Murderous plants: Victorian Gothic,Darwin and modern insights into vegetable carnivory

Darwin's interest in carnivorous plants was in keeping with the Victorian fascination with Gothic horrors, and his experiments on them were many and varied, ranging from what appears to be idle curiosity (e.g. what will happen if I place a human hair on a Drosera leaf?) to detailed investigations of mechanisms. Mechanisms for capture and digestion of prey vary greatly among the six (or more) lineages of flowering plants that have well‐developed carnivory, and some are much more active than others. Passive carnivory is common in some groups, and one, Roridula (Roridulaceae) from southern Africa, is so passively carnivorous that it requires the presence of an insect intermediate to derive any benefit from prey trapped on its leaves. Other groups not generally considered to be carnivores, such as Stylidium (Stylidiaceae), some species of Potentilla (Rosaceae), Proboscidea (Martyniaceae) and Geranium (Geraniaceae), that have been demonstrated to both produce digestive enzymes on their epidermal surfaces and be capable of absorbing the products, are putatively just as ‘carnivorous’ as Roridula. There is no clear way to discriminate between cases of passive and active carnivory and between non‐carnivorous and carnivorous plants – all intermediates exist. Here, we document the various angiosperm clades in which carnivory has evolved and the degree to which these plants have become ‘complete carnivores’. We also discuss the problems with definition of the terms used to describe carnivorous plants. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 161 , 329–356.     

The impacts of rising CO2 concentrations on Australian terrestrial species and ecosystems

The increasing atmospheric concentration of carbon dioxide ([CO₂]) contributes to global warming and the accompanying shifts in climate. However, [CO₂] itself has the potential to impact on Australia's terrestrial biodiversity, due to its importance in the photosynthetic process, which underlies all terrestrial food webs. Here, we review our knowledge regarding the impacts of elevated [CO₂] on native terrestrial species and ecosystems, and suggest key areas in which we have little information on this topic. Experimental information exists for 70 (or less than 0.05%) of Australia's native terrestrial plant and animal species. Of these, 68 are vascular plants. The growth of Australian woody species is more reliably increased by elevated [CO₂] than it is in grasses. At the species level, the most overwhelming responses to increased [CO₂] are a reduction in plant nitrogen concentration and an increase in the production of secondary metabolites. This is of particular concern for Australia's unique herbivorous and granivorous marsupials, for which no information is available. While many plant species also displayed increased growth rates at higher [CO₂], this was far from universal, indicating that changes in community structure and function are likely, leading to alterations of habitat quality. Future research should be directed to key knowledge gaps including the relationship between [CO₂], fire frequency and fire tolerance and the impacts of increasing [CO₂] for Australia's iconic browsing mammals. We also know virtually nothing of the impacts of the increasing [CO₂] on Australia's unique shrublands and semi‐arid/arid rangelands. In conclusion, there is sufficient information available to be certain that the increasing [CO₂] will affect Australia's native biodiversity. However, the information required to formulate predictions concerning the long‐term future of almost all organisms is far in excess of that currently available.     

Impacts of climate on prey abundance account for fluctuations in a population of a northern wader at the southern edge of its range

Understanding the mechanisms by which climate change will affect animal populations is vital for adaptive management. Many studies have described changes in the timing of biological events, which can produce phenological mismatch. Direct effects on prey abundance might also be important, but have rarely been studied. We examine the likely importance of variation in prey abundance in driving the demographics of a European golden plover ( Pluvialis apricaria ) population at its southern range margin. Previous studies have correlated plover productivity with the abundance of their adult cranefly (Tipulidae) prey, and modelled the phenology of both plover breeding and cranefly emergence in relation to temperature. Our analyses demonstrate that abundance of adult craneflies is correlated with August temperature in the previous year. Correspondingly, changes in the golden plover population are negatively correlated with August temperature 2 years earlier. Predictions of annual productivity, based on temperature-mediated reductions in prey abundance, closely match observed trends. Modelled variation in annual productivity for a future scenario of increasing August temperatures predicts a significant risk of extinction of the golden plover population over the next 100 years, depending upon the magnitude of warming. Direct effects of climate warming upon cranefly populations may therefore cause northward range contractions of golden plovers, as predicted by climate envelope modelling. Craneflies are an important food source for many northern and upland birds, and our results are likely to have wide relevance to these other species. Research into the potential for habitat management to improve the resilience of cranefly populations to high temperature should be an urgent priority.     

