Herbivores equalize the seedling height growth of three dominant tree species in an African tropical rain forest

Determining the impact of insect herbivores on forest tree seedlings and saplings is difficult without experimentation in the field. Moreover, this impact may be heterogeneous in time and space because of seasonal rainfall and canopy disturbances, or ‘gaps’, which can influence both insect abundance and plant performance. In this study we used fine netting to individually protect seedlings of Microberlinia bisulcata, Tetraberlinia bifoliolata and Tetraberlinia korupensis trees (Fabaceae = Leguminosae) from insects in 41 paired gap-understorey locations across 80 ha of primary rain forest (Korup, Cameroon). For all species, growth in height and leaf numbers was negligible in the understorey, where M. bisulcata had the lowest survival after c. 2 years. In gaps, however, all species responded positively with pronounced above-ground growth across seasons. When exposed to herbivores their seedling height growth was similar, but in the absence of herbivores, M. bisulcata significantly outgrew both Tetraberlinia species and matched their leaf numbers. This result suggests that insect herbivores might play an important role in maintaining species coexistence by mitigating sapling abundance of the more palatable M. bisulcata, which in gaps was eaten the most severely. The higher ratio in static leaf damage of control-to-caged M. bisulcata seedlings in gaps than understorey locations was consistent with the Plant Vigour Hypothesis. This result, however, did not apply to either Tetraberlinia species. For M. bisulcata and T. korupensis, but not T. bifoliolata (the most shade-tolerant species), caging improved relative seedling survival in the understory locations compared to gaps, providing restricted support for the Limiting Resource Model. Approximately 2.25 years after treatments were removed, the caged seedlings were taller and had more leaves than controls in all three species, and the effect remained strongest for M. bisulcata. We conclude that in this community the impact of leaf herbivory on seedling growth in gaps is strong for the dominant M. bisulcata, which coupled to a very low shade-tolerance contributes to limiting its regeneration. However, because gaps are common to most forests, insect herbivores may be having impacts upon functionally similar tree species that are also characterized by low sapling recruitment much more widely than currently appreciated. An implication for the restoration and management of M. bisulcata populations in forests outside of Korup is that physical protection from herbivores of new seedlings where the canopy is opened by gaps, or by harvesting, should substantially increase its subcanopy regeneration, and thus, too, its opportunities for adult recruitment.

Hybridization increases invasive knotweed success

Hybridization is one of the fundamental mechanisms by which rapid evolution can occur in exotic species. If hybrids show increased vigour, this could significantly contribute to invasion success. Here, we compared the success of the two invasive knotweeds, Fallopia japonica and F.sachalinensis, and their hybrid, F.x bohemica, in competing against experimental communities of native plants. Using plant material from multiple clones of each taxon collected across a latitudinal gradient in Central Europe, we found that knotweed hybrids performed significantly better in competition with a native community and that they more strongly reduced the growth of the native plants. One of the parental species, F.sachalinensis, regenerated significantly less well from rhizomes, and this difference disappeared if activated carbon was added to the substrate, which suggests allelopathic inhibition of F.sachalinensis regeneration by native plants. We found substantial within-taxon variation in competitive success in all knotweed taxa, but variation was generally greatest in the hybrid. Interestingly, there was also significant variation within the genetically uniform F.japonica, possibly reflecting epigenetic differences. Our study shows that invasive knotweed hybrids are indeed more competitive than their parents and that hybridization increased the invasiveness of the exotic knotweed complex.

The prospect of applying chemical elicitors and plant strengtheners to enhance the biological control of crop pests

An imminent food crisis reinforces the need for novel strategies to increase crop yields worldwide. Effective control of pest insects should be part of such strategies, preferentially with reduced negative impact on the environment and optimal protection and utilization of existing biodiversity. Enhancing the presence and efficacy of native biological control agents could be one such strategy. Plant strengthener is a generic term for several commercially available compounds or mixtures of compounds that can be applied to cultivated plants in order to ‘boost their vigour, resilience and performance’. Studies into the consequences of boosting plant resistance against pests and diseases on plant volatiles have found a surprising and dramatic increase in the plants' attractiveness to parasitic wasps. Here, we summarize the results from these studies and present new results from assays that illustrate the great potential of two commercially available resistance elicitors. We argue that plant strengtheners may currently be the best option to enhance the attractiveness of cultivated plants to biological control agents. Other options, such as the genetic manipulation of the release of specific volatiles may offer future solutions, but in most systems, we still miss fundamental knowledge on which key attractants should be targeted for this approach.

