Diversity partitioning in Permian and Early Triassic benthic ecosystems of the Western USA: a comparison

This paper compares the relative contributions of within-habitat diversity [alpha-diversity] and between-habitat-diversity [beta-diversity] to regional diversity [gamma-diversity] in marine benthic communities of the western US before and after the end-Permian mass extinction. We found that presumably cool-water faunas from the Permian Gerster Limestone and the Park City Formation had low alpha- and beta-diversities, comparable to those of low diverse faunas of the Early Triassic. In contrast, tropical Permian faunas had much higher alpha-diversities and a variable pattern of beta-diversity: Whereas faunas of space-limited bioherms show a positive correlation between beta-diversity and gamma-diversity, beta-diversity in level-bottom faunas is elevated only when gamma-diversity is very high (>250 species). This contrasting pattern probably reflects differential effects of interspecific competition on habitat partitioning. In low-competitive level-bottom faunas, species are able to coexist until competition forces species into their ecological optima, thereby increasing beta-diversity. This effect occurs at much lower gamma-diversities in more competitive reef-bound faunas, causing the observed positive correlation between beta- and gamma-diversity. We suggest that differences in the level of interspecific competition and hence diversity partitioning between Permian and Triassic benthic communities result from the higher average metabolic rates in the Mesozoic mollusc-dominated benthos in contrast to their Permian counterparts.     

Why was there a delayed radiation after the end‐Palaeozoic extinctions?

Data from widespread dysaerobic facies, carbon/sulphur ratios and cerium anomalies suggest that the early Triassic was a time when anoxic conditions spread widely over epicontinental seas. These conditions, associated with marine transgression following the latest Permian regression, are likely to be a prime cause of the mass extinction of Palaeozoic marine faunas. The occurrence of many Lazarus taxa in the Middle and Upper Triassic indicates, however, that the extinctions at the end of the Permian were less severe than has been widely assumed, and that the turnover from Palaeozoic to Mesozoic faunas was considerably extended in time, being finally accomplished only after the end‐Triassic mass extinction event.     

Evolutionary morphology of oblique ribs of bivalves

Fossil bivalves bearing oblique ribs first appeared in the Mid Ordovician but their diversity remained low during the Palaeozoic. The diversity soon increased after the Early Triassic, peaking in the Early Cretaceous. The Palaeozoic–Mesozoic record is dominated by burrowing bivalves (mainly pholadomyoids and trigonioids), which developed oblique ribs with symmetric profiles, probably adapted for shell reinforcement, although there are indications that the ribs of trigonioids also enhanced burrowing efficiency. After the Paleocene, the main groups of burrowing bivalves were veneroids (primarily tellinoideans and lucinoideans) and nuculoids, which generated oblique ribs of the shingled type, adapted to increase burrowing efficiency. The inferred change in function at the Mesozoic/Cenozoic boundary can be correlated with an increase in mean mobility of the bivalve faunas bearing oblique ribs through time. This implies a major ecological cause for the observed temporal patterns, which forced bivalve faunas to burrow more rapidly and efficiently. In particular, either the Phanerozoic increase in the diversity of durophagous predators or the accelerating rate of sediment reworking (both being a consequence of the Mesozoic Marine Revolution), or both, could have provided the necessary evolutionary force.     

Environmental preferences of brachiopods and bivalves across major climatic changes during the Late Palaeozoic ice age (Pennsylvanian,western Argentina)

