The evolution of methods for establishing evolutionary timescales

The fossil record is well known to be incomplete. Read literally, it provides a distorted view of the history of species divergence and extinction, because different species have different propensities to fossilize, the amount of rock fluctuates over geological timescales, as does the nature of the environments that it preserves. Even so, patterns in the fossil evidence allow us to assess the incompleteness of the fossil record. While the molecular clock can be used to extend the time estimates from fossil species to lineages not represented in the fossil record, fossils are the only source of information concerning absolute (geological) times in molecular dating analysis. We review different ways of incorporating fossil evidence in modern clock dating analyses, including node-calibrations where lineage divergence times are constrained using probability densities and tip-calibrations where fossil species at the tips of the tree are assigned dates from dated rock strata. While node-calibrations are often constructed by a crude assessment of the fossil evidence and thus involves arbitrariness, tip-calibrations may be too sensitive to the prior on divergence times or the branching process and influenced unduly affected by well-known problems of morphological character evolution, such as environmental influence on morphological phenotypes, correlation among traits, and convergent evolution in disparate species. We discuss the utility of time information from fossils in phylogeny estimation and the search for ancestors in the fossil record. This article is part of the themed issue ‘Dating species divergences using rocks and clocks’

Disparities in the analysis of morphological disparity

Analyses of morphological disparity have been used to characterize and investigate the evolution of variation in the anatomy, function and ecology of organisms since the 1980s. While a diversity of methods have been employed, it is unclear whether they provide equivalent insights. Here, we review the most commonly used approaches for characterizing and analysing morphological disparity, all of which have associated limitations that, if ignored, can lead to misinterpretation. We propose best practice guidelines for disparity analyses, while noting that there can be no ‘one-size-fits-all’ approach. The available tools should always be used in the context of a specific biological question that will determine data and method selection at every stage of the analysis

Digestive and appendicular soft-parts, with behavioural implications, in a large Ordovician trilobite from the Fezouata Lagerstätte, Morocco

Trilobites were one of the most successful groups of marine arthropods during the Palaeozoic era, yet their soft-part anatomy is only known from a few exceptionally-preserved specimens found in a handful of localities from the Cambrian to the Devonian. This is because, even if the sclerotized appendages were not destroyed during early taphonomic stages, they are often overprinted by the three-dimensional, mineralised exoskeleton. Inferences about the ventral anatomy and behavioural activities of trilobites can also be derived from the ichnological record, which suggests that most Cruziana and Rusophycus trace fossils were possibly produced by the actions of trilobites. Three specimens of the asaphid trilobite Megistaspis (Ekeraspis) hammondi, have been discovered in the Lower Ordovician Fezouata Konservat-Lagerstätte of southern Morocco, preserving appendages and digestive tract. The digestive structures include a crop with digestive caeca, while the appendages display exopodal setae and slight heteropody (cephalic endopods larger and more spinose than thoracic and pygidial ones). The combination of these digestive structures and the heteropody has never been described together among trilobites, and the latter could assist in the understanding of the production of certain comb-like traces of the Cruziana rugosa group, which are extraordinarily abundant on the shallow marine shelves around Gondwana.This work has been supported by the Spanish Ministry of Economy and Competitiveness, project number CGL2012- 39471/BTE.Peer reviewe

A xandarellid artiopodan from Morocco – a middle Cambrian link between soft-bodied euarthropod communities in North Africa and South China

NB. A corrigendum [correction] for this article was published online on 09 May 2017; this has been attached to this article as an additional file. This work is licensed under a Creative Commons Attribution 4.0 International License. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in the credit line; if the material is not included under the Creative Commons license, users will need to obtain permission from the license holder to reproduce the material. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/ © The Author(s) 2017. The attached file is the published version of the article

