Insect traces on early Permian plants of India

The study of fossil leaves from the Glossopteris flora of the Lower Permian Gondwana sequence of the Barakar Formation of the Raniganj Coalfield, India, has revealed the presence of different types of insect traces. The structures are compatible with a variety of insect activities on fossil leaves. Nibbled and cuspate margins, trench marks, obliterated surfaces, blotch marks, and holes of various shape constitute feeding traces, whereas egg pouches along the midrib and irregularly distributed oviposition marks are traces of egglaying. Although fossil records of both insects and their activities in the Glossopteris flora of India remain scarce, the different types of insect traces identified in the present study demonstrate the existence of a diverse insect fauna during the Permian Period of India.     

An efficient method for estimating vein density of Glossopteris and its application

Glossopteris-type leaves are the most abundant floristic element from the Gondwanan continent and are recorded throughout the Permian, which was a period of extreme icehouse-to-hothouse climatic global change. Fossil leaf traits can be useful for the reconstruction of palaeoenvironments and identification of climatic changes throughout geological time, but the conservative morphology of Glossopteris leaves has thus far made them difficult to use for this purpose. If the characters of Glossopteris can be better quantified then it should make them useful for tracking environmental changes over a wide geographical area and over a long time interval. Venation density is a highly variable leaf trait that might be useful for this purpose. This trait can be calculated, usually as vein length per centimetre squared, but this can be a time-consuming procedure. In this paper we propose a new rapid method to estimate venation density in a conical sector of Glossopteris leaf lamina using an accurate linear model whose predictors are three linear venation densities, measured as veins per centimetre. In addition to substantially reducing the data collection time, it is less biased and more reproducible than methods applied previously with this leaf type. Using this more robust method, preliminary results significantly distinguish the venation densities of leaves produced in wet and drier ecosystems, matching a pattern similar to modern plants. This is the first survey using a large sample size to reveal that environmental stress controlled the vein architecture of Palaeozoic plants, in a manner similar to plants in modern ecosystems.     

Hypersensitivity of Fagus sylvatica L. against leaf galling insects

Hypersensitivity is known as a localized resistance of plants against pathogens. It also can be detected in response to galling insects, i.e., in the area immediately adjacent to the site of oviposition and attempted penetration by the galling larva. This host response includes morphological and histological changes that cause the death of the attacked tissue. It is observed as a rounded dark brown halo around the gall induction site. We provide the first observation on the occurrence and possible relevance of this induced mechanism by which one of the most common tree species in Germany, Fagus sylvatica L., resists attack by two of its most common galling insects, Mikiola fagi and Hartigiola annulipes (Diptera: Cecidomyiidae). Galls induced by these cecidomyiids were extremely common in the studied area in beech forests around Darmstadt, Germany. The availability of resources (leaves on a stem) was a poor predictor of attack by the galling insects as well as for gall abundance (galls successfully formed). Hypersensitive reaction was the most important factor acting against the galling population studied. More than 77% of the attempts of the insects to induce galls on F. sylvatica resulted in failure and consequently the death of the galling larvae. Therefore, few live galls remained to be found and destroyed by natural enemies. This corroborates the view that in galling insect–host plant system interactions plant-driven factors may play a major role in determining herbivore failure and success, and perhaps the resulting community structure.     

Levels of polyphenol oxidase activity in leaves of Milicia seedlings is indicative of the resistance to Phytolyma lata-induced gall disease

