Factors underlying the late seasonal appearance of the lepidopterous leaf-mining guild on oak

Abstract. 1. Most lepidopterous leaf mining species found on the oak Quercus robur in Britain develop in summer. At this time of year, externally feeding caterpillars remove little leaf area since most of these free living folivores are predominantly spring feeders.
2. I forced Phyllonorycter harrisella (L.) miners to oviposit in spring, then exposed developing larvae to a wide range of leaf damage levels.
3. Leaf miner survivorship and mean female pupal weight were significantly greater in the experimental spring generation on undamaged oak leaves, this being when oak foliage is of the highest nutritional quality.
4. Leaf miner survivorship in all generations is correlated with natural leaf damage levels. Experimental leaf damage also reduces miner survivorship.
5. Leaf damage reduces miner survivorship by increasing the probability of larval death due to wound induced responses.
6. The role of asymmetrical competitive interactions between caterpillars and leaf-miners in determining the late seasonal appearance of miners is discussed. It is suggested that wound induced responses may play an important part in the structuring of phytophagous insect communities.     

Spatial, bottom-up, and top-down effects on the abundance of a leaf miner

Many plant and animal species have higher densities at the centre of their distribution, with a gradual decline in abundance towards the edge of the range, though reasons for this pattern is not well known. We examined the abundance of the leaf miner Cameraria sp. nova over the range of its host plant Quercus myrtifolia in Florida and addressed how bottom-up and top-down factors varied over its whole distribution. Leaf miner densities, plant quality and natural enemy effects on mine survivorship were evaluated in 40 sites and spatially structured models were used to determine the effects of spatial location on the abundance of Cameraria and effects of both bottom-up (tannin concentration, foliar nitrogen, soil nitrogen, and leaf area) and top-down factors (larval parasitism and predation) on abundance and survivorship. Cameraria mines were, on average, three times more abundant on edge/coastal sites compared to centre/inland sites and did not support the hypothesis of higher abundance on the centre of the distribution. Differences in plant quality, larval parasitism and successful emergence of mines on edge versus central sites might be partially responsible for this finding. A trend surface equation with latitude and longitude combined explained almost 52% of the variation in Cameraria density and a trend surface map also revealed peaks of Cameraria abundance on the edges of the plant distribution. Correlograms also indicated a significant spatial structure of Cameraria as mines were positively spatially autocorrelated at small distances (≈122 km). Partial regression analyses indicated that 69% of the variation in Cameraria abundance was explained by the effects of latitude, longitude, elevation and percentage of foliar nitrogen. Our results indicated that variation in Cameraria abundance was mostly explained by spatial position and significant effects of bottom-up and top-down factors were not detected in our large-scale study.     

Patterns of intra- and interspecific association in leaf-mining insects on three oak host species

Abstract. 1. We determined mortality and distributional patterns of leaf miners on three oak host species (Quercus falcata, Q.nigra and Q.hemisphaerica) in northern Florida, U.S.A.
2. Patterns of intra- and interspecific occurrence within leaves, and mortality of five most abundant leaf miner species were analysed as a test of competition.
3. Miners co-occurred on leaves more often that expected by chance (P<0.05) in six of ten possible species combinations and log-linear model analysis showed no negative higher-order interactions.
4. All five miner species had highly clumped distributions between leaves (P<0.01).
5. Leaf miner survival was less than expected for four of five species when co-occurring on leaves with conspecifics than when mining with heterospecifics or alone (P<0.05).
6. We conclude that interspecific competition is unapparent within this leaf miner guild and that intraspecific competition occurs in four of the five major leaf miner species. We discuss leaf miner selection of common leaves, perhaps based on chemical/physical leaf characters, as a cause of intra- and interspecific aggregation.     

