The beetle tree of life reveals that Coleoptera survived end‐Permian mass extinction to diversify during the Cretaceous terrestrial revolution

Here we present a phylogeny of beetles (Insecta: Coleoptera) based on DNA sequence data from eight nuclear genes, including six single‐copy nuclear protein‐coding genes, for 367 species representing 172 of 183 extant families. Our results refine existing knowledge of relationships among major groups of beetles. Strepsiptera was confirmed as sister to Coleoptera and each of the suborders of Coleoptera was recovered as monophyletic. Interrelationships among the suborders, namely Polyphaga (Adephaga (Archostemata, Myxophaga)), in our study differ from previous studies. Adephaga comprised two clades corresponding to Hydradephaga and Geadephaga. The series and superfamilies of Polyphaga were mostly monophyletic. The traditional Cucujoidea were recovered in three distantly related clades. Lymexyloidea was recovered within Tenebrionoidea. Several of the series and superfamilies of Polyphaga received moderate to maximal clade support in most analyses, for example Buprestoidea, Chrysomeloidea, Coccinelloidea, Cucujiformia, Curculionoidea, Dascilloidea, Elateroidea, Histeroidea and Hydrophiloidea. However, many of the relationships within Polyphaga lacked compatible resolution under maximum‐likelihood and Bayesian inference, and/or lacked consistently strong nodal support. Overall, we recovered slightly younger estimated divergence times than previous studies for most groups of beetles. The ordinal split between Coleoptera and Strepsiptera was estimated to have occurred in the Early Permian. Crown Coleoptera appeared in the Late Permian, and only one or two lineages survived the end‐Permian mass extinction, with stem group representatives of all four suborders appearing by the end of the Triassic. The basal split in Polyphaga was estimated to have occurred in the Triassic, with the stem groups of most series and superfamilies originating during the Triassic or Jurassic. Most extant families of beetles were estimated to have Cretaceous origins. Overall, Coleoptera experienced an increase in diversification rate compared to the rest of Neuropteroidea. Furthermore, 10 family‐level clades, all in suborder Polyphaga, were identified as having experienced significant increases in diversification rate. These include most beetle species with phytophagous habits, but also several groups not typically or primarily associated with plants. Most of these groups originated in the Cretaceous, which is also when a majority of the most species‐rich beetle families first appeared. An additional 12 clades showed evidence for significant decreases in diversification rate. These clades are species‐poor in the Modern fauna, but collectively exhibit diverse trophic habits. The apparent success of beetles, as measured by species numbers, may result from their associations with widespread and diverse substrates – especially plants, but also including fungi, wood and leaf litter – but what facilitated these associations in the first place or has allowed these associations to flourish likely varies within and between lineages. Our results provide a uniquely well‐resolved temporal and phylogenetic framework for studying patterns of innovation and diversification in Coleoptera, and a foundation for further sampling and resolution of the beetle tree of life.     

Colloidal crystal-like structure of sporopollenin in the megaspore walls of Recent Selaginella and similar fossil spores

HEMSLEY, A. R., COLLINSON, M. E. & BRAIN, A. P. R., 1992. Colloidal crystal-like structure of sporopollenin in the megaspore walls of Recent Selaginella and similar fossil spores. TEM and SEM studies of iridescent fossil Erlansonisporites and Recent Selaginella spores reveal a wall composed of three regions. The central region consists of close-packed particles in a semi-crystalline arrangement. This organization is compared with that of aggregated Iridoviridae, prolamellar bodies, lipid micelles and precious opal. Our observations suggest that the structure of this wall region (and that of the remaining regions) can be explained by derivation from colloidal mixtures. It is concluded that colloids and their precursors may play a far more significant part in spore and pollen wall structure than was previously believed.     