1. Phytolyma lata-induced galling has remained one of the foremost challenges limiting the regeneration of Milicia species in nurseries and degraded forests in West Africa. Although planting of Milicia progenies resistant to Phytolyma attack has been identified as an opportunity for long-term control of the pest, the absence of biomarkers for early detection of resistance has limited the selection of resistant seedlings for planting.
2. We compared phenolic metabolites and oxidative enzymes in resistant and susceptible Milicia progenies with the aim of identifying potential biomarkers for resistance to Phytolyma-induced galling in seedlings.
3. Levels of total phenolics and ortho-dihydroxyphenolics did not significantly differ between the susceptible and resistant saplings or among seedlings. Similarly, peroxidase activity did not differ among sapling progenies or among seedlings. Thus, no clearly defined role could be ascribed to phenolics and peroxidase in resistance of Milicia species to P. lata-induced galling. However, polyphenol oxidase (PPO) activity was over two-fold higher in the resistant than the tolerant or susceptible progenies, suggesting that higher levels of PPO activity corresponded with an increased level of resistance to galling.
4. With further validation, PPO activity in leaves of Milicia seedlings can be used as a biomarker of resistance to Phytolyma-induced galling.

     

Distribution and oviposition preference of galling sawflies in arctic Canada

The distribution and ecology of insects in arctic regions are poorly known. The aim of this study was to determine the distribution of galling sawflies in the Canadian arctic and their oviposition preference. The Swedish Tundra Northwest 1999 expedition visited 17 sites in the Canadian arctic. We determined the occurrence of galling sawflies at all the sites and studied the oviposition preference of two leaf-galling sawflies, Eupontania arctica and Pontania nivalis, on Salix reticulata and S. glauca, respectively. Galling sawflies were abundant at only one site, the mainland site at Ivvavik National Park. Only a few galls in total were found at the remaining sites, suggesting that galling sawflies are rare in the higher arctic, and potential explanations for this pattern are discussed. Shoots with leaf galls were longer than shoots without galls on both S. reticulata and S. glauca. These differences could not be explained by a higher number of leaves on longer shoots. This suggests that long shoots are preferred by sawflies because of faster development and better survival of larvae on long shoots.     

Management of Root-Knot Nematode,Meloidogyne Incognita,by integration of Paecilomyces Lilacinus with organic materials in Chilli

Studies were made to determine the efficacy of Paecilomyces lilacinus in management of root-knot nematode (Meloidogyne incognita) in soil amended with various organic matters. The soil amendments with organic additives except gram and rice husks significantly reduced the multiplication of M. incognita and the root galling caused by root-knot nematode which consequently increased the plant growth. The greatest improvement in plant growth and reduced reproduction factor and root galling was recorded in soil amendment with leaves of Calotropis procera while the least was in kail saw dust. The best protection against M. incognita was observed on the integration of organic additives with P. lilacinus, which resulted increased plant growth and reduced population build-up of nematodes and root gallings. The leaves of C. procera with P. lilacinus were most effective than all other organic materials used among the different integrated approaches. The organic amendments also increased the parasitism of P. lilacinus on M. incognita.     

Assessment of three strategies for the management of Meloidogyne arenaria on carrot in Mexico using Pochonia chlamydosporia var. mexicana under greenhouse conditions

Three control strategies for management of Meloidogyne arenaria were evaluated on carrot under greenhouse conditions. The control strategies tested were: i) incorporation of fresh broccoli; ii) carbofuran (Furadan®); and iii) Pochonia chlamydosporia var. mexicana, isolate Pcp21. Each strategy was evaluated separately and in combination (16 treatment combinations), each with three replicates. The experiment was done between February and April 2014 and repeated over the same time period in 2015. Each replicate experimental unit consisted of a 1?kg pot filled with tyndallised soil, to which broccoli, fungus and carbofuran were added according to treatment. Carrot seeds were sown into each pot and inoculated with M. arenaria eggs according to treatment. Pots were then maintained in the glasshouse for 12 weeks at 25?±?5°C. After this time the fresh root weight, root length, percent reduction in root galling, P. chlamydosporia var. mexicana colony forming units (CFU)/g of soil, and CFU/g root were all measured. Results showed that isolate Pcp21 of P. chlamydosporia var. mexicana significantly reduced root galling percentage by 50 and 78% in 2014 and 2015, respectively. Application of the fungus in combination with broccoli or carbofuran also reduced root galling. The number of CFU/ g of soil or root remained the same over the 12 weeks of the experiment. This isolate of P. chlamydosporia significantly reduced damage caused by M. arenaria on carrot and is a promising agent for management of this nematode.     

