Carlo Romano

Abstract: Classic descriptions and a previous reconstruction of the Middle Triassic ray-finned fish Birgeria stensioei Aldinger, 1931 from the Besano Formation (= Grenzbitumenzone) of Monte San Giorgio (Canton Ticino, Switzerland) and Besano (Lombardy, Italy) relied either on poor material or on a single specimen only. This study of B. stensioei is based on 67 specimens from that region in the Southern Alps including the lectotype of B. stensioei. The reappraisal of B. stensioei indicates that this species differs from the other known species ...

The Permian and Triassic were key time intervals in the history of life on Earth. Both periods are marked by a series of biotic crises including the most catastrophic of such events, the end-Permian mass extinction, which eventually led to a major turnover from typical Palaeozoic faunas and floras to those that are emblematic for the Mesozoic and Cenozoic. Here we review patterns in Permian-Triassic bony fishes, a group whose evolutionary dynamics are understudied. Based on data from primary literature, we analyse changes in their taxonomic diversity and body size (as a proxy for trophic position) and explore their response to Permian-Triassic events. Diversity and body size are investigated separately for different groups of Osteichthyes (Dipnoi, Actinistia, 'Palaeopterygii', 'Subholostei', Holostei, Teleosteomorpha), within the marine and freshwater realms and on a global scale (total diversity) as well as across palaeolatitudinal belts. Diversity is also measured for different palaeogeographical provinces. Our results suggest a general trend from low osteichthyan diversity in the Permian to higher levels in the Triassic. Diversity dynamics in the Permian are marked by a decline in freshwater taxa during the Cisuralian. An extinction event during the end-Guadalupian crisis is not evident from our data, but 'palaeopterygians' experienced a significant body size increase across the Guadalupian-Lopingian boundary and these fishes upheld their position as large, top predators from the Late Permian to the Late Triassic. Elevated turnover rates are documented at the Permian-Triassic boundary, and two distinct diversification events are noted in the wake of this biotic crisis, a first one during the Early Triassic (dipnoans, actinistians,…

In the wake of the end-Permian mass extinction, the Early Triassic (~251.9 to 247 million years ago) is portrayed as an environmentally unstable interval characterized by several biotic crises and heavily depauperate marine benthic ecosystems. We describe a new fossil assemblage—the Paris Biota—from the earliest Spathian (middle Olenekian, ~250.6 million years ago) of the Bear Lake area, southeastern Idaho, USA. This highly diversified assemblage documents a remarkably complex marine ecosystem including at least seven phyla and 20 distinct metazoan orders, along with algae. Most unexpectedly, it combines early Paleozoic and middle Mesozoic taxa previously unknown from the Triassic strata, among which are primitive Cambrian-Ordovician leptomitid sponges (a 200–million year Lazarus taxon) and gladius-bearing coleoid cephalopods, a poorly documented group before the Jurassic (~50 million years after the Early Triassic). Additionally, the crinoid and ophiuroid specimens show derived anatomical characters that were thought to have evolved much later. Unlike previous works that suggested a sluggish postcrisis recovery and a low diversity for the Early Triassic benthic organisms, the unexpected composition of this exceptional assemblage points toward an early and rapid post-Permian diversification for these clades. Overall, it illustrates a phylogenetically diverse, functionally complex, and trophically multileveled marine ecosystem, from primary producers up to top predators and potential scavengers. Hence, the Paris Biota highlights the key evolutionary position of Early Triassic fossil ecosystems in the transition from the Paleozoic to the Modern marine evolutionary fauna at the dawn of the Mesozoic era.

A new, marine osteichthyan (bony fish) fauna from the Early Triassic of northern India is presented. The material was collected in situ at localities within Pin Valley (Lahaul and Spiti District, Himachal Pradesh, India) and is dated as middle-late Dienerian (one specimen possibly earliest Smithian). The new ichthyofauna includes a lower jaw of the predatory basal ray-finned fish Saurichthys, a nearly complete specimen of a parasemionotid neopterygian (cf. Watsonulus cf. eugnathoides), as well as further articulated and disarticulated remains (Actinopterygii indet., Actinistia indet.), and thus comprises the most complete Triassic fish fossils known from the Indian subcontinent. Saurichthys is known from many Triassic localities and reached a global distribution rapidly after the Late Permian mass extinction event. Parasemionotidae, a species-rich family restricted to the Early Triassic, also achieved widespread distribution during this epoch. Comparison of the Spiti material with other parasemionotid species reveals similarities with Watsonulus eugnathoides from Madagascar. However, taxonomic ambiguities within Parasemionotidae prevent a specific attribution of the Spiti specimen. The new material also includes an isolated actinistian urohyal exhibiting morphology distinct from any previously described urohyal. Marine Dienerian black shale deposited on continental shelves are common not only in the Himalayas but also in other geographic regions. Anoxic depositional settings provide ideal preservational conditions for vertebrate fossils, suggesting that additional ichthyofaunas could still be discovered in marine Dienerian strata of other localities. The study of Early Triassic fish assemblages, including the presented one, is fundamental for our understanding of the great osteichthyan diversification after the Late Permian mass extinction event.

