Richard A. Fortey

Disparity as an evolutionary index; a comparison of Cambrian and Recent arthropods

Disparity is a measure of the range or significance of morphology in a given sample of organisms, as opposed to diversity, which is expressed in terms of the number (and sometimes ranking) of taxa. At present there is no agreed definition of disparity, much less any consensus on how to measure it. Two possible categories of metric are considered here, one independent of any hypothesis of relationship (phenetics), the other constrained within an evolutionary framework (cladistics). The Early Cambrian radiation was clearly a period of significant morphologic and taxonomic diversification. However, we question the interpretation of its first generation products as numerous body plans at the highest level. Four phenetic and two cladistic measures have been used to compare disparity among Cambrian arthropods with that in the living fauna. Phenetic methods assessing character-state variability and the amount of morphological attribute space occupied yield similar results for Cambrian and Recent arthropods. Assessments of disparity within a taxonomic framework rely on the identification of particular characters that delineate higher level body plans. This requires a phylogenetic interpretation, a cladistic investigation of hierarchical structure in the data. Both sets of arthropods fall within the same major clades, and within this cladistic framework the amount of character-state evolution in the two groups is comparable. None of these methods identifies markedly greater disparity among the Cambrian compared with the Recent taxa. Although measures of disparity are applied here to a consideration of the Cambrian radiation, the metrics clearly have a much wider potential for estimating macroevolutionary trends independently from existing taxonomic frameworks. Geometric morphometry is ideal for measuring morphological variety at lower taxonomic levels, but it requires the recognition of homologous landmarks in all the forms under comparison, or the identification of entire homologous structures. Conventional phenetics has much wider application as it can operate on data coded as discrete homologous character states (this facility is also a requirement of cladistics), which are a more appropriate basis for comparing disparity in markedly dissimilar forms.

Palaeontological evidence bearing on global Ordovician-Silurian continental reconstructions

Abstract The discreteness or otherwise of major Ordovician and Silurian terranes can be recognised by the shallow-water benthic faunas which lived upon them. Their borders are often indicated by the disposition of progressively shallow- to deep-water assemblages at the terrane edge as well as by structural features. Their positions relative to each other in the Early Palaeozoic can be best indicated by a combination of palaeomagnetic and faunal evaluation: the latter is the topic of this paper. Faunal evaluation is now possible quantitatively as well as quantitatively. Global palaeobiogeography is reviewed for the period as deduced from faunal evidence. There was one supercontinent, Gondwana, which stretched from West Gondwana (todays southern Europe and North Africa) at high latitudes to tropical East Gondwana (Australasia and adjacent areas), with intermediate palaeolatitudes in the Middle East and South America. Around Gondwana, especially to its north, were a large number of peri-Gondwanan terranes, particularly Avalonia, Perunica, parts of Turkey and Arabia and Sibumasu. In addition, there were the substantial independent continents of Laurentia, Baltica, Siberia, Annamia, North China and South China. Analysis of the shallow-water benthos, particularly trilobites and brachiopods, provides distinctive signatures for palaeo-position in most cases. Despite a large faunal turnover particularly corresponding with the latest Ordovician glacial event, the progressive evolution of the ecologies of benthic shelly faunas were also much influenced by changing geographies during the 80-Ma period. In the early Ordovician, oceans were at their widest, enabling Baltica and Laurentia to have different signatures from either East or West Gondwana. Siberia in early Ordovician times had faunal contact with Laurentia and East Gondwana, but in the mid-Ordovician, there were more endemics, and by the late Silurian, it was the only continent of substance in the northern hemisphere (hosting the Tuvaella Fauna). South China has varied faunal links but seems best treated as at the edge of the peri-Gondwanan collage for most of the period. We show how faunas document the early Ordovician rift of Avalonia from West Gondwana and its movement and subsequent collisions, first with Baltica in the end Ordovician and then with Laurentia in the early Silurian. Faunas also support the postulated movement of the Precordillera of South America from Laurentia in the early Ordovician to intermediate- to high-latitude Gondwana in the Silurian. We examine peripheral terranes bordering Iapetus to demonstrate their pre-collision positions. Analysis of some of the many terranes now forming Kazakhstan and adjacent areas in central Asia today reveals that the benthic faunas there have more affinity with Gondwanan and peri-Gondwanan faunas than with Baltica or Siberia, and thereby challenge structural models postulating an Early Palaeozoic Kipchak arc.