The Inductive Role of Bacterial Symbionts in the Morphogenesis of a Squid Light Organ

SYNOPSIS. The association of the sepiolid squid Euprymna scolopeswith its marine luminous bacterial symbiont Vibrio fischeriis an emerging model system to study the initiation and developmentof bacterial symbioses in higher animals, in particular theinfluence of bacteria on the ontogenic development of symbiotic-specifichost tissues. Experiments comparing the development of juvenilesquid infected with symbiotic V. fischeri with that of uninfectedjuveniles suggest postembryonic development of the light organrequires cell-cell interactions with the bacterial symbionts.The presence of symbiotic bacteria induces specific morphologicalchanges by affecting such fundamental processes as cell deathand cell differentiation. The surface of the juvenile organis largely composed of ciliated cells that appear to facilitateinfection of the light organ. These cells begin to undergo celldeath within hours of infection with symbiotic V. fischeri.Within three days the epithelial cells that form the bacteriacontainingcrypts of the light organ increase in size; these cells do notappear mitotically active, and may represent a terminally differentiatedstate. The light organs of uninfected juvenile E. scolopes,however, do not exhibit any of these early postembryonic developmentalevents but remain in a state of arrested morphogenesis.     

Studies on the Glasshouse Carnation: Effects of Light and Temperature on the Growth and Development of the Flower

Experiments were concerned with the growth and development offlowers of carnation (Dianthus) var. White Sim from the timethe flower buds became visible up to anthesis. Rates of growthin size of the flower were decreased by either low temperaturesor low light intensities but only low temperatures caused anappreciable delay in anthesis. Effects of low light intensitycould be simulated by partial defoliation and appeared to bemediated through effects on photosynthesis. Temperature, however,acted directly on processes occurring in the flower bud. The rate of development of the flower, in the sense of progresstowards anthesis, appeared to be independent of levels of assimilatesavailable for growth. It is suggested that processes controllingdevelopment in the flower regulate the partition of assimilatesbetween the flower and the remainder of the shoot system. Seasonal variations in rates of flower development under glasshouseconditions are considered to be largely attributable to variationsin the temperature of the flower buds.     

Effects of Temperature Variation at Different Times on Growth and Yield of Sugar Beet and Barley

Sugar-beet and barley were grown in pots outdoors (environmentN) and, for five successive 4-week periods starting at sowing,batches of plants were transferred to three growth rooms whosetemperatures were either similar to the outdoor mean (environment^M), or 3° C hotter (environment H) or 3° C colder (environmentC). Some plants were harvested immediately after treatment;others were returned to environment N and harvested when mature. At the end of period 1, sugar-beet plants from environment Mhad more dry weight and leaf area than those outdoors. Immediatelyafter spending later periods in environment M, plants had smallerleaves and similar dry weight to those continuously outdoors.These differences disappeared by maturity. Warmth in the growthrooms (i.e. the difference H—C) during periods 1, 2, and3, while leaf area was increasing, increased the number andsize of leaves and usually also dry weight; in later periodsit had no effect. The effects induced during periods 2 and 3,but not period 1, persisted to maturity to give greater totaland root dry weight and yield of sugar. The final effects ondry weight were much larger than those immediately after treatment,and were the result of differences in growth outdoors aftertreatment which depended on differences in leaf area; the efficiencyof the leaves was not affected by previous treatment. Transferring barley to environment M from N had inconsistentimmediate effects on leaf area and dry weight which disappearedby the final harvest. Transfer during periods 2 and 3, beforethe ears had started emerging, increased shoot number and delayeddevelopment. The proportion of the ears that ripened and theyield of grain were usually less for plants that had spent aperiod in environment M than for plants permanently outdoors,which also had some green ears. Warmth in the growth rooms duringperiods 1 and 2 increased dry weight and leaf area immediately,but had negligible effects at maturity because the increasesin leaf area did not persist after ear emergence. Warmth laterhastened death of leaves, decreased total dry weight immediatelyand also at maturity, but increased the proportion of ears thatripened and hence usually grain weight. Variation in leaf areaduration after ear emergence (D), determined by effects on thetime the ears emerged and the rate the leaves died, accountedfor most of the variation in grain yield, but warmth after theears emerged decreased grain yield less than proportionallyto the decrease in D. Net assimilation rate (E) of sugar-beet was greater than ofbarley, and decreased less with age. E of both species was usuallygreater in environment M than outdoors in spite of less radiation.It was only slightly affected by temperature. Nitrogen and potassium uptake were increased by treatments thatincreased dry weight. The percentage contents suggest that extrauptake was a consequence and not a cause of the increase indry weight.