A gall-inducing arthropod drives declines in canopy tree photosynthesis

Mature forest canopies sustain an enormous diversity of herbivorous arthropods; however, with the exception of species that exhibit large-scale outbreaks, canopy arthropods are thought to have relatively little influence on overall forest productivity. Diminutive gall-inducing mites (Acari; Eriophyoidae) are ubiquitous in forest canopies and are almost always highly host specific, but despite their pervasive occurrence, the impacts of these obligate parasites on canopy physiology have not been examined. We have documented large declines in photosynthetic capacity (approx. 60%) and stomatal conductance (approx. 50%) in canopy leaves of mature sugar maple (Acer saccharum) trees frequently galled by the maple spindle gall mite Vasates aceriscrumena. Remarkably, such large impacts occurred at very low levels of galling, with the presence of only a few galls (occupying approx. 1% of leaf area) compromising gas-exchange across the entire leaf. In contrast to these extreme impacts on the leaves of adult trees, galls had no detectible effect on the gas-exchange of maple saplings, implying large ontogenetic differences in host tolerance to mite galling. We also found a significant negative correlation between canopy tree radial increment growth and levels of mite galling. Increased galling levels and higher physiological susceptibility in older canopy trees thus suggest that gall-inducing mites may be major drivers of “age-dependent” reductions in the physiological performance and growth of older trees.     

Vigour of a dioecious shrub and attack by a galling herbivore

1. The pattern of attack by the leaf‐galling insect Neopelma baccharidis (Homoptera: Psyllidae) was studied in three populations of the dioecious shrub Baccharis dracunculifolia (Asteraceae) in south‐eastern Brazil. The plant vigour hypothesis, which predicts higher rates of attack and increased herbivore performance on the longest plant shoots, was tested. This work also provides further information for the study of differential herbivory in dioecious plants. 2. In total, 9200 shoots were collected randomly from 46 male and 47 female plants belonging to the three populations. Shoot length, number of leaves per shoot, rate of galling, and survival of psyllids did not differ between male and female plants. Another population on the Campus of the Federal University of Minas Gerais was used only to determine the pattern of shoot growth. 3. The hypothesis of sex‐mediated herbivory was not corroborated in this study. 4. The frequency of galling increased with increasing shoot length, as predicted by the plant vigour hypothesis. Nevertheless, the number of oviposition sites (leaf buds) increased with shoot length. 5. The performance of the galling herbivore was not related to shoot length in the plant populations studied. 6. In conclusion, Neopelma baccharidis did not select shoots based on length only.     

Evolution of the glossopterid gymnosperms from Permian Gondwana

Our understanding of the glossopterid gymnosperms from the Permian of Gondwana has recently been greatly enhanced through the study of anatomically preserved plant remains in cherts from several localities in Australia and Antarctica. Studies ofGlossopteris leaves, leaf-stem attachment, stem structure and wood anatomy,Vertebraria roots, ovule- and pollen-bearing reproductive structures and putative glossopterid pollen and ovules demonstrate that the glossopterids were a diverse group of sophisticated gymnosperms. Comparative studies of different Permian chert floras show that while some glossopterid elements are widespread, other taxa appear to be more localized in their distribution. As information from petrified plant-bearing cherts becomes available, it can be used to reconstruct whole glossopterid plants and establish their paleogeographic distribution.     