A revision of Ordovician series and stages from the historical type area

This paper is a revision of the standard chronostratigraphy of the Ordovician of the historical type area in England and Wales. The revision is a response to the need for more precise definitions of series and stages, especially for practical international correlation. In some cases this has entailed moving horizons for Series bases away from those in the classical sections. The scheme will be used as the standard for a forthcoming revision of the Correlation Chart of the British Ordovician System. The Ordovician is divided into five series: Tremadoc, Arenig, Llanvirn, Caradoc and Ashgill. The Llanvirn is extended to include part of the classical Llandeilo Series, which is included within it as a stage. Where stage subdivisions of standard series have not yet been defined, as in the Tremadoc, they are proposed in this paper. However, Bancroft’s fine stadial divisions of the Caradoc, which have been criticized for their brevity and local utility, are combined into four new stages, which will have wider application. The former Caradoc subdivisions are retained as substages for the purposes of regional correlation.

PROBLEMS OF REPRODUCIBILITY: DOES GEOLOGICALLY ANCIENT DNA SURVIVE IN AMBER-PRESERVED INSECTS?

Apparently ancient DNA has been reported from amber–preserved insects many millions of years old. Rigorous attempts to reproduce these DNA sequences from amber– and copal–preserved bees and flies have failed to detect any authentic ancient insect DNA. Lack of reproducibility suggests that DNA does not survive over millions of years even in amber, the most promising of fossil environments.

Late Ordovician global warming—The Boda event

There is substantial evidence for mid-Ashgillian global warming before the latest Ordovician Hirnantian glaciation, as shown by the movement of previously lower latitude benthic faunas such as trilobites and brachiopods to progressively higher latitudes and by an increase in endemic faunas at low latitudes. This is shown by faunal dynamics on several paleocontinents and endorsed by published evidence of warmer-water sediments from Gondwana and elsewhere. We term this global warming the Boda event.

New uppermost Cambrian U–Pb date from Avalonian Wales and age of the Cambrian–Ordovician boundary

A crystal-rich volcaniclastic sandstone in the lower Peltura scarabaeoidesZone at Ogof-ddu near Criccieth, North Wales, yields a U-Pb zircon age of 491 ± 1 Ma. This late Late Cambrian date indicates a remarkably young age for the Cambrian-Ordovician boundary whose age must be less than 491 Ma. Hence the revised duration of the post-Placentian (trilobite-bearing) Cambrian indicates that local trilobite zonations allow a biostratigraphic resolution comparable to that provided by Ordovician graptolites and Mesozoic ammonites.

A trilobite fauna from the highest Shineton Shales in Shropshire, and the correlation of the latest Tremadoc

A rich fauna dominated by trilobites and calcified chordates has been collected in Shropshire from the Arenaceous Beds, the highest member of the Tremadoc Shineton Shale Formation, and hitherto regarded as poorly fossiliferous. This fauna shows that shelf conditions persisted longer in Shropshire than has been supposed. It is likely that even younger Tremadoc is cut out at the unconformity below the Caradoc in the Shineton Inlier. The correlation of the later Tremadoc is reviewed, and the nomenclature of British Tremadoc biozones is revised. The trilobites described here are a mixture of previously known and new forms. The name Shumardia (Conophrys) salopiensis Callaway is revived for British material traditionally assigned to the Scandinavian species Shumardia pusilla (Sars), from which it is distinct. The new taxa Litagnostus meniscus sp.nov., Apatokephalus sarculum sp.nov. and Skljarella cracens sp.nov. are described. The type species of Asaphellus, A. homfrayi and Leptoplastides, L. salteri , are redescribed, and Geragnostus callavei, Pseudokainella impar , and Parapilekia sp. are recorded. New information on the ontogeny of S. (C.) salopiensis and A. homfrayi is given. Litagnostus and Skljarella are recorded from the British Isles for the first time.