Environmental controls on water use efficiency during severe drought in an Ozark Forest in Missouri,USA

To accurately predict ecosystem responses induced by climate warming at local‐to‐global scales, models are in need of more precise knowledge of response during periods of environmental stress such as drought. In this paper, we studied environmental control of canopy‐level water use efficiency (WUE) during drought at an eddy flux site in an oak‐hickory forest in central Missouri, USA. Two consecutive severe droughts in the summers of 2006 and 2007 afforded coverage of a broad range of environmental conditions. We stratified data to obtain subranges that minimized cross‐correlations among putative WUE‐controlling factors. Our results showed that WUE was subject to control by atmospheric saturation deficit (ASD), soil water potential (SWP) and the ratio of diffuse to total photosynthetically active radiation (I_f/I_t). Generally, WUE was found to scale with 1/(ASD)^0.5, consistent with predictions from stomatal optimization theory. In contrast, SWP and I_f/I_t were related to WUE in a linear fashion. ASD was better correlated with WUE than either of the other two factors. It was also observed that the relationship between WUE and any single controlling factor was subject to influence of the other two. One such example was an opposite response of WUE to SWP between low and high ASD values, suggesting a breakdown of stomatal optimality under severe environmental stresses and a shift from optimal stomatal regulation to nonstomatal regulation at leaf scale. We have demonstrated that different data handling (stratified vs. nonstratified) or selection (hourly vs. daily) could lead to different conclusions on the relationship between WUE and its controls. For this reason, we recommend modelers to be cautious when applying WUE‐response formulas at environmental conditions or at time scales different from those at which they are derived.     

Variation in local abundance and species richness of stream fishes in relation to dispersal barriers: implications for management and conservation

1. Barriers to immigration, all else being equal, should in principle depress local abundance and reduce local species richness. These issues are particularly relevant to stream‐dwelling species when improperly designed road crossings act as barriers to migration with potential impacts on the viability of upstream populations. However, because abundance and richness are highly spatially and temporally heterogeneous and the relative importance of immigration on demography is uncertain, population‐ and community‐level effects can be difficult to detect. 2. In this study, we tested the effects of potential barriers to upstream movements on the local abundance and species richness of a diverse assemblage of resident stream fishes in the Monongahela National Forest, West Virginia, U.S.A. Fishes were sampled using simple standard techniques above‐ and below road crossings that were either likely or unlikely to be barriers to upstream fish movements (based on physical dimensions of the crossing). We predicted that abundance of resident fishes would be lower in the upstream sections of streams with predicted impassable barriers, that the strength of the effect would vary among species and that variable effects on abundance would translate into lower species richness. 3. Supporting these predictions, the statistical model that best accounted for variation in abundance and species richness included a significant interaction between location (upstream or downstream of crossing) and type (passable or impassable crossing). Stream sections located above predicated impassable culverts had fewer than half the number of species and less than half the total fish abundance, while stream sections above and below passable culverts had essentially equivalent richness and abundance. 4. Our results are consistent with the importance of immigration and population connectivity to local abundance and species richness of stream fishes. In turn, these results suggest that when measured at appropriate scales (multiple streams within catchments), with simple protocols amenable to use by management agencies, differences in local abundance and species richness may serve as indicators of the extent to which road crossings are barriers to fish movement and help determine whether road‐crossing improvements have restored connectivity to stream fish populations and communities.     