The interplay between plant traits and herbivore attack: a study of a stem galling midge in the neotropics

1. The relationship between plant traits and the frequency of attack by a stem galling midge, Neolasioptera sp. (Diptera: Cecidomyiidae), on Eremanthus erythropappus (Asteraceae) was studied. The morphological changes of the host after a galler attack and the potential effects of these changes on attacks by the next generation of gallers were analysed. The study was conducted in the Serra do Japi, São Paulo, south-eastern Brazil. 2. Galled branches were significantly longer, thicker, and had more leaves than ungalled branches. Accordingly, gall establishment was higher in the longer and more foliose branches. Hence, it is suggested that ovipositing females are maximizing their performance by selecting larger branches. 3. Galled branches were larger than ungalled branches of the same age. Two hypotheses, not necessarily exclusive, can explain this pattern: (1) the plant vigour hypothesis that the females are choosing the more vigorous, fast-growing branches, which still remain more vigorous after galling; or (2) the resource regulation hypothesis that galling increases branch growth rates, thus increasing resource quality for forthcoming conspecifics. 4. Co-occurrence frequencies of current and past generation galls showed that the likelihood of a branch being galled increased when it, or the branch from which it stemmed, had been galled before. The data indicated that this preference was conditioned by the number of previous attacks. Heavier attack intensities, such as one gall in the same branch and another in the branch from which it stemmed, decreased the probability of further galling. 5. The suggested links between herbivore attack and plant traits indicate that studies on host selection by phytophagous insects must take into account that herbivory itself may change the plant traits that are postulated to be selected by the insects.     

Limits of stratigraphic distribution of Glossopteris in India: Discussion

The stratigraphic limits of Glossopteris in India have been reviewed by Sarbadhikari (1974) and are inferred to range from the Sakmarian-Artinskian (early Permian) to the Permo-Triassic transition. The lower limit in peninsular India is revised as Asselian with the help of Eurydesma-dominated fauna. Glossopteris was well established during the Asselian in the Salt Range but appeared during the late Sakmarian-Artinskian in the Himalayas. In peninsular India, Glossopteris flourished well during the middle Triassic and continued at least upto the Rhaetic, in association with Dicroidium. The Glossopteris-Dicroidium assemblage thus is not a time-restricted transitional zone. Dicroidium with associated forms characterises a flora which is distinct from the older Glossopteris flora and the younger Ptillophylum flora. A three-fold biostratigraphic subdivision of the Indian Gondwana thus has been advocated.     

Morphological studies on the genusClematis linn

In four-sepaled flowers ofClematis the sepal is supplied by three main traces. The basic pattern of the vascular supply to sepals is found inC. alpina var.ochotensis which invariably has six-bundled pedicels. It is as follows: the median traces to the first pair of opposite sepals, as well as all the lateral traces, arise directly from pedicel bundles, while those to the second pair are formed secondarily, after fusion and subsequent division of two adjacent pedicel bundles. As to the manner of origin of the median traces, the pattern is similar to that of the vascular supply to foliage leaves. This gives further evidence for the generally accepted view that the sepals ofClematis, like foliage leaves, are decussately arranged. In most other species such asC. apiifolia, C. stans, etc. the number of pedicel bundles tends to be reduced from six to four so as to coincide with that of the sepals, so patterns are much simplified and specialized: all the traces arise directly from pedicel bundles. InC. japonica an iconsistent pattern is observed, since the number of pedicel bundles from which sepal traces arise is much higher and varied.     

Timing of growth determines fitness and performance of a galling insect on willow

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A QUANTITATIVE STUDY OF SIEVE-TUBE DIFFERENTIATION IN VEGETATIVE SHOOT APICES OF COLEUS

Quantitative methods, with round-the-clock collecting of large samples in successive years, have uncovered several new phenomena of sieve-tube differentiation in young leaves of Coleus vegetative shoots. In small leaves (1–350 μ), there are no sieve tubes in the leaf itself, but they differentiate acropetally in the two traces to each leaf. Regression lines fitted to the data for leaf length vs. most distal position of sieve tubes in the traces support the view that differentiation is steady and acropetal, but they also reveal that differentiation in the traces falls steadily farther behind elongation of the leaf. Leaves more than 500 μ long have sieve tubes close to their tips. An intensive search of leaves of intermediate lengths revealed an isolated locus of sieve-tube differentiation. These relationships were reproducible year after year. Every plant with discontinuous strands of sieve tubes in the second leaf pair had discontinuous xylem in the third. This isolated locus was not seen before, probably because of small samples and daytime collections; most of our cases were from night collections.     

