area served as a refuge from some of the effects of the end-Permian extinction. The composition of the lower Fremouw Formation fauna implies a community structure similar to that of the ecologically anomalous Lystrosaurus Assemblage Zone of South Africa, providing additional evidence for widespread ecological disturbance in the extinction’s aftermath.

The oldest caseid synapsid from the Late Pennsylvanian of Kansas, and the evolution of herbivory in terrestrial vertebrates.

The origin and early evolution of amniotes (fully terrestrial vertebrates) led to major changes in the structure and hierarchy of terrestrial ecosystems. The first appearance of herbivores played a pivotal role in this transformation. After an early bifurcation into Reptilia and Synapsida (including mammals) 315 Ma, synapsids dominated Paleozoic terrestrial vertebrate communities, with the herbivorous caseids representing the largest vertebrates on land. Eocasea martini gen. et sp. nov., a small carnivorous caseid from the Late Carboniferous, extends significantly the fossil record of Caseidae, and permits the first clade-based study of the origin and initial evolution of herbivory in terrestrial tetrapods. Our results demonstrate for the first time that large caseid herbivores evolved from small, non-herbivorous caseids. This pattern is mirrored by three other clades, documenting multiple, independent, but temporally staggered origins of herbivory and increase in body size among early terrestrial tetrapods, leading to patterns consistent with modern terrestrial ecosystem.

Macropredatory ichthyosaur from the Middle Triassic and the origin of modern trophic networks

The biotic recovery from Earth’s most severe extinction event at the Permian-Triassic boundary largely reestablished the preextinction structure of marine trophic networks, with marine reptiles assuming the predator roles. However, the highest trophic level of todays marine ecosystems, i.e., macropredatory tetrapods that forage on prey of similar size to their own, was thus far lacking in the Paleozoic and early Mesozoic. Here we report a top-tier tetrapod predator, a very large (>8.6 m) ichthyosaur from the early Middle Triassic (244 Ma), of Nevada. This ichthyosaur had a massive skull and large labiolingually flattened teeth with two cutting edges indicative of a macropredatory feeding style. Its presence documents the rapid evolution of modern marine ecosystems in the Triassic where the same level of complexity as observed in today’s marine ecosystems is reached within 8 My after the Permian-Triassic mass extinction and within 4 My of the time reptiles first invaded the sea. This find also indicates that the biotic recovery in the marine realm may have occurred faster compared with terrestrial ecosystems, where the first apex predators may not have evolved before the Carnian.

A new large caseid (Synapsida, Caseasauria) from the Permian of Rodez (France), including a reappraisal of “Casea” rutena Sigogneau-Russell & Russell, 1974

ABSTRACT The description of a new large caseid, Ruthenosaurus russelbrum n. gen., n. sp. from the Early Permian (upper Cisuralian to lower Lopingian) of the Rodez Basin, France, is based on a partial postcranial skeleton that was initially collected at the same time as the holotype of “Cásea” rutena Sigogneau-Russell & Russell, 1974. Despite its distinctly larger size than “C.” rutena, the holotype of Ruthenosaurus n. gen. clearly represents an immature individual, as shown most clearly by the lack of fusion of the neural arches to their respective vertebral centra and incomplete ossification of the ends of the limb elements, including the absence of an ossified olecranon on the ulna. Nonetheless, Ruthenosaurus n. gen. is diagnosed by several autapomorphic characters, including dorsal vertebrae with anteriorly tilting neural spines and a diamond-shaped outline in transverse section, a first sacral rib with a dorsoventrally expanded distal head, and a low iliac blade with a poorly developed anterior process. The new taxon is further distinguishable from the only other known French caseid, “Casea” rutena, by the shape of the distal part of the humerus, including an ectepicondylar notch rather than a fully enclosed foramen, the specific shape of the ulna, and the overall robustness of the specimen. The taxonomic status of “Cased” rutena is discussed and it is concluded that this species should be moved into a new genus named Euromycter n. gen. The occurrence of the large-sized Ruthenosaurus n. gen. in France increases our knowledge about the early diversity of this clade in Europe.