Saurichthys, characterized by a long slender body and an elongated rostrum, is one of the most iconic genera of Late Paleozoic–Early Mesozoic fishes. The genus was particularly speciose in the Triassic, with a global distribution in both marine and freshwater habitats. Here, we describe two new species from the Middle Triassic Besano Formation of Monte San Giorgio, Switzerland, Saurichthys breviabdominalis sp. nov. and Saurichthys rieppeli sp. nov. S. breviabdominalis is characterized by a proportionately long operculum, short abdominal region and rib-like mid-lateral scales, whereas S. rieppeli is divergent from other Middle Triassic saurichthyids in the block-like haemal arches, fringing fulcra on the pelvic and unpaired fins, and reduction of the squamation to a single row in the abdominal region. Phylogenetic analysis places S. rieppeli in a basal position relative to congeners from the Alpine Triassic, and supports previous hypotheses regarding the convergent evolution of reduced squamation within saurichthyids. S. breviabdominalis forms a monophyletic group with species from the same locality, suggesting divergence in sympatry. This finding has implications for our understanding of disparity and character evolution in saurichthyid fishes, as well as ecomorphological divergence and resource partitioning between closely related fishes in Triassic marine ecosystems.

The Permian and Triassic were key time intervals in the history of life on Earth. Both periods are marked by a series of biotic crises including the most catastrophic of such events, the end-Permian mass extinction, which eventually led to a major turnover from typical Palaeozoic faunas and floras to those that are emblematic for the Mesozoic and Cenozoic. Here we review patterns in Permian–Triassic bony fishes, a group whose evolutionary dynamics are understudied. Based on data from primary literature, we analyse changes in their taxonomic diversity and body size (as a proxy for trophic position) and explore their response to Permian–Triassic events. Diversity and body size are investigated separately for different groups of Osteichthyes (Dipnoi, Actinistia, ‘Palaeopterygii’, ‘Subholostei’, Holostei, Teleosteomorpha), within the marine and freshwater realms and on a global scale (total diversity) as well as across palaeolatitudinal belts. Diversity is also measured for different palaeogeographical provinces. Our results suggest a general trend from low osteichthyan diversity in the Permian to higher levels in the Triassic. Diversity dynamics in the Permian are marked by a decline in freshwater taxa during the Cisuralian. An extinction event during the end-Guadalupian crisis is not evident from our data, but ‘palaeopterygians’ experienced a significant body size increase across the Guadalupian–Lopingian boundary and these fishes upheld their position as large, top predators from the Late Permian to the Late Triassic. Elevated turnover rates are documented at the Permian–Triassic boundary, and two distinct diversification events are noted in the wake of this biotic crisis, a first one during the Early Triassic (dipnoans, actinistians, ‘palaeopterygians’, ‘subholosteans’) and a second one during the Middle Triassic (‘subholosteans’, neopterygians). The origination of new, small taxa predominantly among these groups during the Middle Triassic event caused a significant reduction in osteichthyan body size. Neopterygii, the clade that encompasses the vast majority of extant fishes, underwent another diversification phase in the Late Triassic. The Triassic radiation of Osteichthyes, predominantly of Actinopterygii, which only occurred after severe extinctions among Chondrichthyes during the Middle–Late Permian, resulted in a profound change within global fish communities, from chondrichthyan-rich faunas of the Permo-Carboniferous to typical Mesozoic and Cenozoic associations dominated by actinopterygians. This turnover was not sudden but followed a stepwise pattern, with leaps during extinction events.

Examining the geological past of our planet allows us to study periods of severe climatic and biological crises and recoveries, biotic and abiotic ecosystem fluctuations, and faunal and floral turnovers through time. Furthermore, the recovery dynamics of large predators provide a key for evaluation of the pattern and tempo of ecosystem recovery because predators are interpreted to react most sensitively to environmental turbulences. The end-Permian mass extinction was the most severe crisis experienced by life on Earth, and the common paradigm persists that the biotic recovery from the extinction event was unusually slow and occurred in a step-wise manner, lasting up to eight to nine million years well into the early Middle Triassic (Anisian) in the oceans, and even longer in the terrestrial realm. Here we survey the global distribution and size spectra of Early Triassic and Anisian marine predatory vertebrates (fishes, amphibians and reptiles) to elucidate the height of trophic pyramids in the aftermath of the end-Permian event. The survey of body size was done by compiling maximum standard lengths for the bony fishes and some cartilaginous fishes, and total size (estimates) for the tetrapods. The distribution and size spectra of the latter are difficult to assess because of preservation artifacts and are thus mostly discussed qualitatively. The data nevertheless demonstrate that no significant size increase of predators is observable from the Early Triassic to the Anisian, as would be expected from the prolonged and stepwise trophic recovery model. The data further indicate that marine ecosystems characterized by multiple trophic levels existed from the earliest Early Triassic onwards. However, a major change in the taxonomic composition of predatory guilds occurred less than two million years after the end-Permian extinction event, in which a transition from fish/amphibian to fish/reptile-dominated higher trophic levels within ecosystems became apparent.

Sun et al. (Reports, 19 October 2012, p. 366) reconstructed Permian to Middle Triassic equatorial seawater temperatures. After correct temporal positioning of their data points, their presumed trends of temperature changes, and hence their assumption of a one-to-one relationship between putative "lethally hot" seawater temperatures and a disputable equatorial "eclipse" of some organisms, are no longer supported by their data.

Recovery from the end-Permian mass extinction is frequently described as delayed, with complex ecological communities typically not found in the fossil record until the Middle Triassic epoch. However, the taxonomic diversity of a number of marine groups, ranging from ammonoids to benthic foraminifera, peaked rapidly in the Early Triassic. These variations in biodiversity occur amidst pronounced excursions in the carbon isotope record, which are compatible with episodes of massive CO2 outgassing from the Siberian Large Igneous Province. Here we present a high-resolution Early Triassic temperature record based on the oxygen isotope composition of pristine apatite from fossil conodonts. Our reconstruction shows that the beginning of the Smithian substage of the Early Triassicwas marked by a cooler climate, followed by an interval of warmth lasting until the Spathian substage boundary. Cooler conditions resumed in the Spathian. We find the greatest increases in taxonomic diversity during the cooler phases of the early Smithian and early Spathian. In contrast, a period of extreme warmth in the middle and late Smithian was associated with floral ecological change and high faunal taxonomic turnover in the ocean. We suggest that climate upheaval and carboncycle perturbations due to volcanic outgassing were important drivers of Early Triassic biotic recovery.

A new marine fish assemblage from the late Smithian (Olenekian, Early Triassic) Anasibirites beds of the Thaynes Formation collected near Georgetown (Bear Lake County, south-east Idaho, USA) comprises actinopterygians (Saurichthys cf. elongatus, Actinopterygii indet.), sarcopterygians (Actinistia indet.), and possibly chondrichthyans. We review the global fossil record of the Triassic lower actinopterygian Saurichthys, which is used herein for a case study of trends in morphological adaptations as well as variations in palaeogeographic distribution and diversity dynamics of Early Mesozoic fishes. In the Early Triassic, Saurichthys already occupied a top position in marine food webs, with some species achieving body lengths of up to 1.5 m. Distribution of morphological characters in Saurichthys during the Triassic suggests trends towards a reduction in squamation, stiffening of the fins and axial skeleton, shortening of the postorbital portion of the cranium, and reduction in the number of dermal skull bones. The postcranial adaptations in particular helped to improve the fast-start ability of these ambush predatory fishes. The palaeogeographic range of Saurichthys changed from a virtually global distribution in the Early Triassic (indicating rapid dispersal within marine and freshwater ecosystems after the end-Permian mass extinction) to an occurrence mainly restricted to the north-western Tethys in the Late Triassic, and also towards increasing rarity within continental deposits. Modifications in the palaeogeographic distribution were accompanied by successive loss in global species richness and were possibly related to intra-Triassic extinction events, environmental alterations and/or competition.

"Present paper gives an updated summary of research history on the Chondrichthyes and Osteichthyes of the Early Triassic  (Griesbachian, Dienerian, Smithian, Spathian) and primarily of the early Anisian. Early Triassic and Anisian marine and freshwater ichthyofaunas are found on all continents except South America, and much more fish assemblages are known from the Northern than from the Southern Hemisphere. The Early Triassic and the Anisian are times of major importance for the phylogeny of the Chondrichthyes and Osteichthyes. After the end-Permian mass extinction the surviving groups of the cartilaginous and bony fishes recovered, and many new forms appeared in the Early Triassic. The neoselachians as well as close relatives of the teleosteans evolved, clades to which nearly all extant fishes belong. Present publication also provides a revised data base for the distribution of Early Triassic and early Anisian chondrichthyan and osteichthyan fishes in time and space on which future research on their paleobiodiversity shall be guided."