Canopy cover mediates interactions between a specialist caterpillar and seedlings of a neotropical tree

1 Light availability may be crucial for understanding dynamics of plant–herbivore interactions in temperate and tropical forest communities. This is because local light availability can influence both tree seedling tolerance and susceptibility to herbivory – yet how they mediate levels of insect herbivory that vary with the density of host population is virtually unknown. Here we tested predictions of three key, non-mutually exclusive hypotheses of plant–herbivore interactions: the Limiting Resource Model (LRM), the Plant Vigour Hypothesis (PVH), and the Janzen-Connell Mechanism (JCM). 2 In an Amazonian forest, we planted Swietenia macrophylla seedlings (c. 5 months old) into natural canopy gaps and the shaded understorey and simulated the damage patterns of the specialist herbivore moth, Steniscadia poliophaea, by clipping seedling leaves. Over the next 8 months, we monitored seedling performance in terms of growth and survivorship and also quantified herbivory to new young leaves on a seasonal basis. 3 In support of the LRM, severe leaf damage (≥ 50%) was lethal for Swietenia macrophylla seedlings in the understorey, but in gaps only reduced seedling growth. In support of the PVH, gap seedlings suffered greater post-simulated herbivory (up to 100% defoliation) by S. poliophaea caterpillars than their understorey counterparts. 4 Adding a novel dimension to the Janzen–Connell hypothesis, we found that early wet season herbivory of seedlings in gaps increased with conspecific adult density within a 125-m radius; whereas in the understorey only those seedlings within 50 m of a Swietenia tree were attacked by caterpillars. 5 Synthesis. These results suggest lepidopterans that need young leaves for food may forage more widely in forests to find seedlings in light-rich canopy gaps. Moths may achieve this successfully by being first attracted to gaps, and then searching within them for suitable hosts. A conceptual model, integrating conspecific adult tree density with light-driven changes in seedling tolerance/vigour and their susceptibility to herbivory and mortality, is presented. Spatial variation in the light available to tree seedlings often affects their tolerance and vigour, which may have important consequences for leaf-chewing insects and the scale of density-dependent herbivory in forests.

Dinámica poblacional de planococcus ficus sign. (Hemiptera-Pseudcoccidae) en viñedos : Mendoza (Argentina)

Planococcus ficus Signoret es un insecto perteneciente al orden Hemiptera, familia Pseudococcidae, cuya población ha aumentado en los últimos años en Mendoza. Ataca gran variedad de hospederos y está distribuido en las principales zonas vitícolas del mundo. Su acción disminuye el vigor general de las plantas, perjudica la calidad de los frutos y las características organolépticas de los vinos elaborados con racimos infestados. El objetivo de este trabajo fue realizar estudios bioecológicos durante la temporada 2004-2005, analizando la forma de pasaje invernal y el número de generaciones anuales. Se marcaron y muestrearon plantas con los signos característicos del ataque. Se recolectó quincenalmente material vegetal infestado. En invierno: trozos de corteza. En primavera-verano: brotes, hojas y racimos. Los resultados indicaron que pasa el invierno principalmente como huevos agrupados en masas algodonosas, bajo la corteza. Se distinguieron seis generaciones, movilizándose conforme al aumento de las temperaturas hacia los brotes y hojas y, a partir de la tercera generación, a los racimos, permaneciendo en todas las partes aéreas de la planta el resto de la temporada hasta la caída de hojas. Con la disminución de las temperaturas retornó hacia la corteza del tronco para invernar.

Influencia de seis portainjertos de vid sobre el comportamiento vitícola de la cv. Malbec y estudio de las relaciones hídricas que se establecen