During the late Palaeozoic ice age (LPIA), ice‐proximal marine regional communities record contrasting responses to climate change compared to ice‐distal communities. However, there is still much to be understood in distal regions in order to fully understand the palaeobiological consequences of the LPIA. Here, were analyse brachiopod and bivalve environmental preferences along the bathymetric gradient during a major glacial event and the subsequent non‐glacial interval in western Argentina. Median environmental breadths did not change with the reassembly of communities during the non‐glacial interval. Moreover, bivalves and brachiopod immigrants show similar environmental breadths although they tend to have immigrated from different palaeogeographical regions. These patterns reinforce the idea that the worldwide marine fauna was probably culled of stenotopic taxa during the LPIA. On the other hand, analysis of the preferred depths of survivors and immigrants sheds light on the substantial modification of the bathymetric diversity gradient. Among different possible explanations, the immigration of taxa with affinities for deep environments is the only one supported. In addition, results underscore the observation that the higher turnover in the offshore environment was probably driven by immigration rather than extinction. Finally, stability in environmental preferences at a regional scale is not mirrored by stability in survivors’ individual preferences, because survivors’ preferred depth is not correlated during the glacial and non‐glacial intervals. Moreover, the amount of change in survivors preferred depth is not related to their environmental breadth, nor to their occupancy. These patterns suggest: (1) instability in realized niches; and (2) individual responses of survivor genera.     

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.     

Diversity dynamics of echinoderms and evolution of marine communities

A compound analysis of two global paleontological databases (Sepkoski??s database (SDB) and The Paleobiology Database) allowed the recognition of a number of previously undescribed trends in the evolution of the phylum Echinodermata. Paleozoic echinoderms, dominated by sessile epibenthic filter feeders, played an important role in benthic communities, especially in the Ordovician and Carboniferous. Paleozoic echinoderms typically showed an increased rate of genus renewal, which significantly decreases in the Meso-Cenozoic. After the P-T crisis the echinoderms became dominated by motile taxa, while the role of infaunal forms increased. During the global turnover in the benthic communities at the K-T boundary, which was accompanied by a sharp increase in the mean alpha-diversity, many marine organisms became inhabitants of much richer (compared to the Mesozoic) communities. However, of all echinoderms, this trend is observed only in crinoids. In contrast to most large taxa, echinoderms do not show positive correlation between the duration of genera and alpha-diversity of communities, which included these genera. During the Phanerozoic the geographical distribution of echinoderms showed a sharp paleolatitudinal gradient, i.e., each period was characterized by one paleolatitudinal zone with the maximum diversity of echinoderms, and the diversity rapidly decreasing to the north and to the south of this zone. The zone of the maximum diversity of echinoderms, like of entire marine biota, during the Phanerozoic gradually moved from the tropics of the southern hemisphere to the middle latitudes of the northern hemisphere.     

Hyperbolic growth of marine and continental biodiversity through the phanerozoic and community evolution

