The Linear Angiosperm Phylogeny Group (LAPG) III: a linear sequence of the families in APG III

The publication of the third Angiosperm Phylogeny Group (APG) classification (APG III. 2009. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161: 128–131) has resulted in the need for a revised systematic listing of the accepted families. This linear APG III (LAPG III) sequence of families is presented here. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 161 , 128–131.     

Patterns of self compatibility, inbreeding depression, outcrossing, and sex allocation in a marine bryozoan suggest the predominating influence of sperm competition

Sex allocation by simultaneous hermaphrodites is theoretically influenced by selfing rate, which is in turn influenced by the benefits of enhanced genomic transmission and reproductive assurance relative to the cost of inbreeding depression. The experimental investigation of these influences in seed plants has a rich pedigree, yet although such an approach is equally relevant to colonial invertebrates, which globally dominate subtidal communities on firm substrata, such studies have been scarce. We reared self‐compatible genets of the marine bryozoan Celleporella hyalina s.l. in the presence and absence of allosperm, and used molecular genetic markers for paternity analysis of progeny to test theoretical predictions that: (1) genets from focal populations with high selfing rates show less inbreeding depression than from focal populations with low selfing rates; (2) genets whose selfed progeny show inbreeding depression prefer outcross sperm (allosperm); and (3) genets bias sex allocation toward female function when reared in reproductive isolation. Offspring survivorship and paternity analysis were used to estimate levels of inbreeding depression and preference for outcrossing or selfing. Sex allocation was assessed by counting male and female zooids. As predicted, inbreeding depression was severe in selfed progeny of genets derived from the populations with low self‐compatibility rates, but, with one exception, was not detected in selfed progeny of genets derived from the populations with higher self‐compatibility rates. Also, as predicted, genets whose selfed progeny showed inbreeding depression preferred outcrossing, and a genet whose selfed progeny did not show inbreeding depression preferred selfing. Contrary to prediction, sex allocation in the majority of genets was not influenced by reproductive isolation. Lack of economy of male function may reflect the over‐riding influence of allosperm‐competition in typically dense breeding populations offering good opportunity for outcrossing. We suggest that hermaphroditism may be a plesiomorphic character of the crown group Bryozoa, prevented by phylogenetic constraint from being replaced by gonochorism and therefore not necessarily adaptive in all extant clades. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 519–531.     

Annual growth of Cassiope tetragona as a proxy for Arctic climate: developing correlative and experimental transfer functions to reconstruct past summer temperature on a millennial time scale

Annual growth of the polar evergreen shrub Cassiope tetragona on Svalbard was evaluated as a proxy for Arctic summer temperatures. Transfer functions were derived from temperature‐growth correlations of shoots and from a temperature‐growth response, obtained from experimental warming using open top chambers (OTC) in high Arctic tundra vegetation at Isdammen approximately 1.5 km southeast of Longyearbyen, Svalbard (78°N, 15 E) and in Longyeardalen, 3 km west of Isdammen from 2004 to 2006. Air temperatures, monitored throughout the summer months, were 1.3 °C higher inside the OTCs than in the control plots. Annual stem growth was measured by tagging stems and leaves, and in the lab with shoots harvested from OTCs and control plots. Annual growth parameters assessed were leaf production, sum of length and weight of individual leaves, and stem length increment derived from leaf scar distances and the distances between wintermarksepta in the stem. Wintermarksepta are formed at the end of the summer growth period when the pith is narrowing and consist of dense and dark tissue ( Fig. 1b ). The variation of annual growth in a 34‐year site chronology (based on Cassiope shoots from the surroundings of the OTCs and control plots) correlated strongly with the mean summer temperature on Svalbard. The number of leaf pairs, leaf length and stem length also increased in the OTC warmed plots in the second and third year of warming. Transfer functions were derived from the temperature‐annual growth correlations from a single shoot from Longyeardalen, from the cross‐dated Isdammen site chronology and from the growth response to experimental warming. Based on leaf scar distances and distances between wintermarksepta of well‐preserved subfossil shoots in arctic tundra soil, annual stem length increase was assessed for the layers of a soil core collected at the Isdammen site. Based on the derived transfer functions summer temperature of the period relating to the 15 cm deep tundra soil core layer, radiocarbon dated at 4230±40 bp , may have been 3.0 °C lower than the present‐day 6.2 °C value. These results indicate that the transfer functions can be used to reconstruct past temperatures, beyond the time range of instrumental temperature and ice core records of Svalbard.

Figure 1 Open in figure viewer PowerPoint (a) Morphology of shoot of Cassiope tetragona collected May 26, 2007, Longyeardalen. Removal of the front row of leaves shows four leaf pairs of the 2006 summer, the leaf primordium for the 2007 growing season and the stem length increase summer 2006. (b) Wintermarksepta, darker colored than the pith tissue, indicating the winterperiod in a longitudinal section of an air‐dried shoot collected August 26, 2006, Longyeardalen.     