Molecular data and species distribution models reveal the Pleistocene history of the mayfly Ameletus inopinatus (Ephemeroptera: Siphlonuridae)1

1. We investigated the Pleistocene and Holocene history of the rare mayfly Ameletus inopinatus EATON 1887 (Ephemeroptera: Siphlonuridae) in Europe. We used A. inopinatus as a model species to explore the phylogeography of montane, cold‐tolerant aquatic insects with arctic–alpine distributions. 2. Using species distribution models, we developed hypotheses about the species demographic history in Central Europe and the recolonisation history of Fennoscandia. We tested these hypotheses using mitochondrial cytochrome oxidase I (mtCOI) sequence data and compared our genetic results with previously generated microsatellite data to explore genetic diversity distributions of A. inopinatus. 3. We observed old lineages, deep splits and almost complete lineage sorting of mtCOI sequences among mountain ranges. These results support a periglacial survival, i.e. persistence at the periphery of Pleistocene glaciers in Central Europe. 4. There was strong differentiation between the Fennoscandian and all other populations, indicating that Fennoscandia was recolonised from a refugium not accounted for in our sampling. High degrees of population genetic structure within the northern samples suggest that Fennoscandia was recolonised by more than one lineage. However, this structure was not apparent in previously published microsatellite data, consistent with secondary contact without sexual incompatibility or with sex‐biased dispersal. 5. Our demographic analyses indicate that (i) the separation of northern and Central European lineages occurred during the early Pleistocene; (ii) Central European populations have persisted independently throughout the Pleistocene and (iii) the species extended its range about 150 000 years ago.     

Leaf‐litter colonisation and breakdown in relation to stream typology: insights from Mediterranean low‐order streams

1. Scant information is available on leaf breakdown in streams of arid and semiarid regions, including the Mediterranean, where environmental heterogeneity can be high and the relationship between stream characteristics and leaf breakdown is poorly known. We tested the hypotheses that differences in leaf breakdown metrics would be substantially higher between mountain and lowland Mediterranean streams than among streams within each subregion and that variability among streams would be substantially higher in the lowlands, because permanent reaches in the semiarid lowland streams are rare and isolated. 2. We compared leaf breakdown and associated dynamics of nutrients, fungi and invertebrates in low‐order Mediterranean streams draining sub‐humid forests in the Sierra Nevada Mountains and nearby semiarid lowlands of south‐eastern Spain. Streams differed between the two subregions mainly in water ion content, temperature and riparian tree cover. We detected higher environmental heterogeneity among streams within the lowlands compared to the Sierra Nevada mountain range. In the lowlands, breakdown coefficients (k) of alder leaves spanned almost the entire range reported for this species from temperate streams, overlapping with less variable breakdown coefficients in the Sierra Nevada. 3. The high variability of k values among the lowland sites appeared to be caused primarily by variability in the composition and abundance of a few leaf‐consuming invertebrate taxa, particularly the snail Melanopsis praemorsa. Fungal and nutrient dynamics were less variable among sites within each subregion. 4. These results indicate that the critical condition for stream functional assessment of well‐constrained breakdown rates, or related metrics, could be met at reference sites within homogenous bio‐geo‐climatic regions such as the Sierra Nevada. By contrast, in heterogeneous areas such as the semiarid lowland streams, natural variability of breakdown rates can greatly exceed the magnitude of effects expected in response to anthropogenic disturbances.     