Among diverse models that are used to describe and interpret the changes in global biodiversity through the Phanerozoic, the exponential and logistic models (traditionally used in population biology) are the most popular. As we have recently demonstrated (Markov, Korotayev, 2007), the growth of the Phanerozoic marine biodiversity at genus level correlates better with the hyperbolic model (widely used in demography and macrosociology). Here we show that the hyperbolic model is also applicable to the Phanerozoic continental biota at genus and family levels, and to the marine biota at species, genus, and family levels. There are many common features in the evolutionary dynamics of the marine and continental biotas that imply similarity and common nature of the factors and mechanisms underlying the hyperbolic growth. Both marine and continental biotas are characterized by continuous growth of the mean longevity of taxa, by decreasing extinction and origination rates, by similar pattern of replacement of dominant groups, by stepwise accumulation of evolutionary stable, adaptable and "physiologically buffered" taxa with effective mechanisms of parental care, protection of early developmental stages, etc. At the beginning of the development of continental biota, the observed taxonomic diversity was substantially lower than that predicted by the hyperbolic model. We suggest that this is due, firstly, to the fact that, during the earliest stages of the continental biota evolution, the groups that are not preserved in the fossil record (such as soil bacteria, unicellular algae, lichens, etc.) played a fundamental role, and secondly, to the fact that the continental biota initially formed as a marginal portion of the marine biota, rather than a separate system. The hyperbolic dynamics is most prominent when both marine and continental biotas are considered together. This fact can be interpreted as a proof of the integrated nature of the biosphere. In the macrosociological models, the hyperbolic pattern of the world population growth arises from a non-linear second-order positive feedback between the demographic growth and technological development (more people - more potential inventors - faster technological growth - the carrying capacity of the Earth grows faster - faster population growth - more people - more potential inventors, and so on). Based on the analogy with macrosociological models and diverse paleontological data, we suggest that the hyperbolic character of biodiversity growth can be similarly accounted for by a non-linear second-order positive feedback between the diversity growth and community structure complexity. The feedback can work via two parallel mechanisms: 1) decreasing extinction rate (more taxa- higher alpha diversity, or mean number of taxa in a community - communities become more complex and stable - extinction rate decreases - more taxa, and so on) and 2) increasing origination rate (new taxa facilitate niche construction; newly formed niches can be occupied by the next "generation" of taxa). The latter possibility makes the mechanisms underlying the hyperbolic growth of biodiversity and human population even more similar, because the total ecospace of the biota is analogous to the "carrying capacity of the Earth" in demography. As far as new species can increase ecospace and facilitate opportunities for additional species entering the community, they are analogous to the "inventors" of the demographic models whose inventions increase the carrying capacity of the Earth. The hyperbolic growth of the Phanerozoic biodiverstiy suggests that "cooperative" interactions between taxa can play an important role in evolution, along with generally accepted competitive interactions. Due to this "cooperation", the evolution of biodiversity acquires some features of a self-accelerating process. Macroevolutionary "cooperation" reveals itself in: 1) increasing stability of communities that arises from alpha diversity growth; 2) ability of species to facilitate opportunities for additional species entering the community.     

Alpha, beta, or gamma: where does all the diversity go?

Global taxonomic richness is affected by variation in three components: within-community, or alpha, diversity, between-community, or beta, diversity; and between-region, or gamma, diversity. A data set consisting of 505 faunal lists distributed among 40 stratigraphic intervals and six environmental zones was used to investigate how variation of alpha and beta diversity influenced global diversity through the Paleozoic, and especially during the Ordovician radiations. As first shown by Bambach (1977), alpha diversity increased by 50 to 70 percent in offshore marine environments during the Ordovician and then remained essentially constant of the remainder of the Paleozoic. The increase is insufficient, however, to account for the 300 percent rise observed in global generic diversity. It is shown that beta diversity among level, soft-bottom communities also increased significantly during the early Paleozoic. This change is related to enhanced habitat selection, and presumably increased overall specialization, among diversifying taxa during the Ordovician radiations. Combined with alpha diversity, the measured change in beta diversity still accounts for only about half of the increase in global diversity. Other sources of increase are probably not related to variation in gamma diversity but rather to appearance and/or expansion of organic reefs, hardground communities, bryozoan thickets, and crinoid gardens during the Ordovician.     

Diversity and structure of epifaunal communities on mollusc valves,Buzzards Bay,Massachusetts

The diversity, homogeneity and population structure of epifauna living on dead bivalve shells in a shallow marine bay are examined. Twenty-eight shells of Mercenaria mercenaria were artificially emplaced on each of three different sediment types in Buzzards Bay, Massachusetts, for one year. Living faunas (53,000 individuals assignable to 106 species) and preservable faunas (16,000 individuals assignable to 25 species) which colonized these shells are compared.Living epifaunas demonstrate a moderately good fit to the Preston truncated lognormal distribution but deviate consistently from the MacArthur broken-stick model. Homogeneity and diversity of living faunas are greater on shells resting on coarse sediments. The rarefaction methodology overestimates faunal diversity in both living and preservable faunas.Preservable epifaunas posses a higher homogeneity but one which is generally parallel to that of the living communities from which they are derived. The diversity of preservable epifaunas does not reflect the diversity of the living faunas from which they are derived.Abundant species living on shells are more likely to possess preservable hard parts than are rare species. This is indicative of the evolutionary success of various protective devices in epibenthic communities exposed to predation and environmental vagaries.     