The Pattern of Global Bio-Events During the Ordovician Period

The 70 Ma Ordovician Period is characterized by extensive epeiric seas, paleocontinent dispersal, intervals of intense volcanism and black shale deposition, a greenhouse climate state deteriorating to a brief icehouse state, strong faunal provincialism, and profound changes to the biota including the changeover from the Cambrian Fauna to the Paleozoic Fauna. Although many invertebrate phyla diversify during the Ordovician, precise biostratigraphic and global biogeographic data are provided best by conodonts, trilobites and graptolites. These three groups are used in this chapter to recognize five major bio-events four of which correspond closely to Series boundaries: Basal Tremadoc (BTc), Basal Arenig (B’Ag), Basal Llanvirn (B’Ln), Basal Caradoc (B’Cc) and Upper Ashgill (U’Al). Most of these correspond to significant eustatic events and the latter to the terminal Ordovician glaciation. The first four are each characterized by extinctions but these are overshadowed by a rapid innovation event with a radiation of a more diversified fauna; the U’Al is a severe extinction event, second only to the terminal Permian event in the entire Phanerozoic. Compared to many other Phanerozoic systems, the Ordovician is a period of considerable biologic, climatic and oceanographic complexity within which the balance between the forcing processes that produced the major and minor events is still not well understood.

TRILOBITE SYSTEMATICS: THE LAST 75 YEARS

Abstract The progress achieved in trilobite systematics over the last 75 years is briefly reviewed. Different approaches to phylogenetics have influenced the way trilobites have been classified. Classical evolutionary taxonomy, the stratigraphical approach, and cladistics have all contributed in different ways to the current classification, which has evolved piecemeal, and is still unsatisfactory is some ways. Nonetheless, progress towards a phylogenetic classification has been made, especially as the result of information from ontogenies provided by well-preserved silificified material. Trilobites are a well-defined clade within a larger arachnomorph group. Agnostida have been excluded from Trilobita, but are perhaps best considered as specialised trilobites, at least until limbs of eodiscids are described. The outstanding problems in classification of each trilobite order are reviewed. Most are concerned with the recognition of the appropriate Cambrian sister taxa, and the discovery of the relevant ontogenies. It is very likely that post-Cambrian clades “root” deeply into the Cambrian. The coherence, or otherwise, of Proetida, Asaphida, Corynexochida and the lichid/odontopleurid groups will be resolved by such studies. The problems of paraphyly in Ptychopariida and Redlichiina may prove more obdurate. The temporal brevity of certain Cambrian family ranges may be partly a taxonomic artefact. The possibility of a late Cambrian gap in the record on some clades should be considered.

Wonderful strife: systematics, stem groups, and the phylogenetic signal of the Cambrian radiation

Abstract Goulds Wonderful Life (1989) was a landmark in the investigation of the Cambrian radiation. Gould argued that a number of experimental body plans (“problematica”) had evolved only to become extinct, and that the Cambrian was a time of special fecundity in animal design. He focused attention on the meaning and significance of morphological disparity versus diversity, and provoked attempts to quantify disparity as an evolutionary metric. He used the Burgess Shale as a springboard to emphasize the important role of contingency in evolution, an idea that he reiterated for the next 13 years. These ideas set the agenda for much subsequent research. Since 1989 cladistic analyses have accommodated most of the problematic Cambrian taxa as stem groups of living taxa. Morphological disparity has been shown to be similar in Cambrian times as now. Konservat-Lagerstätten other than the Burgess Shale have yielded important new discoveries, particularly of arthropods and chordates, which have extended the range of recognized major clades still further back in time. The objective definition of a phylum remains controversial and may be impossible: it can be defined in terms of crown or total group, but the former reveals little about the Cambrian radiation. Divergence times of the major groups remain to be resolved, although molecular and fossil dates are coming closer. Although “superphyla” may have diverged deep in the Proterozoic, “explosive” evolution of these clades near the base of the Cambrian remains a possibility. The fossil record remains a critical source of data on the early evolution of multicellular organisms.