THE UPPER CAMBRIAN TRILOBITE OLENUS AT ANDRARUM,SWEDEN: A CASE OF ITERATIVE EVOLUTION?

Abstract:  The morphological changes in four species-level lineages of the Late Cambrian trilobite Olenus from the Alum Shale at Andrarum, south-east Scania, Sweden, have been analysed with the aim of re-examining the classical study of iterative evolution by Kaufmann (1933 a ). New material comprising 647 pygidia was collected for a restudy of Olenus , and for comparison the skeletal development of 918 cephala and 568 pygidia from the associated Homagnostus obesus were included. The re-examination shows that (1) only pygidia of the O. transversus – O. truncatus – O. wahlenbergi lineage display a statistically significant directional change towards a narrower shape; (2) all other lineages, including the agnostids, show net morphological stasis; (3) hence iterative directional evolution cannot be confirmed. Based on morphological investigations it is tempting to conclude that the lineage O. transversus – O. truncatus – O. wahlenbergi represents just one species displaying gradual morphological changes. Whether this is an evolutionary or an ecophenotypic phenomenon remains uncertain. The vertical distributions of olenids and agnostids are compared by using vanadium/(vanadium + nickel) ratios from the Olenus Zone of the nearby Gislövshammar-2 drill-core. It appears that incursions of olenids were linked to increases in bottom-water oxygen levels and that Homagnostus obesus was tolerant of lower dysoxia than olenids.     

From success to persistence: Identifying an evolutionary regime shift in the diverse Paleozoic aquatic arthropod group Eurypterida,driven by the Devonian biotic crisis

Mass extinctions have altered the trajectory of evolution a number of times over the Phanerozoic. During these periods of biotic upheaval a different selective regime appears to operate, although it is still unclear whether consistent survivorship rules apply across different extinction events. We compare variations in diversity and disparity across the evolutionary history of a major Paleozoic arthropod group, the Eurypterida. Using these data, we explore the group's transition from a successful, dynamic clade to a stagnant persistent lineage, pinpointing the Devonian as the period during which this evolutionary regime shift occurred. The late Devonian biotic crisis is potentially unique among the “Big Five” mass extinctions in exhibiting a drop in speciation rates rather than an increase in extinction. Our study reveals eurypterids show depressed speciation rates throughout the Devonian but no abnormal peaks in extinction. Loss of morphospace occupation is random across all Paleozoic extinction events; however, differential origination during the Devonian results in a migration and subsequent stagnation of occupied morphospace. This shift appears linked to an ecological transition from euryhaline taxa to freshwater species with low morphological diversity alongside a decrease in endemism. These results demonstrate the importance of the Devonian biotic crisis in reshaping Paleozoic ecosystems.     

Revision of <Emphasis Type="Italic">Pseudorhaptagnostus</Emphasis> Lermontova (?Trilobita,Agnostida)

The revision of the type collections on the genus Pseudorhaptagnostus Lermontova, 1940 has shown the validity of the type species. Originally indicated type species P. simplex Lermontova, 1951 appeared to be erroneously combination of cephala and pygidia from different species; recombined set of cephala and pygidia for P. simplex synonymized it with P. punctatus Lermontova, 1940, and therefore renamed the type species as the latter one. Also Pseudorhaptagnostus has shown to differ from its close morphological affinities Neoagnostus, Norilagnostus, Machairagnostus, Idolagnostus. New introduction of their morphological difference allowed us to rearrange the species assignment of these genera.     

EXTINCTION SPACE—A METHOD FOR THE QUANTIFICATION AND CLASSIFICATION OF CHANGES IN MORPHOSPACE ACROSS EXTINCTION BOUNDARIES

Three main modes of extinction are responsible for reductions in morphological disparity: (1) random (caused by a nonselective extinction event); (2) marginal (a symmetric, selective extinction event trimming the margin of morphospace); and (3) lateral (an asymmetric, selective extinction event eliminating one side of the morphospace). These three types of extinction event can be distinguished from one another by comparing changes in three measures of morphospace occupation: (1) the sum of range along the main axes; (2) the sum of variance; and (3) the position of the centroid. Computer simulations of various extinction events demonstrate that the pre‐extinction distribution of taxa (random or normal) in the morphospace has little influence on the quantification of disparity changes, whereas the modes of the extinction events play the major role. Together, the three disparity metrics define an “extinction‐space” in which different extinction events can be directly compared with one another. Application of this method to selected extinction events (Frasnian‐Famennian, Devonian‐Carboniferous, and Permian‐Triassic) of the Ammonoidea demonstrate the similarity of the Devonian events (selective extinctions) but the striking difference from the end‐Permian event (nonselective extinction). These events differ in their mode of extinction despite decreases in taxonomic diversity of similar magnitude.     