The development of an Early Ordovician hard ground community in response to rapid sea-floor calcite precipitation

The Kanosh Shale (Upper Arenig, Lower Ordovician) of west-central Utah. USA. contains abundant carbonate hardgrounds and one of the earliest diverse hardground communities. The hardgrounds were formed through a combination of processes including the development of early digenetic nodules in clay sediments which were exhumed and concentrated as lags by storms. These cobble deposits. together with plentiful biogenic metrical. were cemented by inorganically precipitated calcite on the sea floor. forming intraformational conglomerate hardgrounds. Echinoderms may have -played a critical role in the development of hardground faunas since their disarticulated calcite ossicles were rapidly cemented by syntaxial overgrowths. forming additional cobbles and hardgrounds. The echinoderms thus may have taphonomically facilitated the development of some of the hard substrates they required. A significant portion of the hardground cements may have been derived from the early dissolution of aragonitic mollusk shells. Kanosh hardground species include the earliest bryozoans recorded on hardgrounds and large numbers of stemmed echinoderms. primarily rhipidocystid cocrinoids. Bryozoans and echinoderms covered nearly equal areas of the hardground surfaces. and there was a distinct polarization between species which preferred the upper. exposed portions of the hardgrounds and others which were most common on undercut. overhang surfaces. The Kanosh Shale hardground fossils combine elements of Late Cambrian assemblages and Middle Ordovician faunas, thus confirming predicted trends in hardground community evolution. especially the replacement of cocrinoids by bryozoans and. to a lesser extent, by other stemmed echinoderms, especially crinoids. The Kanosh community marks the transition from the Cambrian Fauna to The Paleozoic Fauna in The hardground ecosystem. *Carbonate hardgrounds, aragonite dissolution, calcite cement, Echinodermara, Trepostomata, Nicholsonclla. Dianulites. Porifpra. taphonomic facilitation, Utah. Pogonip Group, Kanosh Shale. Ordovician.     

EVOLUTION OR REVOLUTION OF AMMONOIDS AT MESOZOIC SYSTEM BOUNDARIES

1. Biological revolutions at major stratigraphical boundaries have been given numerous explanations involving endogenous biological, exogenous ecological, physical, and cosmic, as well as sedimentary or chemical factors. In an attempt to elucidate the true nature of these faunal revolutions and to assess the possible influence of biological and/or physical factors, the evolution of ammonites at the boundaries of Mesozoic stratigraphical Systems is reviewed. It is believed that the more detailed data now available can give a clearer impression of evolutionary events at these boundaries. 2. It can be demonstrated that there is neither an abrupt and world-wide extinction, nor a spontaneous replacement by new elements at these caesuras as had been generally supposed to have occurred at the Triassic-Jurassic boundary, for example. Instead, one can recognize three distinct phases in the sequence of events: (1) a continuous disappearance of the ‘antique’ faunal elements; (2) a similarly continuous, gradual, and largely synchronous appearance of, or substitution by, qualitatively distinguishable ‘modern’ elements in small populations, yet in various parallel lineages (mosaic evolution); (3) a quite revolutionary, and quantitatively very sudden, diversification of these new elements, occurring at or with some delay above the boundary. 3. Thus one can demonstrate both continuous evolution of the modern faunas (‘preadaptational phase’), as well as ‘discontinuous’ spontaneous revolution, which does not produce qualitatively new characters and must be explained by diversification or adaptive radiation. This means that no further explanation by internal factors or by higher mutation rates resulting from the impact of cosmic rays becomes necessary. It is believed that, preceded by high extinction rates, world-wide ecological factors promoting higher niche diversity suffice to explain these adaptive radiations. The high degree of provincialism, endemism and specialization of the ‘antique’ faunas and the constant survival of smooth oxycones — regarded as inhabitants of a deep-sea environment — demonstrate that marine regressions and transgressions were the most effective ecological factors. 4. If there is not too much time involved between the two events, the caesura (Faunenschnitt) between final extinction of the old faunas and the radiation of the new is the most appropriate point by which to define System boundaries.     

