Seventeen years of carbon dioxide enrichment of sour orange trees: final results

The long-term responses of trees to elevated CO₂ are especially crucial (1) to mitigating the rate of atmospheric CO₂ increase, (2) to determining the character of future forested natural ecosystems and their spread across the landscape, and (3) to determining the productivity of future agricultural tree crops. Therefore, a long-term CO₂-enrichment experiment on sour orange trees was started in 1987, and the final results after 17 years are reported herein. Four sour orange trees ( Citrus aurantium L.) were grown from seedling stage at 300 μmol mol⁻¹ CO₂ above ambient in open-top, clear-plastic-wall chambers at Phoenix, AZ. Four control trees were similarly grown at ambient CO₂. All trees were supplied ample water and nutrients comparable with a commercial orchard. After a peak 2–4 years into the experiment, there was a productivity plateau at about a 70% enhancement of annual fruit and incremental wood production over the last several years of the experiment. When summed over the duration of the experiment, there was an overall enhancement of 70% of total biomass production. Much of the enhancement came from greater numbers of fruits produced, with no change in fruit size. Thicker trunks and branches and more branches and roots were produced, but the root/shoot ratio was unaffected. Also, there was almost no change in the elemental composition of the biomass produced, perhaps in part due to the minimal responsiveness of root-symbiotic arbuscular mycorrhizal fungi to the treatment.     

Radiating in a river: systematics, molecular genetics and morphological differentiation of viviparous freshwater gastropods endemic to the Kaek River, central Thailand (Cerithioidea, Pachychilidae)

Speciation in the context of adaptive radiation is regarded as a key process in the creation of biodiversity. While several lacustrine species flocks provide ideal models for elucidating the underlying evolutionary mechanisms, riverine radiations are both rarely known and studied. The Kaek River, a third‐order tributary of the Nan River and Chao Praya drainage in central Thailand, harbours an exceptional endemic species assemblage of morphologically distinct, viviparous pachychilid gastropods. Our systematic revision, combining a morphological and molecular genetics approach, reveals the sympatric existence of at least seven species of the genus Brotia that is widespread in rivers of South‐east Asia where usually only two species at the most coexist. At eight locations along a 100‐kilometre stretch of the Kaek River, we found the syntopic occurrence of two to three species that are separated by specific habitat preferences and exhibit trophic specialization in their radula morphology. Phylogenetic analyses (using MP, NJ, ML and Bayesian inference statistics) of partial COI and 16S sequence data of 17 samples from six species occurring sympatrically and parapatrically, respectively, in the Kaek River drainage (plus the type species B. pagodula as outgroup) indicate monophyly of all these endemic species. Brotia solemiana, which also occurs in the Loei River, a tributary of the Mekong drainage system, was found to be sister to all other Kaek River pachychilids. The distinctive morphotypes, proposed here to represent biospecies, do not show high levels of genetic variation consistent with long periods of reproductive isolation. This suggests a relatively recent origin of this intrariverine radiation and rapid morphological divergence in the Kaek River Brotia. Recent diversification combined with ecological separation and trophic specialization parallels conditions found, albeit on a more specious level, in the lacustrine species flock of the closely related pachychilid genus Tylomelania, which is endemic to ancient lakes on the Indonesian island of Sulawesi. We discuss and compare the allopatric and ecological aspects of speciation in this unique riverine radiation and outline a putative historical biogeography of the Kaek River species, employing the most recent geological and palaeohydrological data for Thailand. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 275–311.     

High larval density does not induce a prophylactic immune response in a butterfly

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Phylogeny of the gastropod superfamily Cerithioidea using morphology and molecules

The Cerithioidea is an ecologically important superfamily of basal Caenogastropoda with speciose marine, brackish water, and freshwater lineages primarily in tropical, subtropical, and warm temperate regions of the world. They often represent significant components of the communities where they occur and have given rise to several spectacular endemic radiations in rivers and ancient lakes. Earlier attempts to resolve the phylogenetic history of the group have been based on smaller taxon and character subsets with incongruent results. Here the monophyly and phylogeny of the group is evaluated with expanded morphological and molecular (16S, 28S rRNA) data sets. For morphological analyses, 151 characters (shell, operculum, radula, alimentary tract, kidney, nervous system, reproductive anatomy, and sperm ultrastructure) were scored for 47 cerithioideans (representing 17 families) and nine outgroup taxa. To test monophyly of the Cerithioidea, extended molecular data sets of 16S and 28S sequences for 57 and 44 taxa, respectively, were compiled using new and previously published sources. For combined analyses, a pruned molecular data set was combined with the morphological partition. The morphological data were analysed alone using only parsimony; molecular and simultaneous analyses were performed using both parsimony and Bayesian inference. The effect of excluding unconserved regions of the alignments was also explored. All analyses, with the exception of the individual 16S and 28S data sets, support monophyly of the Cerithioidea as currently formulated. Of the 12 families represented by more than one terminal, only two (Planaxidae, Potamididae) are always supported as monophyletic; Batillariidae, Cerithiidae, Pachychilidae, Pleuroceridae, Semisulcospiridae, Thiaridae, and Turritellidae are monophyletic in most but not all topologies. The combination of diverse data sources (morphology, 16S and 28S sequences) and inclusion of unconserved regions of the alignments improved the recovery of monophyletic families. At deeper levels, a consensus is beginning to emerge in the recognition of three main assemblages, but whether these represent clades or grades is still unclear; the resolution of these assemblages and the branching order within them are sensitive to exclusion of unconserved regions and choice of optimality criterion. No clear conclusion is reached with respect to the number of freshwater invasions, with two invasions supported on some topologies and three supported on others. Progress toward a robust and stable resolution of cerithioidean relationships will require (1) strategically coordinated sampling for additional morphological and molecular data; (2) comprehensive anatomical treatments for several poorly documented limnic lineages (e.g. Melanopsidae, Thiaridae) and comparative data for poorly understood organ systems (e.g. renal system); (3) the addition of poorly known, minute, and/or rare marine taxa, to provide novel character combinations, insight into putative homologies, and to help anchor basal nodes and break up long branches. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 43–89.     

