Andrew J. Ross

Purbeck–Wealden (early Cretaceous) climates

A multidisciplinary colligation including new data and analysis of the evidence for the climates of southern Britain during c. 140 Ma. to c. 120 Ma BP (Berriasian-Barremian — ? earliest Aptian). The climate was at first hot, semi-arid and ‘Mediterranean’ (rather than ‘monsoonal’) in type, probably with seasonally opposed winds (E/W). An irregular long-term trend of increasing rainfall in the moister seasons is evident. This was probably associated with establishment of predominant westerlies during the Jurassic-Cretaceous transition and slightly lower average annual temperatures thereafter until Barremian times. Causes proposed are frequent changes in the regional climatic system due to technically induced adjustments of relief under the special conditions of the semi-enclosed Purbeck–Wealden archipelago and increasing proximity of the widening Protoatlantic sea.

Insect origination and extinction in the Phanerozoic

Abstract Insects (Superclass Hexapoda) are the most successful group of living terrestrial arthropods and the richness of their fossil record is only just beginning to be realized with the recent publication of two extensive databases. Hexapods first appeared in the Early Devonian, post-dating the Cambrian ‘explosion’ of marine arthropods by some 140 Ma. The earliest Hexapoda belong to primitively wingless taxa; however, these ‘Apterygota’ comprise less than 1% of all hexapod species. The appearance of winged insects (Pterygota) in the mid-Carboniferous was accompanied by a major adaptive radiation of non-holometabolous hexapods at ordinal level This was supplemented in the Permian by the radiation of insects with complete metamorphosis (Holometabola). The number of insect orders present in this period was similar to that at the present day. The family data suggest four major periods of origination in the Phanerozoic, with peaks in the Permo-Carboniferous, Early Jurassic, Early Cretaceous and Eocene. Unlike the Tertiary, the Palaeozoic and Cretaceous peaks are accompanied by considerable turnover of families; they are followed by reduced palaeodiversity in the Early Triassic and Late Cretaceous and Palaeocene. The former decline may be linked with the general extinction at the Permian/Triassic boundary and the latter with the rise of the angiospermous flowering plants. Insect generic data for the Phanerozoic reflect the pattern shown by families but not orders. In general, insect diversity may be explained by an overall trend towards low extinction and steady origination at a subordinal level since the Palaeozoic.

Fossil evidence for key innovations in the evolution of insect diversity

Explaining the taxonomic richness of the insects, comprising over half of all described species, is a major challenge in evolutionary biology. Previously, several evolutionary novelties (key innovations) have been posited to contribute to that richness, including the insect bauplan, wings, wing folding and complete metamorphosis, but evidence over their relative importance and modes of action is sparse and equivocal. Here, a new dataset on the first and last occurrences of fossil hexapod (insects and close relatives) families is used to show that basal families of winged insects (Palaeoptera, e.g. dragonflies) show higher origination and extinction rates in the fossil record than basal wingless groups (Apterygota, e.g. silverfish). Origination and extinction rates were maintained at levels similar to Palaeoptera in the more derived Polyneoptera (e.g. cockroaches) and Paraneoptera (e.g. true bugs), but extinction rates subsequently reduced in the very rich group of insects with complete metamorphosis (Holometabola, e.g. beetles). Holometabola show evidence of a recent slow-down in their high net diversification rate, whereas other winged taxa continue to diversify at constant but low rates. These data suggest that wings and complete metamorphosis have had the most effect on family-level insect macroevolution, and point to specific mechanisms by which they have influenced insect diversity through time.

Tomographic reconstruction of neopterous Carboniferous insect nymphs

Two new polyneopteran insect nymphs from the Montceau-les-Mines Lagerstätte of France are presented. Both are preserved in three dimensions, and are imaged with the aid of X-ray micro-tomography, allowing their morphology to be recovered in unprecedented detail. One–Anebos phrixos gen. et sp. nov.–is of uncertain affinities, and preserves portions of the antennae and eyes, coupled with a heavily spined habitus. The other is a roachoid with long antennae and chewing mouthparts very similar in form to the most generalized mandibulate mouthparts of extant orthopteroid insects. Computer reconstructions reveal limbs in both specimens, allowing identification of the segments and annulation in the tarsus, while poorly developed thoracic wing pads suggest both are young instars. This work describes the morphologically best-known Palaeozoic insect nymphs, allowing a better understanding of the juveniles’ palaeobiology and palaeoecology. We also consider the validity of evidence from Palaeozoic juvenile insects in wing origin theories. The study of juvenile Palaeozoic insects is currently a neglected field, yet these fossils provide direct evidence on the evolution of insect development. It is hoped this study will stimulate a renewed interest in such work.