Extant‐only comparative methods fail to recover the disparity preserved in the bird fossil record

Most extant species are in clades with poor fossil records, and recent studies of comparative methods show they have low power to infer even highly simplified models of trait evolution without fossil data. Birds are a well‐studied radiation, yet their early evolutionary patterns are still contentious. The fossil record suggests that birds underwent a rapid ecological radiation after the end‐Cretaceous mass extinction, and several smaller, subsequent radiations. This hypothesized series of repeated radiations from fossil data is difficult to test using extant data alone. By uniting morphological and phylogenetic data on 604 extant genera of birds with morphological data on 58 species of extinct birds from 50 million years ago, the “halfway point” of avian evolution, I have been able to test how well extant‐only methods predict the diversity of fossil forms. All extant‐only methods underestimate the disparity, although the ratio of within‐ to between‐clade disparity does suggest high early rates. The failure of standard models to predict high early disparity suggests that recent radiations are obscuring deep time patterns in the evolution of birds. Metrics from different models can be used in conjunction to provide more valuable insights than simply finding the model with the highest relative fit.     

Evolution of generic and species diversity in agnathans (Heterostraci: Orders Cyathaspidiformes,Pteraspidiformes)

The evolution of generic and species diversity in agnathans (orders Cyathaspidiformes and Pteraspidiformes) is analyzed. Cyathaspids appeared in the Silurian and achieved the peak of diversity in the first half of the Early Devonian (Lochkovian Stage and its analogues), as did pteraspids. Cyathaspids are absent from later beds. Pteraspids persisted to the end of the Early Devonian, although they sharply decreased in number in the second half of the Early Devonian (Pragian Stage and its analogues in Europe and America). Adaptive features of cyathaspids and pteraspids are considered. Some data on accompanying groups are discussed. It is proposed that these heterostracan orders became extinct mostly because of archaic locomotor adaptations and insufficient protection from predators. The insufficient protection primarily concerns juvenile cyathaspids. Cyathaspid and pteraspid genera included in the diagrams of changes in taxonomic diversity are listed.     

New crinoids from the Baltic region (Estonia): fossil tip‐dating phylogenetics constrains the origin and Ordovician–Silurian diversification of the Flexibilia (Echinodermata)

This study documents previously unknown taxonomic and morphological diversity among early Palaeozoic crinoids. Based on highly complete, well preserved crown material, we describe two new genera from the Ordovician and Silurian of the Baltic region (Estonia) that provide insight into two major features of the geological history of crinoids: the early evolution of the flexible clade during the Great Ordovician Biodiversification Event (GOBE), and their diversification history surrounding the end‐Ordovician mass extinction. The unexpected occurrence of a highly derived sagenocrinid, Tintinnabulicrinus estoniensis gen. et. sp. nov., from Upper Ordovician (lower Katian) rocks of the Baltic palaeocontinent provides high‐resolution temporal, taxonomic and palaeobiogeographical constraints on the origin and early evolution of the Flexibilia. The Silurian (lower Rhuddanian, Llandovery) Paerticrinus arvosus gen. et sp. nov. is the oldest known Silurian crinoid from Baltica and thus provides the earliest Baltic record of crinoids following the aftermath of the end‐Ordovician mass extinction. A Bayesian ‘fossil tip‐dating’ analysis implementing the fossilized birth–death process and a relaxed morphological clock model suggests that flexibles evolved c. 3 million years prior to their oldest fossil record, potentially involving an ancestor–descendant relationship (via ‘budding’ cladogenesis or anagenesis) with the paraphyletic cladid Cupulocrinus. The sagenocrinid subclade rapidly diverged from ‘taxocrinid’ grade crinoids during the final stages of the GOBE, culminating in maximal diversity among Ordovician crinoid faunas on a global scale. Remarkably, diversification patterns indicate little taxonomic turnover among flexibles across the Late Ordovician mass extinction. However, the elimination of closely related clades may have helped pave the way for their subsequent Silurian diversification and increased ecological role in post‐Ordovician Palaeozoic marine communities. This study highlights the significance of studies reporting faunas from undersampled palaeogeographical regions for clade‐based phylogenetic studies and improving estimates of global biodiversity through geological time.     