Seeing the forest for the trees: long-term exposure to elevated CO2 increases some herbivore densities

The effects of elevated CO₂ on plant growth and insect herbivory have been frequently investigated over the past 20 years. Most studies have shown an increase in plant growth, a decrease in plant nitrogen concentration, an increase in plant secondary metabolites and a decrease in herbivory. However, such studies have generally overlooked the fact that increases in plant production could cause increases of herbivores per unit area of habitat. Our study investigated leaf production, herbivory levels and herbivore abundance per unit area of leaf litter in a scrub‐oak system at Kennedy Space Center, Florida, under conditions of ambient and elevated CO₂, over an 11‐year period, from 1996 to 2007. In every year, herbivory, that is leafminer and leaftier abundance per 200 leaves, was lower under elevated CO₂ than ambient CO₂ for each of three species of oaks, Quercus myrtifolia, Quercus chapmanii and Quercus geminata. However, leaf litter production per 0.1143 m² was greater under elevated CO₂ than ambient CO₂ for Q. myrtifolia and Q. chapmanii, and this difference increased over the 11 years of the study. Leaf production of Q. geminata under elevated CO₂ did not increase. Leafminer densities per 0.1143 m² of litterfall for Q. myrtifolia and Q. chapmanii were initially lower under elevated CO₂. However, shortly after canopy closure in 2001, leafminer densities per 0.1143 m² of litter fall became higher under elevated CO₂ and remained higher for the remainder of the experiment. Leaftier densities per 0.1143 m² were also higher under elevated CO₂ for Q. myrtifolia and Q. chapmanii over the last 6 years of the experiment. There were no differences in leafminer or leaftier densities per 0.1143 m² of litter for Q. geminata. These results show three phenomena. First, they show that elevated CO₂ decreases herbivory on all oak species in the Florida scrub‐oak system. Second, despite lower numbers of herbivores per 200 leaves in elevated CO₂, increased leaf production resulted in higher herbivore densities per unit area of leaf litter for two oak species. Third, they corroborate other studies which suggest that the effects of elevated CO₂ on herbivores are species specific, meaning they depend on the particular plant species involved. Two oak species showed increases in leaf production and herbivore densities per 0.1143 m² in elevated CO₂ over time while another oak species did not. Our results point to a future world of elevated CO₂ where, despite lower plant herbivory, some insect herbivores may become more common.     

Fire frequency influences composition and structure of the shrub layer in an Australian subcoastal temperate grassy woodland

Little is known about the relationship between fire regimes and plant diversity in Australia's temperate grassy woodlands. The effect of fire frequency on shrubs in grassy woodland remnants across Western Sydney's Cumberland Plain was examined. Shrub species richness and composition were compared in sites that had experienced a high, moderate or low frequency of fire over the previous 20 years. Nine sites were surveyed, three in each fire frequency category; most sites, including all low‐fire‐frequency sites, had burnt 9–36 months prior to sampling. Fire frequency had a profound effect on the composition and structure of the shrub layer. Per cent frequency and density of the prickly shrub Bursaria spinosa (Pittosporaceae) was considerably higher in low‐fire‐frequency sites than where fires had occurred at least once a decade. In sites where fire had been absent for decades prior to a recent fire, this species dominated the landscape, while elsewhere it occurred as clumps in a grassy matrix. Per cent frequency of other native shrubs, particularly obligate seeders, was greatest at moderate fire frequencies. Exotic shrubs were recorded most often where fire had been rare. While ordination clearly separated out the low‐fire‐frequency sites, complete separation between high‐ and moderate‐fire‐frequency blocks was not achieved. The increase in Bursaria in the absence of fire mirrors the encroachment of woody plants into a range of grassy ecosystems around the world. The sensitivity of obligate seeder species, many of them short‐lived legumes with fire‐cued seeds, to both very frequent and very infrequent fire shows the vulnerability of these species to extreme fire regimes, despite the safeguards conferred by hard‐seededness. Competition from Bursaria, as well as loss of viable seed in the soil, may have contributed to the low frequency of these species after a long inter‐fire interval.     

Ontogenesis of Dileptus terrenus and Pseudomonilicaryon brachyproboscis (Ciliophora, Haptoria)

ABSTRACT. Dileptids are haptorid ciliates with a conspicuous proboscis belonging to the oral apparatus and carrying a complex, unique ciliary pattern. We studied development of body shape, ciliary pattern, and nuclear apparatus during and after binary fission of Dileptus terrenus using protargol impregnation. Additional data were obtained from a related species, Pseudomonilicaryon brachyproboscis . Division is homothetogenic and occurs in freely motile condition. The macronucleus is homomeric and condenses to a globular mass in mid-dividers. The proboscis appears in late mid-dividers as a small convexity in the opisthe's dorsal brush area and maturates post-divisionally. The oral and dorsal brush structures develop by three rounds of basal body proliferation. The first round generates minute anarchic fields that will become circumoral kinetofragments, while the second round produces the perioral kinety on the right and the preoral kineties on the dorsal opisthe's side. The dorsal brush is formed later by a third round of basal body production. The formation of various Spathidium -like body shapes and ciliary patterns during ontogenesis and conjugation of Dileptus shows a close relationship between spathidiids and dileptids. On the other hand, the peculiarities of the dileptid morphology and ontogenesis indicate a long, independent evolution.     