Manipulation of food resources by a gall-forming aphid: the physiology of sink-source interactions

Summary We examined the capacity of the galling aphid, Pemphigus betae, to manipulate the sink-source translocation patterns of its host, narrowleaf cottonwood (Populus angustifolia). A series of ¹⁴C-labeling experiments and a biomass allocation experiment showed that P. betae galls functioned as physiologic sinks, drawing in resources from surrounding plant sources. Early gall development was dependent on aphid sinks increasing allocation from storage reserves of the stem, and later development of the progeny within the gall was dependent on resources from the galled leaf blade and from neighboring leaves. Regardless of gall position within a leaf, aphids intercepted ¹⁴C exported from the galled leaf (a non-mobilized source). However, only aphid galls at the most basal site of the leaf were strong sinks for ¹⁴C fixed in neighboring leaves (a mobilized source). Drawing resources from neighboring leaves represents active herbivore manipulation of normal host transport patterns. Neighboring leaves supplied 29% of the ¹⁴C accumulating in aphids in basal galls, while only supplying 7% to aphids in distal galls. This additional resource available to aphids in basal galls can account for the 65% increase in progeny produced in basal galls compared to galls located more distally on the leaf and limited to the galled leaf as a food resource. Developing furits also act as skins and compete with aphid-induced sinks for food supply. Aphid success in producing galls was increased 31% when surrounding female catkins were removed.     

Defence,growth and nutrient allocation in the tropical shrub Bauhinia brevipes (Leguminosae)

Herbivory in natural communities can be high, reducing the growth and reproduction of individual plants. To diminish the impact of herbivores, plants use a variety of defensive mechanisms. It is now recognized that the plant world is characterized by a proliferation of secondary metabolites, but models of the evolution of defence strategies assume that plants are constrained by finite resources, and that defences are costly. In the present study, the relationships between defence, growth and nutrient concentration in Bauhinia brevipes Vog. (Leguminosae) in south‐eastern Brazil were investigated. Patterns of herbivory and defence were examined for 170 B. brevipes individuals in 1997 and 1998. Leaf age influenced tannin concentration and herbivore attack on B. brevipes. Mature leaves had higher concentrations of tannins than young leaves, and young leaves were approximately 60% more likely to be attacked than mature leaves. Carbon and nitrogen content explained 36% of the variation in the percentage of attacked leaves in B. brevipes (y = 16.29 – 0.31 C + 0.25 N; r² = 0.36; F = 9.56; P < 0.0001). Tannins acted as defensive compounds on B. brevipes leaves against free‐feeding herbivores, but no clear pattern was observed against galling herbivores. The amounts of nitrogen, carbon, potassium, and aluminium explained approximately 69% of the variation in tannin concentration on B. brevipes leaves (y = 0.62 – 0.16 N – 0.36 K – 0.26 Al + 0.74 C; r² = 0.69; F = 21.18; P < 0.0001). We found an inverse relationship between shoot growth rates and tannin concentration in B. brevipes, indicating that the production of tannins may be costly, therefore offering some evidence for the trade‐off hypothesis. Nevertheless, the weak relationship observed between herbivore damage and tannin concentration indicates that other factors are also involved with herbivore pressure.     