Moving towards a better understanding of iterative evolution: an example from the late Silurian Monograptidae (Graptolithina) of the Baltic Basin

Iterative evolution has proved a difficult evolutionary phenomenon to study and interpret. Inferences of causality vary from study to study and quantitatively based phylogenetic reconstruction has never been attempted. In an effort to better understand iterative evolution we employed stratocladistics, gap analysis and disparity analysis to study the case of the Monograptidae in the aftermath of the late Silurian Cyrtograptus lundgreni extinction event. Our combination of gap analytical and stratocladistic techniques allowed us to elucidate the evolutionary relationships between the studied taxa. Based on our stratocladistic results we recommend the generic reassignment of five monograptid taxa. The stratocladistic results, in conjunction with morphological disparity analysis suggest the presence of a persistent developmental potential for the emergence of iteratively evolving characters. This persistent potential appears to be limited by extrinsic ecological constraints, which would have relaxed in the aftermath of the C. lundgreni extinction event. Our findings indicate that iterative evolution in the late Silurian Monograptidae is a product of the interaction of both intrinsic and extrinsic constraints on the acquisition of the iteratively evolving character, with the exact causality being dependent on the particular character.     

Decoupling of taxonomic diversity and morphological disparity during decline of the Cambrian trilobite family Pterocephaliidae

Although discordance between taxonomic diversity and morphological disparity is common, little is known about the underlying dynamics that drive this decoupling. Early in the history of the Cambrian trilobite family Pterocephaliidae, there was an increase in taxonomic diversity and morphological diversity. As taxonomic diversity declined in the later history of the clade, range of variation stayed high and disparity continued to increase. However, per‐branch rates of morphological evolution estimated from a recent phylogeny decreased with time. Neither within‐trait nor within‐species variation increased or decreased, suggesting that the declining rates of morphological evolution were more likely related to ecological opportunity or niche partitioning, rather than increasing intrinsic constraints. This is further supported by evidence for increased biofacies associations throughout the time period. Thus, the high disparity seen at low taxonomic diversity late in the history of this clade was due to extinction – either random or targeting mean forms – rather than increased rates of morphological evolution. This pattern also provides a scenario that could account for instances of low taxonomic diversity but high morphological disparity in modern groups.     

Patterns,trends, and rates of evolution within the Actinistia

Synopsis The interrelationships of 31 actinistian species (including Latimeria chalumnae) are analyzed based on a cladistic analysis of 75 osteological characters. Inference of evolutionary trends (e.g., modification of body shape and skull morphology) from the phylogenetic patterns demonstrates that the morphology of actinistians is less conservative than has been proposed previously. This empirical cladistic approach supports two distinct tempos of evolution during an evolutionary history of 380 million years. Along a phylogenetic pathway originating with a Devonian stem-species and ending with the living Latimeria chalumnae (including 101 morphological changes and 18 cladogenetic events), the first tempo occurred during the Devonian — Permian periods as a decreasing rate of morphological changes, which was followed by a stabilizing tempo during the Permian — Recent periods. The decreasing tempo is characterized by a sequence of gradual versus quantum temporal changes and low versus faster rates, whereas the stabilizing tempo primarily is gradual and low. In contrast to a common assumption, no significant correlation was found between the rates of morphological evolution and the temporal diversity of species.     

Evolution of taxonomic diversity in amphiaspids (Agnatha,Heterostraci: Amphiaspidiformes) and the causes of extinction in ecologically favorable conditions

Changes in the taxonomic composition of Early Devonian amphiaspids represented in Siberia by two assemblages (from the northwestern Siberian Platform and Taimyr) are analyzed. The study is performed at the generic level and represented by diagrams. Changes in the amphiaspid composition are compared with changes in the development of the Taimyr and platform paleobasins. It is shown that shifts in ecological conditions at the stages of extinction of amphiaspids occurred within the limits of changes of abiotic factors (depth, salinity, etc.) that are usual for heterostracans. The disappearance (extinction) of amphiaspids is attributable to the level of their morphological organization, which caused inefficient adaptations to the developing paleoecosystems. The disappearance (extinction) of groups resulting from inadequate vital adaptations to the changing structure of paleoecosystems is considered to be a general law of evolution.     