El Niño,grazers and fisheries interact to greatly elevate extinction risk for Galapagos marine species

Comparisons between historical and recent ecological datasets indicate that shallow reef habitats across the central Galapagos Archipelago underwent major transformation at the time of the severe 1982/1983 El Niño warming event. Heavily grazed reefs with crustose coralline algae (‘urchin barrens’) replaced former macroalgal and coral habitats, resulting in large local and regional declines in biodiversity. Following recent threat assessment workshops, a total of five mammals, six birds, five reptiles, six fishes, one echinoderm, seven corals, six brown algae and nine red algae reported from coastal environments in Galapagos are now recognized as globally threatened. The 2008 International Union for the Conservation of Nature (IUCN) Red List includes 43 of these species, while two additional species (Galapagos damsel Azurina eupalama and 24‐rayed sunstar Heliaster solaris) not seen for > 25 years also fulfil IUCN threatened species criteria. Two endemic species (Galapagos stringweed Bifurcaria galapagensis and the damselfish A. eupalama) are now regarded as probably extinct, while an additional six macroalgal species (Dictyota galapagensis, Spatoglossum schmittii, Desmarestia tropica, Phycodrina elegans, Gracilaria skottsbergii and Galaxaura barbata) and the seastar H. solaris are possibly extinct. The removal of large lobster and fish predators by artisanal fishing probably magnified impacts of the 1982/1983 El Niño through a cascade of indirect effects involving population expansion of grazing sea urchins. Marine protected areas with adequate enforcement are predicted to ameliorate but not eliminate ecosystem impacts caused by increasing thermal anomalies associated with El Niño and global climate change.     

Resilience of a high‐conservation‐value,semi‐arid grassland on fertile clay soils to burning,mowing and ploughing

In grassland reserves, managed disturbance is often necessary to maintain plant species diversity. We carried out experiments to determine the impact of fire, kangaroo grazing, mowing and disc ploughing on grassland species richness and composition in a nature reserve in semi‐arid eastern Australia. Vegetation response was influenced by winter–spring drought after establishment of the experiments, but moderate rainfall followed in late summer–autumn. Species composition varied greatly between sampling times, and the variability due to rainfall differences between seasons and years was greater than the effects of fire, kangaroo grazing, mowing or disc ploughing. In the fire experiment, species richness and composition recovered more rapidly after spring than autumn burning. Species richness and composition were similar to control sites within 12 months of burning and mowing, suggesting that removal of the dominant grass canopy is unnecessary to enhance plant diversity. Two fires (separated by 3 years) and post‐fire kangaroo grazing had only minor influence on species richness and composition. Even disc ploughing caused only a small reduction in native richness. The minor impact of ploughing was explained by the small areas that were ploughed, the once‐off nature of the treatment, and the high degree of natural movement and cracking in these shrink‐swell soils. Recovery of the composition and richness of these grasslands was rapid because of the high proportion of perennial species that resprout vegetatively after fire and mowing. There appears to be little conservation benefit from fire, mowing or ploughing ungrazed areas, as we could identify no native plant species dependent on frequent disturbance for persistence in this grassland community. However, the ability of the Astrebla‐ and Dichanthium‐dominated grasslands to recover quickly after disturbance, given favourable seasonal conditions, suggests that they are well adapted to natural disturbances (e.g. droughts, fire, flooding and native grazing).     

Secondary seed dispersal of Erodiophyllum elderi,a patchily distributed short‐lived perennial in the arid lands of Australia

We investigated secondary dispersal of propagules of Erodiophyllum elderi (Asteraceae), a short‐lived perennial plant growing in small patches in the arid lands of southern Australia. In spite of its importance for population dynamics, secondary dispersal is a little understood process. We monitored the dispersal of 2280 large woody capitula (seed heads) released in six E. elderi patches for 9 months. Colour‐coded seed heads were located at night using UV light and their distance and direction from the release point were measured. Over the 9‐month period, more seed heads moved, and those that did, moved further in areas with high herbivore activity. Overall dispersal distance across the ground was limited to less than 30 m. Dispersal patterns were related to the topographical slope at the release site: seed heads moved further, and more dispersed on steeper slopes unless the steep slopes had sandy soil in which case seed heads were buried, caught or there was reduced sheet water flow limiting their dispersal potential. After several months, seed head dispersal virtually ceased as seed heads became stuck in the debris and soil after heavy rains or further dispersal became unlikely when seed heads reached locally low‐lying areas. Secondary dispersal patterns suggest two distinctly different influences associated with the presence of herbivores: the direct movement of seed heads by trampling from sheep (an introduced herbivore) and the indirect effect of a reduced standing biomass from grazing. Reduced vegetation cover allows seed head redistribution via sheet water flow during large rainfall events.