Activity, release and flow of digestive enzymes in the cricket, Gryllus bimaculatus

Abstract.  In the cricket Gryllus bimaculatus , proventricular pressure forces a nutrient fluid from the ground food-brei through a sieve at the base of the caecae, and the combination of secreted enzymes and water causes a rapid inflation of the caecae on the first day after imaginal ecdysis. The ceacal region of the midgut is the primary site for the secretion of digestive enzymes. Proteases and amylase flow from the caecae into the mostly empty crop on day 1, and carbohydrate and protein digestion starts as soon as food is present (6 h). Thereafter, much of the amylase activity (but not protease) in the crop is synthesized and released by the crop tissues themselves. Regurgitating proteases and amylases from the caecae into the crop after day 1 is most likely accomplished by temporarily halting proventricular peristalsis and allowing the caecal muscles to contract, forcing caecal contents, including enzymes, forward. The total activity of digestive enzymes in the caecae is virtually identical in 2-day-old fed and unfed females, indicating little or no secretagogue (prandial) regulation of enzyme secretion. Most of the digestive enzymes in the ventricular endoperitrophic space may originate from the mucus dragged from the caecae. Lipase activity is low in all gut regions in both starved and fed females. Head ligation or injection of trypsin modulating oostatic factor, allatostatin A or B fails to indicate any involvement of nerves or hormones in the release of digestive enzymes in the caecae. Gryllus bimaculatus appears to secrete digestive enzymes continuously, and a considerable loss of enzymes may occur at certain times through egestion.     

Species richness estimation and determinants of species detectability in butterfly monitoring programmes

Abstract 1. Species richness is the most widely used biodiversity index, but can be hard to measure. Many species remain undetected, hence raw species counts will often underestimate true species richness. In contrast, capture–recapture methods estimate true species richness and correct for imperfect and varying detectability. 2. Detectability is a crucial quantity that provides the link between a species count and true species richness. For insects, it has hardly ever been estimated, although this is required for the interpretation of species counts. 3. In the Swiss butterfly monitoring programme about 100 transect routes are surveyed seven times a year using a highly standardised protocol. In July 2003, control observers made two additional surveys on 38 transects. Data from these 38 quadrats were analysed to see whether currently available capture–recapture models can provide quadrat‐specific estimates of species richness, and to estimate species detectability in relation to transect, observer, survey, region, and abundance. 4. Species richness over the entire season cannot be estimated using current capture–recapture methods. The species pool was open, preventing use of closed population models, and detectability varied by species, preventing use of current open population models. Assuming a closed species pool during two mid‐season (July) surveys, a Jackknife capture–recapture method was used that accounts for heterogeneity to estimate mean detectability and species richness. 5. In every case, more species were present than were counted. Mean species detectability was 0.61 (SE 0.01) with significant differences between observers (range 0.37–0.83). Species‐specific detection at time t+ 1 was then modelled for those species seen at t for three mid‐season surveys. Detectability averaged 0.50 (range 0.17–0.81) for individual species and 0.65, 0.44, and 0.42 for surveys. Abundant species were detected more easily, although this relationship explained only 5% of variation in species detectability. 6. These are important, although not entirely unexpected, results for species richness estimation of short‐lived animals. Raw counts of species may be misleading species richness indicators unless many surveys are conducted. Monitoring programmes should be calibrated, i.e. the assumption of constant detectability over dimensions of interest needs to be tested. The development of capture–recapture or similar models that can cope with both open populations and heterogeneous species detectability to estimate species richness should be a research priority.     

Subalpine grassland carbon dioxide fluxes indicate substantial carbon losses under increased nitrogen deposition,but not at elevated ozone concentration

Ozone (O₃) and nitrogen (N) deposition affect plant carbon (C) dynamics and may change ecosystem C‐sink/‐source properties. We studied effects of increased background [O₃] (up to [ambient] × 2) and increased N deposition (up to +50 kg ha^?1 a^?1) on mature, subalpine grassland during the third treatment year. During 10 days and 13 nights, distributed evenly over the growth period of 2006, we measured ecosystem‐level CO₂ exchange using a static cuvette. Light dependency of gross primary production (GPP) and temperature dependency of ecosystem respiration rates (R_eco) were established. Soil temperature, soil water content, and solar radiation were monitored. Using R_eco and GPP values, we calculated seasonal net ecosystem production (NEP), based on hourly averages of global radiation and soil temperature. Differences in NEP were compared with differences in soil organic C after 5 years of treatment. The high [O₃] had no effect on aboveground dry matter productivity (DM), but seasonal mean rates of both R_eco and GPP decreased ca. 8%. NEP indicated an unaltered growing season CO₂–C balance. High N treatment, with a +31% increase in DM, mean R_eco increased ca. 3%, but GPP decreased ca. 4%. Consequently, seasonal NEP yielded a 53.9 g C m^?2 (±22.05) C loss compared with control. Independent of treatment, we observed a negative NEP of 146.4 g C m^?2 (±15.3). Carbon loss was likely due to a transient management effect, equivalent to a shift from pasture to hay meadow and a drought effect, specific to the 2006 summer climate. We argue that this resulted from strongly intensified soil microbial respiration, following mitigation of nutrient limitation. There was no interaction between O₃ and N treatments. Thus, during the 2006 growing season, the subalpine grassland lost >2% of total topsoil organic C as respired CO₂, with increased N deposition responsible for one‐third of that loss.