El uso de portainjertos en la vid se ha difundido por su resistencia a filoxera y nemátodos, pero también por su tolerancia a condiciones adversas del suelo. Por otro lado, los portainjertos modifican las relaciones fuente-destino, influyendo en el comportamiento vegetativo y reproductivo de las plantas y en la composición de la uva, lo cuál puede ser utilizado como una herramienta de manejo agronómico. A fin de evaluar si existe un comportamiento diferencial de los portainjertos en cuanto a expresión vegetativa, vigor, rendimiento y composición de la uva, y explicar dichas diferencias en términos de exploración radical, relaciones hídricas, asimilación de carbono, eficiencia en el uso del agua y partición de asimilados se realizó un ensa-yo a campo de cv. Malbec sobre seis portainjertos (3309 C, 1103 P, 140 Ru, SO4, Harmony y Cereza) y a pie franco. Los portainjertos 140 Ru, 1103 P y SO4 tuvieron una mayor tendencia a la producción de uva (mayor Índice de Ravaz), y Franco, Cereza y 3309 C a vegetar, mostrando Harmony una situación intermedia. Las ba-yas sobre el pie Cereza tuvieron un mayor peso (1,96 g) que sobre Harmony (1,75 g). No se encontraron diferencias en los polifenoles de las bayas entre portainjertos. La fotosíntesis de la planta entera (Amax) de Franco, 1103 P y SO4 fue mayor que la de Harmony. La conductancia hidráulica foliar específica (kL) de Harmony fue me-nor que la de Cereza, y su conductancia hidráulica (kH) fue menor que la de Franco, Cereza y SO4. El número de raíces totales de 140 Ru fue mayor que el de 1103 P, SO4 y Harmony. El portainjerto 140 Ru se destacó por privilegiar el desarrollo radi-cal y reproductivo sobre el vegetativo, y por su mayor eficiencia en el uso del agua (EUA). Las diferencias entre portainjertos pueden ser explicadas en parte por dife-rencias en la kL que a su vez incide en el estado hídrico de las plantas (ΨL). De ma-nera que cuando la kL es más baja, el ΨL es menor (i.e., Harmony), y cuando la kL es más alta, el ΨL es mayor (i.e., Franco y Cereza). Mayores ΨL se asocian con mayores superficies foliares.

Stable oxygen isotope ratios and grain size analyses of sediment cores from the Propeller Mound, Northeast Atlantic

The first detailed stratigraphic record from a deep-water carbonate mound in the Northeast Atlantic based on absolute datings (U/Th and AMS 14C) and stable oxygen isotope records reveals that its top sediment sequences are condensed by numerous hiatuses. According to stable isotope data, mainly sediments with an intermediate signal are preserved on the mound, while almost all fully glacial and interglacial sediments have either not been deposited or have been eroded later. The resulting hiatuses reduce the Late Pleistocene sediment accumulation at Propeller Mound to amounts smaller than the background sedimentation. The hiatuses most likely result due to the sweeping of the mound in turn with the re-establishment of vigour interglacial circulation patterns after sluggish current regimes during glacials. Thus, within the discussion if internal, fluid-driven or external environmentally driven processes control the evolution of such carbonate mounds, our findings for Propeller Mound clearly point to environmental forcing as the dominant mechanism shaping deep-water carbonate mounds in the NE Atlantic during the Late Pleistocene and Holocene.

XRF scanning, pollen and spores of ODP Site 108-659

Pliocene vegetation dynamics and climate variability in West Africa have been investigated through pollen and XRF-scanning records obtained from sediment cores of ODP Site 659 (18°N, 21°W). The comparison between total pollen accumulation rates and Ti/Ca ratios, which is strongly correlated with the dust input at the site, showed elevated aeolian transport of pollen during dusty periods. Comparison of the pollen records of ODP Site 659 and the nearby Site 658 resulted in a robust reconstruction of West African vegetation change since the Late Pliocene. Between 3.6 and 3.0 Ma the savannah in West Africa differed in composition from its modern counterpart and was richer in Asteraceae, in particular of the Tribus Cichorieae. Between 3.24 and 3.20 Ma a stable wet period is inferred from the Fe/K ratios, which could stand for a narrower and better specified mid-Pliocene (mid-Piacenzian) warm time slice. The northward extension of woodland and savannah, albeit fluctuating, was generally greater in the Pliocene. NE trade wind vigour increased intermittently around 2.7 and 2.6 Ma, and more or less permanently since 2.5 Ma, as inferred from increased pollen concentrations of trade wind indicators (Ephedra, Artemisia, Pinus). Our findings link the NE trade wind development with the intensification of the Northern Hemisphere glaciations (iNHG). Prior to the iNHG, little or no systematic relation could be found between sea surface temperatures of the North Atlantic with aridity and dust in West Africa.