Fossil vertebrate faunas of the British Rhaetian (latest Triassic)

Rhaetian fossil vertebrate faunas of Britain represent rich but biased samples of taxonomic diversity during uppermost Triassic time. Review of the Westbury Formation, Penarth Group, in particular, reveals a combination of marine, littoral, and terrestrial elements. Minimally, six species of shark are preserved along with a myriacanthid holocephalan, at least four actinopterygian taxa, a characteristic lungfish, ichthyosaurs, plesiosaurs, dinosaurs, and potentially the earliest representative of the Choristodera. Rare mammalian occurrences in the Westbury beds are also possible. Severnichthys gen. nov. is a large osteichthyan, probably a palaeonisciform chondrostean, which historically has been mistaken for a labyrinthodont amphibian. At least two additional actinopterygian species and a holocephalan are known from die Lilstock Formation, and a mammal or mammallike reptile is recorded from the uppermost Blue Anchor Formation. Analysis of element abundance in the disarticulated Westbury Formation assemblage indicates that many parts of some taxa are never preserved while other elements of the same form are common. Such preservational bias suggests that many species may be missing entirely from this long-studied but poorly understood taphocoenosis. Possibly contemporaneous cave faunas from nearby upland areas give a similarly biased picture of the terrestrial fauna during this time of widespread marine transgression.     

Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda)

Living archosaurs comprise birds (dinosaurs) and crocodylians (suchians). The morphological diversity of birds and stem group dinosaurs is tremendous and well-documented. Suchia, the archosaurian group including crocodylians, is generally considered more conservative. Here, we report a new Late Triassic suchian archosaur with unusual, highly specialized features that are convergent with ornithomimid dinosaurs. Several derived features of the skull and postcranial skeleton are identical to conditions in ornithomimids. Such cases of extreme convergence in multiple regions of the skeleton in two distantly related vertebrate taxa are rare. This suggests that these archosaurs show iterative patterns of morphological evolution. It also suggests that this group of suchians occupied the adaptive zone that was occupied by ornithomimosaurs later in the Mesozoic.     

Fish species diversity in southern African estuarine systems: an evolutionary perspective

Synopsis The ichthyofauna of southern African estuaries consists primarily of juvenile marine species that use these habitats as nursery areas. The abundance and biomass of fishes in estuarine systems are typically high but species diversity is generally low, with only a few taxa dominating the community. This relatively low species diversity is attributed to the fact that estuaries in the region are unpredictable environments which lack any degree of permanence and are dominated by mobile marine eurytopes. Although stenotopes, represented mainly by small resident species from the families Gobiidae, Clinidae and Syngnathidae, are present in southern African estuaries, little speciation appears to have occurred. A possible reason for this lack of speciation, apart from the seasonal and annual variability of the abiotic environment, is that the lifetime of individual systems is usually limited to a few thousand years. In addition, fishes utilising southern African estuaries need to remain flexible (eurytopic) in their responses to an external environment which is unlikely to become more stable in the future. Thus the lack of permanence and fluctuating nature of southern African estuaries on both a spatial and temporal scale, together with the dominance of eurytopes in these systems, does not favour the evolution of new species. A preliminary examination of the available literature indicates that a detailed review of estuarine ichthyofaunal communities on a global basis will probably mirror the trends outlined above, and reveal a domination of these dynamic ecosystems by eurytopic taxa with low speciation potential.     