Changes to the Fossil Record of Insects through Fifteen Years of Discovery

The first and last occurrences of hexapod families in the fossil record are compiled from publications up to end-2009. The major features of these data are compared with those of previous datasets (1993 and 1994). About a third of families (>400) are new to the fossil record since 1994, over half of the earlier, existing families have experienced changes in their known stratigraphic range and only about ten percent have unchanged ranges. Despite these significant additions to knowledge, the broad pattern of described richness through time remains similar, with described richness increasing steadily through geological history and a shift in dominant taxa, from Palaeoptera and Polyneoptera to Paraneoptera and Holometabola, after the Palaeozoic. However, after detrending, described richness is not well correlated with the earlier datasets, indicating significant changes in shorter-term patterns. There is reduced Palaeozoic richness, peaking at a different time, and a less pronounced Permian decline. A pronounced Triassic peak and decline is shown, and the plateau from the mid Early Cretaceous to the end of the period remains, albeit at substantially higher richness compared to earlier datasets. Origination and extinction rates are broadly similar to before, with a broad decline in both through time but episodic peaks, including end-Permian turnover. Origination more consistently exceeds extinction compared to previous datasets and exceptions are mainly in the Palaeozoic. These changes suggest that some inferences about causal mechanisms in insect macroevolution are likely to differ as well.

Phylogenetic and environmental context of a Tournaisian tetrapod fauna

The end-Devonian to mid-Mississippian time interval has long been known for its depauperate palaeontological record, especially for tetrapods. This interval encapsulates the time of increasing terrestriality among tetrapods, but only two Tournaisian localities previously produced tetrapod fossils. Here we describe five new Tournaisian tetrapods (Perittodus apsconditus, Koilops herma, Ossirarus kierani, Diploradus austiumensis and Aytonerpeton microps) from two localities in their environmental context. A phylogenetic analysis retrieved three taxa as stem tetrapods, interspersed among Devonian and Carboniferous forms, and two as stem amphibians, suggesting a deep split among crown tetrapods. We also illustrate new tetrapod specimens from these and additional localities in the Scottish Borders region. The new taxa and specimens suggest that tetrapod diversification was well established by the Tournaisian. Sedimentary evidence indicates that the tetrapod fossils are usually associated with sandy siltstones overlying wetland palaeosols. Tetrapods were probably living on vegetated surfaces that were subsequently flooded. We show that atmospheric oxygen levels were stable across the Devonian/Carboniferous boundary, and did not inhibit the evolution of terrestriality. This wealth of tetrapods from Tournaisian localities highlights the potential for discoveries elsewhere.

Raphidiomimula , AN ENIGMATIC NEW COCKROACH IN CRETACEOUS AMBER FROM MYANMAR (BURMA) (INSECTA: BLATTODEA: RAPHIDIOMIMIDAE)

Synopsis An intriguing new genusand species of cockroach (Blattodea), Raphidiomimula burmitica, is described in Cretaceous amber from Myanmar. Raphidiomimidae previously were known only as compression fossils from the Upper Jurassic (Kimmeridgian) of Karatau, Kazakhstan. The structure of the cockroaches, including the apparently raptorial forelegs of Raphidiomimula, suggests they were predatory. The new fossil has several features that are apomorphic and others that are plesiomorphic with respect to the two previously known genera, Raphidiomima and Cameloblatta. The relationships of Raphidiomimidae to other Blattodea and Dictyoptera remain obscure.

The Strawberry Bank Lagerstätte reveals insights into Early Jurassic life

The Strawberry Bank Lagerstätte provides a rich insight into Early Jurassic marine vertebrate life, revealing exquisite anatomical detail of marine reptiles and large pachycormid fishes thanks to exceptional preservation, and especially the uncrushed, 3D nature of the fossils. The site documents a fauna of Early Jurassic nektonic marine animals (five species of fishes, one species of marine crocodilian, two species of ichthyosaurs, cephalopods and crustaceans), but also over 20 species of insects. Unlike other fossil sites of similar age, the 3D preservation at Strawberry Bank provides unique evidence on palatal and braincase structures in the fishes and reptiles. The age of the site is important, documenting a marine ecosystem during recovery from the end-Triassic mass extinction, but also exactly coincident with the height of the Toarcian Oceanic Anoxic Event, a further time of turmoil in evolution.

Corydasialidae fam. n. (Megaloptera) from Baltic amber

A new family of the order Megaloptera from Baltic amber is suggested: Corydasialidae fam. n. The type species is Corydasialis inexspectatus gen. & sp. n. The new family is characterized by some autapomorphic traits: 4 th tarsal segments of the legs are bilobed, underside of the 4 th tarsal lobes with a brush of sucker disc-shaped hairs, in forewings Sc and R1 are not fused and both CuA and CuP distinctly forked.

REDESCRIPTION OF SIMULIDIUM PRISCUM WESTWOOD AND PSEUDOSIMULIUM HUMIDUM (BRODIE) (INSECTA: DIPTERA: RHAGIONIDAE) FROM THE PURBECK LIMESTONE GROUP (LOWER CRETACEOUS) OF ENGLAND

Synopsis The holotype of Simulidium priscum Westwood, 1854 has been rediscovered in the collections of The Natural History Museum, London. It is a single wing from the Lulworth Formation (Berriasian) of Durlston Bay, Dorset. S. priscum and Pseudosimulium humidum (Brodie 1845) from the Lulworth Formation of Wiltshire are redescribed and placed within the Rhagionidae.