New Permian fauna from tropical Gondwana

Terrestrial vertebrates are first known to colonize high-latitude regions during the middle Permian (Guadalupian) about 270 million years ago, following the Pennsylvanian Gondwanan continental glaciation. However, despite over 150 years of study in these areas, the biogeographic origins of these rich communities of land-dwelling vertebrates remain obscure. Here we report on a new early Permian continental tetrapod fauna from South America in tropical Western Gondwana that sheds new light on patterns of tetrapod distribution. Northeastern Brazil hosted an extensive lacustrine system inhabited by a unique community of temnospondyl amphibians and reptiles that considerably expand the known temporal and geographic ranges of key subgroups. Our findings demonstrate that tetrapod groups common in later Permian and Triassic temperate communities were already present in tropical Gondwana by the early Permian (Cisuralian). This new fauna constitutes a new biogeographic province with North American affinities and clearly demonstrates that tetrapod dispersal into Gondwana was already underway at the beginning of the Permian.

Early Evolutionary History of the Synapsida

An exceptionally well-preserved specimen of the bauriid therocephalian Microgomphodon oligocynus from the Burgersdorp Formation (Early-Middle Triassic, Cynognathus Assemblage Zone) of the South African Karoo is described. In addition, a taxonomic revision of bauriid therocephalians from southern Africa, based on firsthand examination of almost all know specimens, is presented. Microgomphodon oligocynus and Bauria cynops are recognized as the only valid species of southern African bauriids. Microgomphodon oligocynus is differentiated from B. cynops on the basis of clear-cut morphological features such as the presence of a complete postorbital bar, pineal foramen, contribution of the vomer to the osseous secondary palate, comparatively large orbits, presence of a lateral fossa on the posterior portion of the horizontal ramus and on the coronoid process of the dentary, and reduced number of postcanines. Procrustes analysis of the two best-preserved specimens of these species allowed recognition of further shape differences: M. oligocynus has a taller but narrower cranium, taller snout, temporal opening more expanded laterally, pterygoid process located more anteriorly, and smaller suborbital vacuity. The mandible of M. oligocynus has a higher symphysis, relatively short corpus, and more laterally-directed coronoid process. Microgomphodon oligocynus is known from the Olenekian to what are probably late Anisian levels in South Africa and Namibia, whereas B. cynops is restricted to the early Anisian of South Africa.

Elevated Extinction Rates as a Trigger for Diversification Rate Shifts: Early Amniotes as a Case Study.

Tree shape analyses are frequently used to infer the location of shifts in diversification rate within the Tree of Life. Many studies have supported a causal relationship between shifts and temporally coincident events such as the evolution of “key innovations”. However, the evidence for such relationships is circumstantial. We investigated patterns of diversification during the early evolution of Amniota from the Carboniferous to the Triassic, subjecting a new supertree to analyses of tree balance in order to infer the timing and location of diversification shifts. We investigated how uneven origination and extinction rates drive diversification shifts, and use two case studies (herbivory and an aquatic lifestyle) to examine whether shifts tend to be contemporaneous with evolutionary novelties. Shifts within amniotes tend to occur during periods of elevated extinction, with mass extinctions coinciding with numerous and larger shifts. Diversification shifts occurring in clades that possess evolutionary innovations do not coincide temporally with the appearance of those innovations, but are instead deferred to periods of high extinction rate. We suggest such innovations did not cause increases in the rate of cladogenesis, but allowed clades to survive extinction events. We highlight the importance of examining general patterns of diversification before interpreting specific shifts.