Sedimentology and Cd/Ca ratios of benthic foraminifera from ODP Site 172-1060 in the western Nort Atlantic

The Western Boundary Undercurrent (WBUC), off eastern America, is an important component of the Atlantic Meridional Overturning circulation and is the principal route for southward transport of North Atlantic waters and southward return of Southern Source Water (SSW). Here a direct flow speed proxy (mean grain size of the sortable silt) is used to infer the vigour of flow of the palaeo-WBUC at Blake Outer Ridge, (ODP Site 1060, depth 3481 m) during Marine Isotope Stage (MIS) 3. The overall shape of the flow speed proxy record shows a complex pattern of variability, with generally more vigorous flow and larger-scale flow variations between 35 and 60 ka than in the younger part of MIS 3 and MIS 2 (b35 ka). Six events of reduced bottom flow vigour (Slow Events, SEs) occur. These appear uncorrelated with Heinrich events, but are instead synchronous with the warming phases of Antarctic Warm Events A-1 to A-4 (with one new one, A-1a and one poorly defined, 'A-0'). This indicates that Antarctic climate exerts a stronger control on deep flow vigour in the North Atlantic during MIS 3 than Northern Hemisphere climate. The correspondence of SEs with Antarctic warming suggests a weaker WBUC flow due to reduced volume flux at SSW source or reduced SSW density. Because the variability of the lower limb of the WBUC was not connected to sharp North Atlantic changes in temperature, it is unlikely that the Dansgaard/Oeschger cycles were associated with a mode of MOC variation involving wholeocean overturn, but more likely with perturbations of only the shallow Glacial Gulf Stream-Glacial Northern Source Intermediate Water cell. Nutrient proxies (benthic carbon isotopes and Cd/Ca of Uvigerina peregrina) at this site show similar trends to the GRIP delta18O record. This correlation has previously been attributed mainly to hydrographic and flow changes but is here shown to be better explained by variations in surface ocean productivity and subsequent decomposition of 12C rich organic material on the sea floor.

Biostratigraphy of selected taxa and radiolarian events in ODP Hole 183-1138A sediments

Well-preserved radiolarian assemblages of late middle Miocene to early Pliocene age are found in Ocean Drilling Program (ODP) Hole 1138A (Cores 183-1138A-12R to 20R), which was rotary drilled into the Central Kerguelen Plateau. The faunas are typical for Antarctic assemblages of this time interval, and the site appears to have been south of the Polar Front during the time period studied. Despite only moderate drilling recovery of the section, most late middle to early Pliocene radiolarian zones are present, although at the sample resolution used, subzones could not be identified. A significant discontinuity in the section is present at the boundary between lithologic Units I and II (between Cores 183-1138A-12R and 13R), corresponding to an interval from at least 4.6 to 6.1 Ma. Mixed late Miocene-early Pliocene assemblages are seen in the base of Core 183-1138A-12R (Sample 183-1138A-12R-3, 20 cm), and the overlying basal Pliocene Tau Zone appears to be absent. It cannot be determined if the discontinuity is due to incomplete recovery of the section and drilling disturbance or if it reflects a primary sedimentary structure - a hiatus or interval of condensed sedimentation.

Locations of geochemical samples and tephra from ODP Site 181-1123

Approximately one thousand sediment samples from ODP Site 1123 on the Chatham Rise, east of New Zealand, have been examined for inorganic elemental concentrations. ODP 1123 provides a record of sediment drift deposition under the Deep Western Boundary Current, the main inflow of deep water to the Pacific Ocean since the Early Oligocene, though a major hiatus spans the late Early Oligocene to the Early Miocene. Normalisation of the elemental concentrations by aluminium was used to allow for the effects of variable carbonate dilution. The elemental ratios were used as proxies for sediment composition and as palaeoceanographic indices. The samples were collected at a resolution designed to sample adequately any variation in elemental ratios at the scale of the Milankovitch orbital cycles. The sampled intervals span the Early Oligocene, Early Miocene, mid-Miocene and Late Pleistocene to Recent. Anomalous Si/Al, K/Al, Ti/Al values in the upper Pleistocene section, often associated with horizons of low carbonate, are attributed to tephras derived from North Island. Not all of the tephras detected geochemically had been detected visually in the cores. A total of 37 tephra events between 1.17 Ma BP and the present are recognised based on this and the shipboard investigations. The tephra events cluster at intervals of approximately 326 000 years (326 ka) perhaps due to variations in eruption frequency on North Island and/or to variations in the regional palaeowind intensity and direction. In the Late Pleistocene to Recent P/Al (inferred nutrient availability), percent calcium carbonate (%CaCO3) and Ba/Al (inferred productivity) varied regularly at a period of 40 000 years with these factors lagging minimum global ice volumes (interglacials). During the mid-Miocene CaCO3, Ba/Al, P/Al and Si/Al all gradually increased with %CaCO3 and P/Al showing regular 138 000-yr cyclicity and Ba/Al showing 44-ka cyclicity. Inferred productivity (Ba/Al) may have been rising in association with increasing nutrient availability (P/Al) at the same time as increased vigour of the Deep Western Boundary Current that was connected to a period of rapid ice-sheet growth in Antarctica. In the Early Miocene P/Al and Si/Al were much higher than subsequently and both %CaCO3 and P/Al exhibited 131 000-yr cycles. By far the highest nutrient levels and inferred productivity at this site apparently occurred during the Early Oligocene as revealed by long-term changes in P/Al and Si/Al. A progressive rise in K/Al, but stable Ti/Al from the Early Oligocene to the Recent probably indicates increased proportions of illite in the clay mineral fraction of the drift sediments caused by increased flux of debris from the Southern Alps.