Phanerozoic changes in the high-rank suprageneric diversity structure of brachiopods: Linear and non-linear effects

Phanerozoic evolution of brachiopods produced many linear (established by a comparison of successive geologic time units) and non-linear (established by a comparison of non-successive geologic time units) effects, which can be examined quantitatively by using the similarity coefficients (Czekanowski's Quantified Coefficient and Gower Index) and correlation tools. The high-rank suprageneric diversity structure accounts for a number of superfamilies in each of 26 orders for every epoch of geological time. The intensity of turnovers in this structure was generally low during the entire Phanerozoic. It was slightly stronger during the Early Paleozoic, but close to zero during the Cenozoic, when the high-rank suprageneric diversity structure of brachiopods stabilized finally. Significant turnovers took place at the Middle Cambrian–Early Ordovician, the Late Ordovician–Early Silurian, the Late Silurian–Early Devonian, the Middle Devonian–Mississippian, and the Permian–Triassic transitions. Influences of mass extinctions, both major like those End Ordovician or Permian/Triassic and minor like Early Jurassic or Jurassic/Cretaceous, on the high-rank suprageneric diversity structure of brachiopods is registered. The strongest was the consequences of the Permian/Triassic catastrophe, which perhaps even reset the brachiopod evolution. No evident direct relationships are established between intensity of turnovers and eustatic fluctuations. However, the changes in the diversity structure recorded with the Gower Index provide evidence that eustatic lowstands were more favorable for intensification in these changes.     

The complex effects of mass extinctions on morphological disparity

Studies of biodiversity through deep time have been a staple for biologists and paleontologists for over 60 years. Investigations of species richness (diversity) revealed that at least five mass extinctions punctuated the last half billion years, each seeing the rapid demise of a large proportion of contemporary taxa. In contrast to diversity, the response of morphological diversity (disparity) to mass extinctions is unclear. Generally, diversity and disparity are decoupled, such that diversity may decline as morphological disparity increases, and vice versa. Here, we develop simulations to model disparity changes across mass extinctions using continuous traits and birth-death trees. We find no simple null for disparity change following a mass extinction but do observe general patterns. The range of trait values decreases following either random or trait-selective mass extinctions, whereas variance and the density of morphospace occupation only decline following trait-selective events. General trends may differentiate random and trait-selective mass extinctions, but methods struggle to identify trait selectivity. Long-term effects of mass extinction trait selectivity change support for phylogenetic comparative methods away from the simulated Brownian motion toward Ornstein-Uhlenbeck and Early Burst models. We find that morphological change over mass extinction is best studied by quantifying multiple aspects of morphospace occupation.     

Did the Late Ordovician mass extinction event trigger the earliest evolution of ‘strophodontoid’ brachiopods?

‘Strophodontoid’ brachiopods represented the majority of strophomenide brachiopods in the Silurian and Devonian periods. They are characterized by denticles developed along the hinge line. The evolution of denticles correlated with the disappearance of dental plates and teeth and were already present when the clade originated in the Late Ordovician. Specimens of Eostropheodonta parvicostellata from the Kuanyinchiao Bed (early–middle Hirnantian, uppermost Ordovician) in the Hetaoba Section, Meitan, Guizhou Province, South China, display clear fossil population variation, during a process of loss of dental plates and the development of denticles. Three phenotypes of E. parvicostellata are recognized in a single fossil bed, likely heralding a speciation process. Non-metric multidimensional scaling (NMDS) based on five key characters of genera of the Family Leptostrophiidae shows a much wider morphospace for Silurian genera than for those in the Devonian. Phylogenetic analysis of the Family Leptostrophiidae supports the NMDS analysis and mostly tracks their geological history. The fossil population differentiation in E. parvicostellata discovered between the two phases of the Late Ordovician mass extinction event (LOME) linked to a major glaciation, suggests a Hirnantian origination of the ‘strophodontoid’ morphology, and links microevolutionary change to a macroevolutionary event.     

The first 50Myr of dinosaur evolution: macroevolutionary pattern and morphological disparity

The evolutionary radiation of dinosaurs in the Late Triassic and Early Jurassic was a pivotal event in the Earth's history but is poorly understood, as previous studies have focused on vague driving mechanisms and have not untangled different macroevolutionary components (origination, diversity, abundance and disparity). We calculate the morphological disparity (morphospace occupation) of dinosaurs throughout the Late Triassic and Early Jurassic and present new measures of taxonomic diversity. Crurotarsan archosaurs, the primary dinosaur 'competitors', were significantly more disparate than dinosaurs throughout the Triassic, but underwent a devastating extinction at the Triassic-Jurassic boundary. However, dinosaur disparity showed only a slight non-significant increase after this event, arguing against the hypothesis of ecological release-driven morphospace expansion in the Early Jurassic. Instead, the main jump in dinosaur disparity occurred between the Carnian and Norian stages of the Triassic. Conversely, dinosaur diversity shows a steady increase over this time, and measures of diversification and faunal abundance indicate that the Early Jurassic was a key episode in dinosaur evolution. Thus, different aspects of the dinosaur radiation (diversity, disparity and abundance) were decoupled, and the overall macroevolutionary pattern of the first 50Myr of dinosaur evolution is more complex than often considered.     

