Since Haeckel's Biogenetic Law (‘ontogeny recapitulates phylogeny’) fell into disrepute early in the twentieth century, there has been intermittent debate, particularly in recent years (de Beer, 1958; Gould, 1977; Alberch et al., 1979; Alberch, 1980; Bonner, 1982; McNamara, 1982a), on the nature of the relationship between an individual's development and phylogenetic history. Important questions under discussion include the following: If a strong causal relationship does exist, what is its nature? How does it work? What is its importance in evolution? How can it be recognized in the fossil record?

The trace fossils of the Tumblagooda Sandstone (?late Silurian) of Kalbarri, Western Australia are spectacular in their variety and preservation. They provide a unique insight into the activities of the early invaders of terrestrial environments, and reveal the presence of a diverse fauna dominated by arthropods. Within the Formation trace fossil assemblages can be related to fluvial, aeolian and marine sand-dominated environments. Two distinct and diverse ichnofaunas are recognised.The Heimdallia–Diplichnites Ichnofauna occurs in sandstones deposited in broad low sinuosity braided fluvial channels, between which were mixed aeolian and waterlain sandsheets, small aeolian dunes and flooded interdune and deflation hollows. Heimdallia is the major bioturbator, favouring shallow pools. Other burrows include Tumblagoodichnus (gen. nov.), Diplocraterion, Skolithos, Beaconites and Didymaulyponomos. Arthropod trackways (Diplichnites) occur on surfaces of waterlain sands and on foreset bedding of aeolian dunes, and represent some of the earliest reported terrestrial trackways. Other trackways include Paleohelcura and Protichnites, and the digging traces Selenichnites and Rusophycus are also present. At least ten types of arthropods are required to produce the observed traces. Myriapods, eurypterids, euthycarcinoids, xiphosurids and scorpionids are considered responsible for the trackway assemblage.The Skolithos–Diplocraterion Ichnofauna occurs at the top of the exposed section in sandstones that overlie a thick fluvial sequence containing few traces. The strata are considered to represent marine influence at a fluvial/marine transition. They show variable trough cross-bedding, complex planar cross-bedding with down-climbing sets, ripple lamination, and fining-up sequences with bioturbated tops. Traces are dominated by crowded Skolithos up to 1 m long, together with two forms of Diplocraterion. Daedalus and Lunatubichnus (gen. nov.) burrows occur in a few beds and Aulichnites trails cover some foreset surfaces of cross-bedding.The trace fossils and the sedimentology of the Tumblagooda Sandstone bear a remarkable similarity to those of the lower part of the Taylor Group of Antarctica, which is probably Devonian in age. It is suggested that the two represent a similar age, stratigraphy, and range of environments on the margins of Gondwana. Large unvegetated fluvial outwash plains with variable aeolian influence were essentially coastal in character and fluvial/marine transitions occur in sand-rich environments. The animals responsible for the traces inhabited coastal areas but many could survive outwith marine influence, and arthropods responsible for some types of Diplichnites trackways walked out of water.The rich diversity of trackways attributable to arthropods illustrate that the invasion of terrestrial environments by arthropods, particularly large forms, was well-established by the beginning of the Devonian. The basis of the food chain was algal and bacterial films which bound the surface sediment in freshwater pools.

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Heterochrony can be defined as change to the timing or rate of development relative to the ancestor. Because organisms generally change in shape as well as increase in size during their development, any variation to the duration of growth or to the rate of growth of different parts of the organism can cause morphological changes in the descendant form. Heterochrony takes the form of both increased and decreased degrees of development, known as "peramorphosis" and "paedomorphosis," respectively. These are the morphological consequences of the operation of processes that change the duration of the period of an individual's growth, either starting or stopping it earlier or later than in the ancestor, or by extending or contracting the period of growth. Heterochrony operates both intra-and interspecifically and is the source of much intraspecific variation. It is often also the cause of sexual dimorphism. Selection of a sequence of species with a specific heterochronic trait can produce evolutionary trends in the form of pera-or paedomorphoclines. Many different life history traits arise from the operation of heterochronic processes, and these may sometimes be the targets of selection rather than morphological features themselves. It has been suggested that some significant steps in evolution, such as the evolution of vertebrates, were engendered by heterochrony. Human evolution was fuelled by heterochrony, with some traits, such as a large brain, being peramorphic, whereas others, such as reduced jaw size, are paedomorphic.

Summary 1. Renewed interest in the role of changes to developmental regulation in organisms has highlighted the importance of heterochrony in the evolution of the Metazoa. 2. Beecher's interpretation of the evolution of the Trilobita as having been principally by peramorphosis is examined, as is the view of later workers, principally Stubblefield and Hupé, that paedomorphosis was a dominant factor in trilobite evolution. 3. Both peramorphosis and paedomorphosis are considered to have been important in trilobite evolution. 4. The role of paedomorphosis in the evolution of major morphological novelties is critically examined. Its importance in changes to the structure of the glabella is discussed and new terms proposed to describe the ontogenetic and phylogenetic state of the glabella. 5. The highly variable nature of early Cambrian trilobites, in particular the large degree of ontogenetic change, is considered, along with possible poor developmental control of the growth and moulting hormonal systems, to have been significant in providing a high degree of intrapopulational morphological variability. Selection of these heterochronic variants was responsible for the rapid diversification of the Trilobita during the Cambrian.