(Appendix) Analysis of benthic foraminifera from ODP Leg 181 and DSDP Leg 90

Canonical correspondence analysis indicates that the distribution of Neogene benthic foraminiferal faunas (>63 µm) in seven DSDP and ODP sites (500-4500 m water depth) east of New Zealand (38-51°S, 170°E-170°W) is most strongly influenced by depth (water mass stratification), and secondly by age (palaeoceanographic changes influencing faunal composition and biotic evolution). Stratigraphic faunal changes are interpretted in terms of the pulsed sequential development of southern, and later northern, polar glaciation and consequent cooling of bottom waters, increased vertical and lateral stratification of ocean water masses, and increased overall and seasonal surface water productivity. Oligocene initiation of the Antarctic Circumpolar Current and Deep Western Boundary Current (DWBC), flowing northwards past New Zealand, resulted in extensive hiatuses throughout the Southwest Pacific, some extending through into the Miocene. Planktic foraminiferal fragmentation index values indicate that carbonate dissolution was significant at abyssal depths throughout most of the Neogene, peaking at upper abyssal depths in the late Miocene (11-7 Ma), with the lysocline progressively deepened thereafter. Miocene abyssal faunas are dominated by Globocassidulina subglobosa and Oridorsalis umbonatus, with increasing Epistominella exigua after 16 Ma at upper abyssal depths. Peak abundances of Epistominella umbonifera indicate increased input of cold Southern Component Water to the DWBC at 7-6 Ma. Faunal association changes imply establishment of the modern Oxygen Minimum Zone (upper Circumpolar Deep Water) in the latest Miocene. Significant latitudinal differences between the benthic foraminiferal faunas at lower bathyal depths indicate the existence of an oceanic front along the Chatham Rise (location of present Subtropical Front), since the early late Miocene at least, with more pulsed productivity (higher E. exigua) along the south side. Modern Antarctic Intermediate Water faunal associations were established north of the Chatham Rise at 10-9 Ma, and south of it at 3-1.5 Ma. Middle-upper bathyal faunas on the Campbell Plateau are dominated by reticulate bolivinids during the early and middle Miocene, indicative of sustained productivity above relatively sluggish, suboxic bottom waters. Faunal changes and hiatuses indicate increased current vigour over the Campbell Plateau from the latest Miocene on. Surface water productivity (food supply) appears to have increased in three steps (at times of enhanced global cooling) marked by substantially increased relative abundance of: (1) Abditodentrix pseudothalmanni, Alabaminella weddellensis, Cassidulina norvangi (16-15 Ma, increased pulsed productivity); (2) Bulimina marginata f. aculeata, Nonionella auris, Trifarina angulosa, Uvigerina peregrina (3-1.5 Ma, increased overall productivity); and (3) Cassidulina carinata (1-0.5 Ma, increased overall productivity). Three intervals of deep-sea benthic foraminiferal taxonomic turnover are recognised (16-15, 11.5-10, 2-0.5 Ma) corresponding to intervals of enhanced global cooling and possible productivity changes. The late Pliocene-middle Pleistocene extinction, associated with increasing Northern Hemisphere glaciation, culminating in the middle Pleistocene climatic transition, was more significant in the study area than the earlier Neogene turnovers.