A quantitative comparison of the ontogeny of two closely-related Upper Devonian phacopid trilobites

The best insight into the development of Devonian phacopids has been obtained from Trimerocephalus lelievrei Crônier & Feist, 1997, a Famennian phacopine from Morocco, where changes in size and shape have been quantified. In this study, a morphometric approach has been used: (1) to retrodeform and then establish patterns of morphological variation in a well preserved but tectonically deformed assemblage belonging to another phacopine species Weyerites ensae (Richter & Richter, 1926), a Famennian phacopine from Thuringia, and (2) to establish patterns of developmental and evolutionary changes within two closely related species: Weyerites ensae and Trimerocephalus lelievrei. The method of retrodeformation using a set of discrete points presumed to be homologous on all studied individuals, has demonstrated that the next analyses are possible on the retrodeformed material as compared to the undeformed material. Morphometric analysis based on outline analysis has permitted demonstration of progressive shape change in agreement with ontogenetic ordination and a comparison of changes in size and shape in Weyerites ensae. The main changes in shape appear to occur in the meraspid period, whereas increase in size takes place mainly in the holaspid period. This pattern, already reported for Trimerocephalus lelievrei, can be generalized for phacopine trilobites from the Late Devonian. Moreover, the comparison of the two ontogenetic trajectories has shown that most of the differences are related to 'structural' changes, probably linked to a relative pre- post-displacement. The results suggest that ecological adaptation may be studied by examining the changes in development that occur within species through time and space.     

Substructure in exines of Artemisia vulgaris (Asteraceae)

Of the Silurian and Devonian macroplant genera described in the literature, about half have been recorded with spores. These in situ spores can be compared at generic level with only about a quarter of the dispersed spore genera described for the same period. Emphasis is here placed on the structure and sculpture of in situ and dispersed Silurian, Gedinnian and Siegenian spores, where records of macroplants with in situ spores are high compared with the Middle and Upper Devonian.A comparative study of the structure and sculpture of the in situ and dispersed spores, together with information from the macroplants recorded, suggests a fairly simple Silurian and Gedinnian flora, with the evolution of a more advanced upland flora during Siegenian times.     

Phylogenetic Clustering of Origination and Extinction across the Late Ordovician Mass Extinction

Mass extinctions can have dramatic effects on the trajectory of life, but in some cases the effects can be relatively small even when extinction rates are high. For example, the Late Ordovician mass extinction is the second most severe in terms of the proportion of genera eliminated, yet is noted for the lack of ecological consequences and shifts in clade dominance. By comparison, the end-Cretaceous mass extinction was less severe but eliminated several major clades while some rare surviving clades diversified in the Paleogene. This disconnect may be better understood by incorporating the phylogenetic relatedness of taxa into studies of mass extinctions, as the factors driving extinction and recovery are thought to be phylogenetically conserved and should therefore promote both origination and extinction of closely related taxa. Here, we test whether there was phylogenetic selectivity in extinction and origination using brachiopod genera from the Middle Ordovician through the Devonian. Using an index of taxonomic clustering (R_CL) as a proxy for phylogenetic clustering, we find that A) both extinctions and originations shift from taxonomically random or weakly clustered within families in the Ordovician to strongly clustered in the Silurian and Devonian, beginning with the recovery following the Late Ordovician mass extinction, and B) the Late Ordovician mass extinction was itself only weakly clustered. Both results stand in stark contrast to Cretaceous-Cenozoic bivalves, which showed significant levels of taxonomic clustering of extinctions in the Cretaceous, including strong clustering in the mass extinction, but taxonomically random extinctions in the Cenozoic. The contrasting patterns between the Late Ordovician and end-Cretaceous events suggest a complex relationship between the phylogenetic selectivity of mass extinctions and the long-term phylogenetic signal in origination and extinction patterns.