Late Devonian - Early Carboniferous vertebrate microremains from the Carnic Alps, northern Italy

Vertebrate microremains from the Late Devonian-Early Carboniferous of the Carnic Alps are predominantly chondrichthyan, with minor placoderm and actinopterygian remains. The faunas are sparse and, with very few exceptions, occur only in conodont-rich pelagic limestones (Pramosio Limestone) representative of the palmatolepid-bispathodid conodont biofacies. Phoebodont and jalodont chondrichthyans, also reflecting open-ocean environments, predominated during the Famennian, and eventually symmoriids seem to predominate during the Early Carboniferous. The presence of Siamodus in this assemblage gives a new locality for this genus known from few regions in the world and allows confirming its stratigraphical range (limpidus Zone) and its relation to deep-water environments. The Late Devonian vertebrate faunas are tropical and cosmopolitan, having much in common with coeval taxa from the North-Gondwanan margins and Asian terranes. Composition of the vertebrate faunas is consistent with the Carnic Alps terrane having occupied a position intermediate between Gondwana and Laurussia, as hypothesized by various authors, but because of sparsity of the taxa represented and the pronounced cosmopolitan nature of both the conodont and vertebrate faunas, the data are not compelling.     

Palaeoecology and evolution of Mesozoic salinity-controlled benthic macroinvertebrate associations

Salinity-controlled benthic macroinvertebrate associations are typical of many Mesozoic marginally marine environments. They can be recognized by abiotic criteria (e.g., environmental setting, specific autigenic minerals), by biotic criteria (faunal composition, diversity, shell morphology, size-frequency histograms, taphonomic features, associated micro fauna and microflora), and by isotope geochemistry of shells. Although salinity-controlled associations must have been widespread in the European German Triassic, very little is known about their ecology. They appear to have been dominated by the bivalve Unionites and the brachiopod Lingula. In the Jurassic, brackish-water associations are characterized by bivalves, in particular neomiodontids, corbulids, mytilids, bakevelliids, isognomonids, and oysters. In the Cretaceous, in addition, corbiculid bivalves and gastropods become increasingly abundant. Salinity-controlled benthic macroinvertebrate associations can be used to reconstruct salinity regimes of ancient environments, but emphasis should be placed on an integrated sedimentological and ecological approach, as salinity is rarely the only parameter influencing faunal composition and diversity. Although the species composition of salinity-controlled benthic associations changes distinctly through time, the composition of morphotypes remains surprisingly constant throughout the Mesozoic and up to the Recent, evidence of a conservative evolution of benthic faunas within marginal marine high-stress environments. □ Salinity, benthic associations, palaeoecology, Mesozoic.     

Evolution of ecospace occupancy by Mesozoic marine tetrapods

Ecology and morphology are different, and yet in comparative studies of fossil vertebrates the two are often conflated. The macroevolution of Mesozoic marine tetrapods has been explored in terms of morphological disparity, but less commonly using ecological‐functional categories. Here we use ecospace modelling to quantify ecological disparity across all Mesozoic marine tetrapods. We document the explosive radiation of marine tetrapod groups in the Triassic and their rapid attainment of high ecological disparity. Late Triassic extinctions led to a marked decline in ecological disparity, and the recovery of ecospace and ecological disparity was sluggish in the Early Jurassic. High levels of ecological disparity were again achieved by the Late Jurassic and maintained during the Cretaceous, when the ecospace became saturated by the Late Cretaceous. Sauropterygians, turtles and ichthyosauromorphs were the largest contributors to ecological disparity. Throughout the Mesozoic, we find that established groups remained ecologically conservative and did not explore occupied or vacant niches. Several parts of the ecospace remained vacant for long spans of time. Newly evolved, radiating taxa almost exclusively explored unoccupied ecospace, suggesting that abiotic releases are needed to empty niches and initiate diversification. In the balance of evolutionary drivers in Mesozoic marine tetrapods, abiotic factors were key to initiating diversification events, but biotic factors dominated the subsequent generation of ecological diversity.