Reconstruction of the pharyngeal corpus of Aphelenchus avenae (Nematoda: Tylenchomorpha), with implications for phylogenetic congruence

The corpus of the pharynx in the nematode Aphelenchus avenae (Nematoda: Tylenchomorpha) was three‐dimensionally reconstructed to address questions of phylogenetic significance. Reconstructed models are based on serial thin sections imaged by transmission electron microscopy. The corpus comprises six classes of radial cells, two classes of marginal cells, and 13 neurones belonging to eight classes. Between the arcade syncytia and isthmus cells, numbers of cell classes along the pharyngeal lumen and numbers of nuclei per cell class correspond exactly between A. avenae and Caenorhabditis elegans. The number of radial cell classes between the arcade syncytia and the dorsal gland orifice (DGO) in A. avenae is also identical with outgroups. Proposed homologies of the pharynx imply that expression of the anterior two cell classes as epithelial or muscular differs within both Rhabditida and Tylenchomorpha. Numbers of neurone cell bodies within the corpus correspond exactly to C. elegans, other free‐living outgroups, and other Tylenchomorpha. Neurone polarity and morphology support conserved relative positions of cell bodies of putative neurone homologues. The configuration of cells in the procorpus, including the length of individual cell classes along its lumen, differs across representatives of three deep Tylenchomorpha lineages. Nonhomology of the procorpus challenges the homology of DGO position within the metacorpus, the primary taxonomic character for circumscribing ‘Aphelenchoidea’. Comparison of A. avenae with Aphelenchoides blastophthorus shows that, despite gross pharynx similarity, these nematodes have several differences in corpus construction at a cellular level. The possibility of convergent evolution of an ‘aphelenchid’ pharynx in two separate lineages would be congruent with molecular‐based phylogeny. Putative homologies and conserved arrangement of pharyngeal neurones in Tylenchomorpha expand the experimental model of C. elegans. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010.     

Effect of oceanographic barriers and overfishing on the population genetic structure of the European spiny lobster (Palinurus elephas)

Defining population structure and genetic diversity levels is of the utmost importance for developing efficient conservation strategies. Overfishing has caused mean annual catches of the European spiny lobster (Palinurus elephas) to decrease alarmingly along its distribution area. In this context, there is a need for comprehensive studies aiming to evaluate the genetic health of the exploited populations. The present study is based on a set of ten nuclear markers amplified in 331 individuals from ten different localities covering most of P. elephas distribution area. Samples from Atlantic and Mediterranean basins showed small but significant differences, indicating that P. elephas populations do not behave as a single panmictic unit but form two partially‐overlapping groups. Despite intense overfishing, our dataset did not recover a recent bottleneck signal, and instead showed a large and stable historical effective size. This result could be accounted for by specific life‐history traits (reproduction and longevity) and the limitations of molecular markers in covering recent timescales for nontemporal samples. The findings of the present study emphasize the need to integrate information on effective population sizes and life‐history parameters when evaluating population connectivity levels from genetic data. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 407–418.     

Moderating mycorrhizas: arbuscular mycorrhizas modify rhizosphere chemistry and maintain plant phosphorus status within narrow boundaries

Pastures often experience a pulse of phosphorus (P) when fertilized. We examined the role of arbuscular mycorrhizal fungi (AMF) in the uptake of P from a pulse. Five legumes (Kennedia prostrata, Cullen australasicum, Bituminaria bituminosa, Medicago sativa and Trifolium subterraneum) were grown in a moderate P, sterilized field soil, either with (+AMF) or without (?AMF) addition of unsterilized field soil. After 9–10 weeks, half the pots received 15 mg P kg^?1 of soil. One week later, we measured: shoot and root dry weights; percentage of root length colonized by AMF; plant P, nitrogen and manganese (Mn) concentrations; and rhizosphere carboxylates, pH and plant‐available P. The P pulse raised root P concentration by a similar amount in uncolonized and colonized plants, but shoot P concentration increased by 143% in uncolonized plants and 53% in colonized plants. Inoculation with AMF decreased the amount of rhizosphere carboxylates by 52%, raised rhizosphere pH by ～0.2–0.7 pH units and lowered shoot Mn concentration by 38%. We conclude that AMF are not simply a means for plants to enhance P uptake when P is limiting, but also act to maintain shoot P within narrow boundaries and can affect nutrient uptake through their influence on rhizosphere chemistry.