Olson's Extinction and the latitudinal biodiversity gradient of tetrapods in the Permian

The terrestrial vertebrate fauna underwent a substantial change in composition between the lower and middle Permian. The lower Permian fauna was characterized by diverse and abundant amphibians and pelycosaurian-grade synapsids. During the middle Permian, a therapsid-dominated fauna, containing a diverse array of parareptiles and a considerably reduced richness of amphibians, replaced this. However, it is debated whether the transition is a genuine event, accompanied by a mass extinction, or whether it is merely an artefact of the shift in sampling from the palaeoequatorial latitudes to the palaeotemperate latitudes. Here we use an up-to-date biostratigraphy and incorporate recent discoveries to thoroughly review the Permian tetrapod fossil record. We suggest that the faunal transition represents a genuine event; the lower Permian temperate faunas are more similar to lower Permian equatorial faunas than middle Permian temperate faunas. The transition was not consistent across latitudes; the turnover occurred more rapidly in Russia, but was delayed in North America. The argument that the mass extinction is an artefact of a latitudinal biodiversity gradient and a shift in sampling localities is rejected: sampling correction demonstrates an inverse latitudinal biodiversity gradient was prevalent during the Permian, with peak diversity in the temperate latitudes.

One hundred years of continental drift: the early Italian reaction to Wegener’s ‘visionary’ theory

Abstract One hundred years ago in 1915 ‘Die Entstehung der Kontinente und Ozeane’ by Alfred Wegener was published, destined to become one of the most controversial geological opus in the first half of the twentieth century. Wegener is the first to combine the most diverse geological (sensu lato) evidences in a single great synthesis. Nonetheless, apart from few upholders, the initial reaction to the drift hypothesis was fierce opposition, and the strongest criticism came from geophysics, the same discipline that, paradoxically, starting from the 1950s led to the Plate Tectonics revolution and, ultimately, to a complete re-evaluation of Wegener’s hypothesis. In the present paper we discuss the initial reaction of Italian scientists to the original continental drift theory, with particular focus on the period between the two world wars. Italian geologists like Fossa-Mancini and Gortani were almost favourable to the new theory, while authors such as Vardabasso and Sacco were neutral or even hostile to the new hypothesis, so iconoclastic for the widely accepted fixist vision of the time. In any case, all these scientists agreed that the new theory had great potential for reopening an enthusiastic debate on issues that were given as established paradigms – the genuine way for progress in science.

Redescription of the geikiid Pelanomodon (Therapsida, Dicynodontia), with a reconsideration of ‘Propelanomodon’

ABSTRACT Cryptodont dicynodonts are some of the most abundant therapsid taxa in the upper Permian fossil record. Despite extensive taxonomic study of these families, the species diversity of geikiid cryptodonts remains problematic, particularly for the set of edentulous taxa in the genera Pelanomodon, Propelanomodon, and Geikia. Here, all known specimens of tuskless geikiid dicynodonts from the upper Permian Karoo Basin of South Africa are reevaluated. The genus Pelanomodon is redescribed based on a series of skulls representing a range of sizes. All tuskless geikiids from the Karoo Basin are considered to represent a single species (Pelanomodon moschops) that exhibited cranial dimorphism as adults. The nominal species Pe. moschops and Pe. rubidgei differ only in degree of facial boss development and are most parsimoniously interpreted as sexual dimorphs. Propelanomodon is considered to represent the juvenile morphotype of Pelanomodon moschops, as indicated by the geographic and stratigraphic overlap of the two taxa, the lack of any small skulls of the Pelanomodon morphotype in well-sampled localities, and the presence of specimens of intermediate size and morphology. ‘Propelanomodon’ specimens are distinctive among juvenile dicynodonts in having a significantly narrower intertemporal region than adults. Analysis of intertemporal width relative to total skull size in Pelanomodon and the abundant cryptodont taxa Aulacephalodon, Oudenodon, and Tropidostoma suggests that Pelanomodon had a unique growth trajectory. Based on available stratigraphic data, Pelanomodon was restricted to the uppermost Permian Daptocephalus Assemblage Zone of the South African Beaufort Group and was a victim of the end-Permian mass extinction. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP