Permian–Triassic Osteichthyes (bony fishes): diversity dynamics and body size evolution

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.     

Volcanic perturbations of the marine environment in South China preceding the latest Permian mass extinction and their biotic effects

The Dongpan section in southern Guangxi Province records the influence of local volcanic activity on marine sedimentation at intermediate water depths (~200-500 m) in the Nanpanjiang Basin (South China) during the late Permian crisis. We analyzed ~100 samples over a 12-m-thick interval, generating palynological, paleobiological, and geochemical datasets to investigate the nature and causes of environmental changes. The section records at least two major volcanic episodes that culminated in deposition of approximately 25- to 35-cm-thick ash layers (bentonites) and that had profound effects on conditions in both the Dongpan marine environment and adjacent land areas. Intensification of eruptive activity during each volcanic cycle resulted in a shift toward conifer forests, increased wildfire intensity, and elevated subaerial weathering fluxes. The resulting increase in nutrient fluxes stimulated marine productivity in the short term but led to a negative feedback on productivity in the longer term as the OMZ of the Nanpanjiang Basin expanded, putting both phytoplankton and zooplankton communities under severe stress. Radiolarians exhibit large declines in diversity and abundance well before the global mass extinction horizon, demonstrating the diachroneity of the marine biotic crisis. The latest Permian crisis, which was probably triggered by the Siberian Traps flood basalts, intensified the destructive effects of the earlier local eruptions on terrestrial and marine ecosystems of the South China craton.     

Permian and early Triassic isotopic records of carbon and strontium in Australia and a scenario of events about the Permian‐Triassic boundary

The negative shift in δ¹³C values of carbonate carbon at the Permian/Triassic boundary is one of the better documented geochemical signatures of a mass extinction event. The similar negative shift in δ¹³C values in organic carbon from Permian/Triassic boundary marine sediments in Austria and Canada is shown to occur also in marine and non‐marine sediments from Australian sedimentary basins. This negative shift in δ¹³C values is used to calibrate Australian sections lacking diagnostic faunal elements identifying the Permian/Triassic boundary. The minimum in the carbonate ⁸⁷Sr/⁸⁶Sr seawater curve from carbonates across the Guadalupian/Ochoan Stage boundary, mainly from North America, is shown to occur also in brachiopod calcite mainly from the Bowen Basin of eastern Australia, hence providing a second calibration point in the Australian sedimentary record. These two geochemical events support a model of a runaway greenhouse developing about the Permian/Triassic boundary; this is inferred to have contributed to the end‐Permian mass extinction.     

Transgressive–regressive sequences on the slope of an isolated carbonate platform (Middle–Late Permian, Laibin, South China)

From the Middle to Late Permian, the Laibin area in Guangxi, South China, was situated on the slope of an isolated carbonate platform, on which continuous marine successions were deposited. Two global stratotype sections for the boundary between the Guadalupian (Middle Permian) and Lopingian (Late Permian) are located at Penglaitan and Tieqiao in the Laibin area, respectively, and thus are chosen for study. At the two locations, 14 facies are recognized in the Maokou and Heshan Formations, and they are further grouped into four facies associations (basin, lower slope, upper slope, and platform margin). Six main transgressive–regressive (TR) sequences are identified in strata from the Roadian (Middle Permian) to the Wuchiapingian (Late Permian). They are conformable marine sequences that were little influenced by regional uplift (Dongwu Movement) and so provide a good record of the sea-level changes in South China at this time. Based on the significant taxonomic selection and controversial marine faunal loss in the end-Guadalupian mass extinction, and the Middle-Late Permian sea-level changes recorded by the TR sequences in the Laibin area, it is suggested that this extinction event might have been triggered by the reduction and loss of shallow-marine habitat area caused by the end-Guadalupian regression. The global cooling and Emeishan volcanism also occurring at this time could have further enhanced this extinction event.     

On the faunal verification of the Permo-Triassic boundary in continental deposits of eastern Europe: 1. Gorokhovets-Zhukov ravine

A reference section of the Permian and Triassic continental deposits of the Zhukov ravine near the town of Gorokhovets (Vladimir Region) is described and new tetrapod localities are characterized. The position of the Permian-Triassic boundary in this section is recognized and its faunal substantiation based on vertebrates is provided for the first time. The Zhukov ravine section is unique in the fact that it shows a thick stratigraphically continuous succession of the Permo-Triassic boundary beds, with three successive tetrapod zones: the terminal Permian Chroniosuchus paradoxus and Archosaurus rossicus zones and the Early Triassic Tupilakosaurus wetlugensis Zone.     

Early Permian bioevent in the fusulinacean fauna of South China

Analysis of a large database of the stratigraphic distribution of fusulinacean Foraminifera reveals an Early Permian event of significant decline of species diversity in South China. Data from Late Carboniferous to Early Permian sections without apparent unconformity in southwest China were evaluated to determine if the apparent pattern of species disappearance was caused by bias in fossil preservation associated with Early Permian sea-level changes. Statistical analysis suggests that the Early Permian event started in the Late Sakmarian with a significant drop of species diversity in the Robustoschwagerina ziyunensis Zone and continued through the Pamirina darvasica Zone of the Artinskian and into the Brevaxina dyhrenfurthi Zone of the Early Kungarian, resulted in a total loss of about 40% species diversity in the fusulinacean fauna. The Early Permian event is the most extensive bioevent in the history of fusulinacean Foraminifera at the species level although it is less significant at the generic level. Because a similar faunal change has been found among the fusulinacean assemblages in North America and in various regions of Tethys, this event may represent a major faunal turn-over in response to the Early Permian changes in sea level and could be of a global nature. Previous recognition of this event was hampered by Early Permian unconformities in North America and other regions of Tethys.     

Did impacts, volcanic eruptions, or climate change affect mammalian evolution?

In recent years, it has become popular to attribute faunal change and mass extinction to impacts, volcanic eruptions, or climatic change. How well do these supposed causes compare to the excellent record of Cenozoic life, especially that of fossil mammals? Two different Cenozoic mammal diversity curves were compared, and important climatic, volcanic, and impact horizons were examined in detail. In no case is there a strong correlation between impacts, eruptions, or climatic events and any episode of mammalian turnover. On the contrary, most of the known impact, eruption, and climatic events of the Cenozoic occurred during intervals of faunal stability. Conversely, episodes of high turnover and faunal change among Cenozoic mammals correlate with no known extrinsic causes. Apparently, extrinsic environmental factors such as impacts, eruptions, and climate change have a minimal effect, and intrinsic biological factors must be more important.     

Evolution of the Permian and Triassic tetrapod communities of Eastern Europe

The Permo-Triassic terrestrial and freshwater tetrapod communities of Eastern Europe are reconstructed as food-webs. The Late Permian theriodont-dinocephalian community (Ocher, Mezen, Isheyevo) changes to a latest Permian theriodont-pareiasaur community (North Dvina, Vyazniki). After a major extinction, the Triassic thecodontian-dicynodont communities appear, a lystrosaurid one in the Early Triassic (Lower and ?Upper Vetluga), and a kannemeyerid one in the later Early Triassic (?Yarenga) and the Mid Triassic (Donguz, Bukobay). Similar stages are represented in the evolution of aquatic communities: the Late Permian temnospondyl community (Ocher, Isheyevo), the latest Permian chroniosuchian one (North Dvina, Vyazniki), the Lower and Middle Triassic new temnospondyl one (from Vetluga to Bukobay). The faunal changes in Eastern Europe are mirrored in other parts of the world, although there are some endemic Russian forms.     

The last diadectomorph sheds light on Late Palaeozoic tetrapod biogeography

Diadectomorpha is a clade of Late Palaeozoic vertebrates widely recognized as the sister group of crown-group Amniota and the first tetrapod lineage to evolve high-fibre herbivory. Despite their evolutionary importance, diadectomorphs are restricted stratigraphically and geographically, with all records being from the Upper Carboniferous and Lower Permian of North America and Germany. We describe a new diadectomorph, Alveusdectes fenestralis, based on a partial skull from the Upper Permian of China. The new species exhibits the derived mechanism for herbivory and is recovered phylogenetically as a deeply nested diadectid. Approximately 16 Myr younger than any other diadectomorph, Alveusdectes is the product of at least a 46 Myr ghost lineage. How much of this time was probably spent in Russia and/or central Asia will remain unclear until a specimen is described that subdivides this cryptic history, but the lineage assuredly crossed this region before entering the relatively isolated continent of North China. The discovery of Alveusdectes raises important questions regarding diadectomorph extinction dynamics including what, if any, ecological factors limited the diversity of this group in eastern Pangea. It also suggests that increased sampling in Asia will likely significantly affect our views of clade and faunal insularity leading up to the Permo-Triassic extinction.     

THE EFFECTS OF SAMPLING BIAS ON PALAEOZOIC FAUNAS AND IMPLICATIONS FOR MACROEVOLUTIONARY STUDIES

Abstract:  Trilobites, a dominant component of marine faunas during the Cambrian and Ordovician and which survived until the end of the Permian (542–251 Ma) have been used in many macroevolutionary analyses. Here, we use a discovery curve to document the sampling history of trilobites, which we consider a proxy for Palaeozoic faunas in general. At higher taxonomic ranks, orders, suborders and superfamilies, the fossil record has been completely sampled, while the family rank also shows a high level of sampling completeness, having reached an asymptote in 1970. Importantly, this levelling-off occurred even though worker effort continued to increase. However, at genus level the sampling record is incomplete, indicating that families should not be used as a proxy for genera. There is little variation among the different subsets of generic data, with the sampling history of different stratigraphic periods and among different orders being very similar. However, there is noticeable variation among geographical regions, caused by variations in worker effort, and this could cause problems when comparing speciation and diversity patterns across faunal provinces. The role of synonyms on sampling history has had little effect.     

Prehistoric fisheries in the Caribbean

We studied faunal remains from archaeological sites on five Caribbean islands, each with an early (1,850-1,280 years B.P.) and late (1,415-560 years B.P.) occupation. On each of these islands (Puerto Rico, St. Thomas, St. Martin, Saba, and Nevis), the mean size of reef fishes in the faunal remains declined from the early to the late occupation. The large samples from sites on St. Thomas and Nevis allowed examination of the size distribution of individual taxa. Samples of obligate reef fishes (Scaridae, Acanthuridae, Lutjanidae, and Serranidae) showed large reductions in size between the early and late occupations. Samples of facultative reef fishes (Carangidae and Clupeidae) showed little change in size frequency distribution. The percentage of estimated reef fish biomass in the total aquatic faunal record sharply declined in the samples from four of the islands, while on Nevis there was a slight increase. The mean trophic level of reef fishes declined from the early to the late occupations on each island. Together these patterns suggest that populations of reef fishes adjacent to occupation sites on these islands were heavily exploited in prehistoric times. Such exploitation resulted in shifts in size structure and species composition among the reef fish fauna. On some islands the decline in reef fish resources corresponded with a shift towards greater exploitation of pelagic species.     

Foraminiferal zonation of late Paleozoic depositional sequences

From the later part of the Devonian through the Permian, calcareous foraminifers became abundant and evolved rapidly. This rapid evolution of taxa forms the basis of a detailed zonation through the Carboniferous and Permian. Comparison of this evolutionary history of foraminifers, their biostratigraphic zonation, and the depositional sequences in which they occur suggests that sea-level events in late Paleozoic depositional history contributed significantly in subdividing a fairly continuous evolutionary record into a succession of about 75 identifiable foraminiferal zones during a 100–125 Myr time span. Although variable in terms of duration and vertical occurrences, the more completely recorded high-stand intervals give brief histories of the foraminiferal evolutionary record and are sandwiched between the poorly recorded or unrecorded low-stand intervals. Many of the individual foraminiferal zones are confined to a single depositional sequence.The late Paleozoic carbonate foraminiferal fossil record, as with the rest of the fossil record, is strongly affected by sediment deposition-nondeposition as a result of major changes in sea level. This incomplete fossil record is the result of repeated depositional breaks because of the way that depositional sequences form. It is not possible to ascribe macromutations, ‘punctuated’ evolution or ‘punctuated gradualism’ as the cause of this evolutionary pattern of the shelf-carbonate fossil record. This pattern is distinctive and we refer to it as ‘sequence evolution’ and ‘sequence extinction’. In the later part of the Middle Permian and in the Late Permian, the fossil record clearly illustrates that a series of faunal losses through ‘sequence extinctions’ progressively exceeded faunal replacements and new species through ‘sequence evolution’, but not a ‘mass extinction’ as is commonly ascribed to the end of the Permian Period. Most Permian faunas became extinct in the interval of 8 to 4 million years before the end of the Late Permian.     

Stepwise and large-magnitude negative shift in δ13Ccarb preceded the main marine mass extinction of the Permian–Triassic crisis interval

Large perturbations to the global carbon cycle occurred during the Permian–Triassic boundary mass extinction, the largest extinction event of the Phanerozoic Eon (542 Ma to present). Controversy concerning the pattern and mechanism of variations in the marine carbonate carbon isotope record of the Permian–Triassic crisis interval (PTCI) and their relationship to the marine mass extinction has not been resolved to date. Herein, high-resolution carbonate carbon isotope profiles (δ¹³C_carb), accompanied by lithofacies, were generated for four sections with microbialite (Taiping, Zuodeng, Cili, and Chongyang) in South China to better constrain patterns and controls on δ¹³C_carb variation in the PTCI and to test hypotheses about the temporal relationship between perturbations to the global carbon cycle and the marine mass extinction event. All four study sections exhibit a stepwise negative shift in δ¹³C_carb during the Late Permian–Early Triassic, with the shift preceding the end-Permian crisis being larger (> 3‰) than that following it (1–2‰). The pre-crisis shifts in δ¹³C_carb are widely correlatable and, hence, represent perturbations to the global carbon cycle. The comparatively smaller shifts following the crisis demonstrate that the marine mass extinction event itself had at most limited influence on the global carbon cycle, and that both Late Permian δ¹³C_carb shifts and the mass extinction must be attributed to some other cause. Their origin cannot be uniquely determined from C-isotopic data alone but appears to be most compatible with a mechanism based on episodic volcanism in combination with collapse of terrestrial ecosystems and soil erosion.     

Brachiopods, biostratigraphy, and correlation of the Permian marine deposits of Mongolia

Six horizons and 11 brachiopod zones are distinguished in the Permian of the Khangai-Khentei (Boreal) and South Mongolian (Tethys) marine basins of Mongolia on the base of monographic study of brachiopods and literary data on other faunal groups. The distinguished in the Permian basins of Mongolia biostratons are correlated with zonal brachiopod scales of neighboring regions of Northeast Asia. Possible bio-geographical relations of the Mongolian Permian marine basins and channels of brachiopods are analyzed. Forty-five brachiopod species are figured including species, which were first described from Mongolia, zonal index species and typical representatives of zonal assemblage.     

Changes in Permo-Triassic terrestrial tetrapod ecological representation in the Beaufort Group (Karoo Supergroup) of South Africa

Nicolas, M. & Rubidge, B.S. 2010: Changes in Permo-Triassic terrestrial tetrapod ecological representation in the Beaufort Group (Karoo Supergroup) of South Africa. Lethaia, Vol. 43, pp. 45–59.
For more than a century, large collections of fossils from the Beaufort Group have been built up at various museums in South Africa and have been handled as separate databases in the individual museums. Because of the unique time-extensive record of continental vertebrate biodiversity represented by the fossils of the Beaufort Group, a single standardized database has been compiled for the fossils collected from the Beaufort Group housed in South African museums. This unique data set has enabled the determination of terrestrial tetrapod ecological representation from the Middle Permian to Middle Triassic Beaufort Group of South Africa. □ Beaufort Group , biodiversity , ecology , Permo-Triassic .     

Ecological succession among late palaeozoic and mesozoic tetrapods

Natural selection and the development of new taxa are associated with ecological replacement and the increase in number of niches with time. Continental faunal interchange was possible globally because of the existence of the super-continent Pangaea during much of the Upper Palaeozoic and Mesozoic. Figures of tetrapod niches vs. time and discussion of this concept for that period are presented for the first time. Four habitat divisions are used, namely marine, fresh-water, lowland and upland.The marine habitat was colonised rather late by tetrapods and these may have been the first predators on the early bony fishes which had diversified in the Permian. The radiation of bony fishes in the Jurassic was followed by a further increase in variety of their reptilian predators. Predators seem to develop some time after the radiation of a new potential prey group.Most early amphibians occupied fresh-water habitats in “crocodile” or “frog” niches, but from the Triassic tetrapods moved from fresh-waters and lowlands into the uplands also.In terrestrial habitats, the replacement of mammal-like reptiles by dinosaurs is tentatively explained in terms of palaeoclimatology and thermoregulatory physiology. Ornithischians capable of dealing with tough vegetation evolved to occupy the new niches produced by the radiation of conifers in the Jurassic. The extinction of dinosaurs appears to have been connected with temperature and habitat changes.Conclusions are supported by a summary of published opinions on the palaeoecological roles of early tetrapods.     

A model of onshore-offshore change in faunal diversity

Onshore-offshore patterns of faunal change occurred at many taxonomic scales during the Paleozoic Era, ranging from replacement of the Cambrian evolutionary fauna by the Paleozoic fauna to the environmental expansion of many orders and classes. A simple mathematical model is constructed to investigate such change. The environmental gradient across the marine shelf-slope is treated as a linear array of discrete habitats, each of which holds a set number of species, as observed in the fossil record. During any interval of time, some portion of the species in each habitat becomes extinct by background processes, with rates of extinction varying among both clades and habitats, as also observed in the record. After extinction, species are replaced from within the habitat and from immediately adjacent habitats, with proportions dependent on surviving species. This model leads to the prediction that extinction-resistant clades will always diversify at the expense of extinction-prone clades. But if extinction intensity is highest in nearshore habitats, extinction-resistant clades will expand preferentially in the onshore direction, build up diversity there, and then diversify outward toward the offshore. Thus, onshore-offshore patterns of diversification may be the expectation for faunal change quite independently of whether or not clades originate onshore. When the model is parameterized for Paleozoic trilobites and brachiopods, numerical solutions exhibit both a pattern of faunal change and a time span for diversification similar to that seen in the fossil record. They also generate structure similar to that seen in global diversification, including logistic patterns of growth, declining origination but constant extinction within clades through time, and declining overall extinction across clades through time.     

Mass extinctions among tetrapods and the quality of the fossil record

The fossil record of tetrapods is very patchy because of the problems of preservation, in terrestrial sediments in particular, and because vertebrates are rarely very abundant. However, the fossil record of tetrapods has the advantages that it is easier to establish a phylogenetic taxonomy than for many invertebrate groups, and there is the potential for more detailed ecological analyses. The relative incompleteness of a fossil record may be assessed readily, and this can be used to test whether drops in overall diversity are related to mass extinctions or to gaps in our knowledge. Absolute incompleteness cannot be assessed directly, but a historical approach may offer clues to future improvements in our knowledge. One of the key problems facing palaeobiologists is paraphyly, the fact that many higher taxa in common use do not contain all of the descendants of the common ancestor. This may be overcome by cladistic analysis and the identification of monophyletic groups. The diversity of tetrapods increased from the Devonian to the Permian, remained roughly constant during the Mesozoic, and then began to increase in the late Cretaceous, and continued to do so during the Tertiary. The rapid radiation of 'modern' tetrapod groups--frogs, salamanders, lizards, snakes, turtles, crocodilians, birds and mammals--was hardly affected by the celebrated end-Cretaceous extinction event. Major mass extinctions among tetrapods took place in the early Permian, late Permian, early Triassic, late Triassic, late Cretaceous, early Oligocene and late Miocene. Many of these events appear to coincide with the major mass extinctions among marine invertebrates, but the tetrapod record is largely equivocal with regard to the theory of periodicity of mass extinctions.     

Chondrichthyan and actinopterygian remains from theLower Permian Copacabana Formation of Bolivia

Two horizons of the marine Lower Permian (Leonardian) Copacabana Formation have yielded fish remains, on the south-western slope of the Jacha Khatawi Hill, Yaurichambi, La Paz department, Bolivia. This fish fauna consists of teeth and scales of chondrichthyans (Eugeneodontida, Petalodontida, ? Bradyodonti, Elasmobranchii) and actinopterygians (? Plastysomidae). The Eugeneodontida are represented by a new species of a large Agassizodontidae, Parahelicoprion mariosuarezi n.sp., based on a large symphysial tooth series which resembles P. clerci from the Lower Permian Arta beds of the Urals. The Petalodontida are represented by a fragment of a large symphysial tooth which may be referred to the pristodontid genus Megactenopetalus, known else-where from the Lower Permian of the U.S.A. and China. Some isolated crushing teeth may questionably be referred to a bradyodont, possibly Helodus or Lagarodus. Some elasmobranch teeth of «Cladodus type also occur in this locality. Some hemispherical teeth of «phyllodont type and some scales are tentatively referred to the actinopterygian family Platysomidae. These findings are the first record of determinable marine Permian fishes in the Andean region of South America. The overall composition of this fauna agrees fairly well with the fish fauna known from the Marine Lower Permian of United States and eastern Asia.     

Delayed recovery of non-marine tetrapods after the end-Permian mass extinction tracks global carbon cycle

During the end-Permian mass extinction, marine ecosystems suffered a major drop in diversity, which was maintained throughout the Early Triassic until delayed recovery during the Middle Triassic. This depressed diversity in the Early Triassic correlates with multiple major perturbations to the global carbon cycle, interpreted as either intrinsic ecosystem or external palaeoenvironmental effects. In contrast, the terrestrial record of extinction and recovery is less clear; the effects and magnitude of the end-Permian extinction on non-marine vertebrates are particularly controversial. We use specimen-level data from southern Africa and Russia to investigate the palaeodiversity dynamics of non-marine tetrapods across the Permo-Triassic boundary by analysing sample-standardized generic richness, evenness and relative abundance. In addition, we investigate the potential effects of sampling, geological and taxonomic biases on these data. Our analyses demonstrate that non-marine tetrapods were severely affected by the end-Permian mass extinction, and that these assemblages did not begin to recover until the Middle Triassic. These data are congruent with those from land plants and marine invertebrates. Furthermore, they are consistent with the idea that unstable low-diversity post-extinction ecosystems were subject to boom-bust cycles, reflected in multiple Early Triassic perturbations of the carbon cycle.