The applicability of the plant vigor and resource regulation hypotheses in explaining Epiblema gall moth–Parthenium weed interactions

A basic question in insect–plant interactions is whether the insects respond to, or regulate plant traits, or a complex mixture of the two. The relative importance of the directions of the influence in insect–plant interactions has therefore been articulated through both the plant vigor hypothesis (PVH) and the resource regulation hypothesis (RRH). This study tested the applicability of these hypotheses in explaining the interactions between Parthenium hysterophorus L. (Asteraceae) and its stem‐galling moth, Epiblema strenuana Walker (Lepidoptera: Tortricidae). Parthenium plants exposed to galling were sampled at three sites in north Queensland, Australia, over a 2‐year period, and the relationship between gall abundance and plant vigor (plant height, biomass, flowers per plant, and branches per plant) was studied. To test the predictions of PVH and RRH, the vigor of parthenium plants protected from galling using insecticides was compared to galled plants and plants that escaped from galling. The vigor of ungalled plants was less than the vigor of galled plants. The higher plant vigor in galled plants was not due to galling, as was evident from insecticide exclusion trials. The insect seemed to preferentially gall the more vigorous plants. These findings support the predictions of the PVH and are contrary to those of RRH. Since gall abundance is linked to plant vigor, galling may have only a limited impact on the vigor of parthenium. This has implications for weed biological control. If the objective of biological control is to regulate the population of a plant by a galling insect, a preference for more vigorous plants by the insect is likely to limit its ability to regulate plant populations. This may explain the paucity of successes against biocontrol of annual weeds using gall insects.     

Leafy branches of Gangamopteris from the Gzhelian–Asselian of westernmost Gondwana

The first reconstructions of glossopterids interpreted this Gondwanan group as arborescent, deciduous plants, with leaves and reproductive structures inserted on short shoots, which were arranged on long branches. The leaves are its most abundant organ in the fossil record, but they are mainly found isolated. The arrangement of the leaves as terminal whorls or tight spirals has been the most accepted phyllotaxis hypothesis. The few examples of leaf impressions preserved in connection with axes correspond mainly to leaves of Glossopteris Brongniart, and mostly without clear evidence of the type of insertion. Several specimens of Gangamopteris McCoy leaves attached to axes from the Bajo de Véliz Formation (Latest Carboniferous–Earliest Cisuralian) facilitate reconstruction of the foliar arrangement of the genus, to date known mostly from isolated leaves. The available evidence from the new specimens confirms a variation from well-spaced to dense helical insertion of the leaves without forming true whorls, and discards the early notion that they were mainly clustered apically on short shoots in a similar manner to the extant Ginkgoales.     

Physiological,biochemical, and fluorescence parameters of senescing sugar beet leaves in the vegetative phase of growth

The phase of vegetative growth of sugar beet (Beta vulgaris L., single-sprout form) was conditionally subdivided into four periods according to leaf number and size (including already withered leaves): (A) 8 ± 1 weeks after seedling emergence (wase) (5–7 leaves); (B) 11 ± 1 wase (10–12 leaves); (C) 14 wase (13–15 leaves); (D) 15 wase (15–18 leaves). It took each next leaf about 1 week to come into view. In the course of leaf senescence, palisade parenchyma became less ordered; cells, vacuoles, and intercellular spaces expanded; leaf area and thickness increased. Chloroplasts became swollen, starch grains (and later osmiophilic globules) accumulated and degraded. In every growth period, the highest levels of soluble carbohydrates (sCH), chlorophyll (Chl (a + b)), soluble protein, and the highest activities of rubisco and soluble carboanhydrase usually preceded the full leaf expansion. In unfolded leaves at the growth period B, the maximum values of biochemical characteristics were as a rule higher than at the growth periods A and C and especially D. The only exception was Chl (a + b) with its peak level somewhat increased with plant age. Occurrence of peak values of individual characteristics depended on plant growth period. These characteristics started diminishing asynchronously, with a minimum in old operational leaves. Only the sCH content in the leaves at the periods C and D was stable. Changes in quantum yield at PSII and nonphotochemical fluorescence quenching reflected the age-associated differences in biochemical characteristics. The results are discussed in the light of the idea that leaf senescence is a normal stage of development directly related to the changes in source-sink relations. Biochemically, this stage comprises the degradation of temporarily stored products and partial utilization of the breakdown products for maintenance of the growth of newly formed leaves and root.