Phylogenetic and Biogeographic Analysis of Sphaerexochine Trilobites

BACKGROUND: Sphaerexochinae is a speciose and widely distributed group of cheirurid trilobites. Their temporal range extends from the earliest Ordovician through the Silurian, and they survived the end Ordovician mass extinction event (the second largest mass extinction in Earth history). Prior to this study, the individual evolutionary relationships within the group had yet to be determined utilizing rigorous phylogenetic methods. Understanding these evolutionary relationships is important for producing a stable classification of the group, and will be useful in elucidating the effects the end Ordovician mass extinction had on the evolutionary and biogeographic history of the group. METHODOLOGY/PRINCIPAL FINDINGS: Cladistic parsimony analysis of cheirurid trilobites assigned to the subfamily Sphaerexochinae was conducted to evaluate phylogenetic patterns and produce a hypothesis of relationship for the group. This study utilized the program TNT, and the analysis included thirty-one taxa and thirty-nine characters. The results of this analysis were then used in a Lieberman-modified Brooks Parsimony Analysis to analyze biogeographic patterns during the Ordovician-Silurian. CONCLUSIONS/SIGNIFICANCE: The genus Sphaerexochus was found to be monophyletic, consisting of two smaller clades (one composed entirely of Ordovician species and another composed of Silurian and Ordovician species). By contrast, the genus Kawina was found to be paraphyletic. It is a basal grade that also contains taxa formerly assigned to Cydonocephalus. Phylogenetic patterns suggest Sphaerexochinae is a relatively distinctive trilobite clade because it appears to have been largely unaffected by the end Ordovician mass extinction. Finally, the biogeographic analysis yields two major conclusions about Sphaerexochus biogeography: Bohemia and Avalonia were close enough during the Silurian to exchange taxa; and during the Ordovician there was dispersal between Eastern Laurentia and the Yangtze block (South China) and between Eastern Laurentia and Avalonia

Exceptionally Preserved Cambrian Trilobite Digestive System Revealed in 3D by Synchrotron-Radiation X-Ray Tomographic Microscopy

The Cambrian ‘Orsten’ fauna comprises exceptionally preserved and phosphatised microscopic arthropods. The external morphology of these fossils is well known, but their internal soft-tissue anatomy has remained virtually unknown. Here, we report the first non-biomineralised tissues from a juvenile polymerid trilobite, represented by digestive structures, glands, and connective strands harboured in a hypostome from the Swedish ‘Orsten’ fauna. Synchrotron-radiation X-ray tomographic microscopy enabled three-dimensional internal recordings at sub-micrometre resolution. The specimen provides the first unambiguous evidence for a J-shaped anterior gut and the presence of a crop with a constricted alimentary tract in the Trilobita. Moreover, the gut is Y-shaped in cross section, probably due to a collapsed lumen of that shape, another feature which has not previously been observed in trilobites. The combination of anatomical features suggests that the trilobite hypostome is functionally analogous to the labrum of euarthropods and that it was a sophisticated element closely integrated with the digestive system. This study also briefly addresses the preservational bias of the ‘Orsten’ fauna, particularly the near-absence of polymerid trilobites, and the taphonomy of the soft-tissue-harbouring hypostome

The Origin and Initial Rise of Pelagic Cephalopods in the Ordovician

BACKGROUND: During the Ordovician the global diversity increased dramatically at family, genus and species levels. Partially the diversification is explained by an increased nutrient, and phytoplankton availability in the open water. Cephalopods are among the top predators of today's open oceans. Their Ordovician occurrences, diversity evolution and abundance pattern potentially provides information on the evolution of the pelagic food chain. METHODOLOGY/PRINCIPAL FINDINGS: We reconstructed the cephalopod departure from originally exclusively neritic habitats into the pelagic zone by the compilation of occurrence data in offshore paleoenvironments from the Paleobiology Database, and from own data, by evidence of the functional morphology, and the taphonomy of selected cephalopod faunas. The occurrence data show, that cephalopod associations in offshore depositional settings and black shales are characterized by a specific composition, often dominated by orthocerids and lituitids. The siphuncle and conch form of these cephalopods indicate a dominant lifestyle as pelagic, vertical migrants. The frequency distribution of conch sizes and the pattern of epibionts indicate an autochthonous origin of the majority of orthocerid and lituitid shells. The consistent concentration of these cephalopods in deep subtidal sediments, starting from the middle Tremadocian indicates the occupation of the pelagic zone early in the Early Ordovician and a subsequent diversification which peaked during the Darriwilian. CONCLUSIONS/SIGNIFICANCE: The exploitation of the pelagic realm started synchronously in several independent invertebrate clades during the latest Cambrian to Middle Ordovician. The initial rise and diversification of pelagic cephalopods during the Early and Middle Ordovician indicates the establishment of a pelagic food chain sustainable enough for the development of a diverse fauna of large predators. The earliest pelagic cephalopods were slowly swimming vertical migrants. The appearance and early diversification of pelagic cephalopods is interpreted as a consequence of the increased food availability in the open water since the latest Cambrian

Strontium isotopic geochemistry of Pan-African/Brasiliano rocks, Chapada copper deposit, Goiás, Brazil

Das Trägergestein der Chapada Kupferlagerstätte in Goiás ist in metamorpher Hoch-Amphibolitfazies ausgebildet. Gesamtgesteins- und individuelle Mineral Rb-Sr-Isochronen mit einem 87 Sr/ 86 Sr-Initialwert (I) von 0,70414±0.00005 ergeben für die Kristallisation der Vulkanite und Magmatite, die die Metamorphose vordatieren, ein Alter von 561±9 Mio Jahre. Die Datierung der Metamorphose während der panafrikanisch/brasilianischen Orogenese (700 bis 450 Mio Jahre) ergibt 532±1 Mio Jahre. Das Mineral-Isochronenalter eines Dioritkörpers porphyrischen Gefüges, der in die Schiefer intrudierte, beträgt 534±16 Mio Jahre (I=0,70415 ±0,00004). Das Ergebnis niedriger Initial-Werte dieser Isochronen ergibt für die Schiefer bei Chapada ein Ausgangs-Material im Bereich des oberen Mantels oder der tieferen Kruste; diese Ausgangswerte sind charakteristisch für magmatische Tätigkeit in Verbindung mit Inselbögen. Das unterkambrische Alter des Trägergesteins bei Chapada steht im Widerspruch zu dem bisher angenommenen mittelproterozoischen Alter und ist ein Hinweis auf eine mögliche Phase intensiverer pan-afrikanisch/brasilianischer magmatischer Aktivität in Zentral-Brasilien als ursprünglich angenommen wurde. Whole rock and mineral Rb-Sr isochrons from the upper amphibolite grade metamorphic host rocks of the Chapada copper deposit in Goiás, Brazil, yield an age of 561±9 Ma, with a 87 Sr/ 86 Sr initial (I) of 0.70414±0.00005, for the crystallization of pre-metamorphic volcanic and igneous rocks and 532 ±1 Ma for their metamorphism during the Pan-African/Brasiliano orogeny (700-450 Ma). A porphyritic diorite stock which intruded the host schists yields a mineral isochron age of 534±16 Ma (I=0.70415±0.00004). The low I values determined from these isochrons suggest that the host schists at Chapada were derived from upper mantle or lower crust material and are characteristic of magmatic products associated with island arcs. The Early Cambrian age of the host rocks at Chapada contradicts the previously assigned mid-Proterozoic age and suggests that Pan-African/Brasiliano magmatic activity may have been more extensive in central Brazil than previously thought. Les roches qui contiennent le gisement de cuivre de Chapada (Goias, Brésil) sont des schistes cristallins appartenent au facies dupérieur des amphibolites. Dans des volcanites et des roches orthomagmatiques appartenant à cet ensemble, des isochrones Rb-Sr sur roches totales et sur minéraux donnent un âge de 561±9 Ma avec un rapport initial de 0,70414±0,00005 pour la cristallisation pré-métamorphique et un âge de 532±1 Ma pour le métamorphisme, lié à l'orogenèse pan-africaine / brésilienne. Un pluton de diorite porphyrique, qui intrude les schistes cristallins donne, par isochrone sur minéraux, un âge de 534±16 Ma (rapport initial de 0,70415±0,00004). Les valeurs basses des rapports initiaux de ces diverses roches suggèrent que les schistes de Chapada sont dérivés de matériaux mantelliques ou crustaux profonds et présentent les caractères des produits magmatiques associés aux arcs insulaires. Leur âge éo-cambrien, en contradiction avec l'âge mésoprotérozoïque admis jusqu'ici, permet de penser que l'activité magmatique pan-africaine/brésilienne dans le centre du Brésil a été plus intense que ce qu'on croyait jusqu'ici. Вмещающие породы зал ежи меди Chapada в Goiбs представляют собой а мфиболитовую фацию в ысшей степени метаморфизм а. Цельная порода и отд ельные минералы дают для кри сталлизации вулкани тов и магматитов до метаморфизма нача льное значение соотн ошения стронция в 0,70414±0,00005, что раз решает принять их возраст в 561± 9 Мио лет, а возраст мет аморфизма в 532±1 Мио, т. е. отнести его к периоду панафриканского -бра зильского орогенеза / 700 до 450 Мио /. При определении в озраста минералов из диоритных тел порфиритовых тек стур, интрудированны х в сланец, получили дат ы в 534±16 Мио лет / 1 = 0,70415 = 0,00004 /. Такое низкое значе ние исходных величин соотношения изотопов разрешает п редполагать, что материал для Chapada при несен был из мантии, или глубинных регион ов коры. Эти исходные з начения характерны для магма тической деятельнос ти типа островных дуг. Во зраст вмещающих поро д считают средне-протерозойск им; это стоит в противо речии с новейшими данными, п о которым он может быт ь только нижнекембрий ским. Это также указыв ает на то, что фаза пан-африка нско-бразильской маг матической активности в централ ьной Бразилии протек ала, возможно, гораздо инт енсивнее, чем принима ли это до сих пор.Peer Reviewedhttp://deepblue.lib.umich.edu/bitstream/2027.42/47849/1/531_2005_Article_BF01830182.pd