Elevated CO2 lowers relative and absolute herbivore density across all species of a scrub-oak forest

The unabated increase in global atmospheric CO(2) is expected to induce physiological changes in plants, including reduced foliar nitrogen, which are likely to affect herbivore densities. This study employs a field-based CO(2 )enrichment experiment at Kennedy Space Center, Florida, to examine plant-herbivore (insect) interactions inside eight open-topped chambers with elevated CO(2) (710 ppm) and eight control chambers with ambient CO(2). In elevated CO(2) we found decreased herbivore densities per 100 leaves, especially of leaf miners, across all five plant species we examined: the oak trees Quercus myrtifolia, Q. geminata, and Q. chapmanii, the nitrogen-fixing vine Galactia elliottii and the shrub Vaccinium myrsinites. Both direct and indirect effects of lowered plant nitrogen may influence this decrease in herbivore densities. Direct effects of lowered nitrogen resulted in increased host-plant related death and an increase in compensatory feeding: per capita herbivore leaf consumption in elevated CO(2) was higher than in ambient CO(2). Indirectly, compensatory feeding may have prolonged herbivore development and increased exposure to natural enemies. For all leaf miners we examined, mortality from natural enemies increased in elevated CO(2). These increases in host-plant induced mortality and in attack rates by natural enemies decreased leaf miner survivorship, causing a reduction in leaf miner density per 100 leaves. Despite increased leaf production in elevated CO(2) from the carbon fertilization effect, absolute herbivore abundance per chamber was also reduced in elevated CO(2). Because insects cause premature leaf abscission, we also thought that leaf abscission would be decreased in elevated CO(2). However, for all plant species, leaf abscission was increased in elevated CO(2), suggesting a direct effect of CO(2) on leaf abscission that outweighs the indirect effects of reduced insect densities on leaf abscission.     

Variation in rates of leaf abscission between plants may affect the distribution patterns of sessile insects

Summary At Sand Lake, Leon County, Florida, mines of Stilbosis quadricustatella, a leaf-mining moth, occur on sand live oak trees (Quercus geminata) over a broad range of densities. Some trees have fewer than 2% of their leaves mined (lightly infested), others up to 70% (heavily infested). Similar levels of infestation are maintained on the same trees year after year. There are no significant negative correlations of miner density per tree with any mortality factor that might explain miner preferences for certain trees. Nor is there a positive correlation with host leaf quality as measured by foliar nitrogen or by total or individual amino acid concentrations of host leaves. Egg-transfer experiments showed that larvae from eggs transferred to lightly infested trees were more likely to be killed by leaf abscission than were those that were transferred to and developed on heavily infested trees. This is the first demonstration that variation in rates of leaf abscission could be an important cause of the observed distribution pattern of sessile insects between conspecific host plants.     

Non-random distribution patterns of leaf miners on oak trees

Summary Leaf-mining Stilbosis quadricustatella larvae are distributed non-randomly within leaves of their host plants, sand live oak (Quercus geminata) and water oak (Q. nigra), in north Florida. Fewer mines are found together on the same side of the mid-vein than separated, on opposite sides of the mid-vein. Larvae do not normally cross the mid-vein but create small blotch-like mines along subsidiary veins. Investigations of the usual mortality factors acting on these leaf-miner populations, including competition, parasitism, and predation, revealed no significant differences in these factors between mines separated by the mid-vein and those on the same side of the leaf. However, early leaf abscission, which kills the larvae present in the leaf, occurs significantly more frequently in cases where larvae are clustered on one leaf side. The reasons for this differential leaf abscission are not yet clear.     

Abundance and mortality of leaf miners on artificially shaded Emory oak

ABSTRACT.

• 1 We measured solar radiation reaching ten Quercus emoryi Torr. trees and recorded densities of four leaf-mining insect species on these trees from June until September 1982.
• 2 The measurements showed that densities of two leaf miner species were negatively correlated with solar radiation.
• 3 In a field experiment, polypropylene shade fabric was suspended 1.3 m above four experimental trees to test for effects of reduced sunlight.
• 4 Leaves of experimentally-shaded trees were heavier and contained lower percentages of proteins and gallotannins than leaves of control, sun trees, while per cent foliar monomeric, polymeric, and total phenols, and water content did not differ between sun and shaded trees.
• 5 Two of four leaf miner species had greater densities on experimentally-shaded trees than sun trees. One of these two species experienced lower survivorship on sun trees owing to high levels of death from unknown causes on sun trees.
• 6 Two leaf miner species had greater densities on sun trees. One of these species had higher survivorship on shaded trees owing to high levels of parasitism on sun trees.
• 7 We conclude that even subtle differences in shading influences leaf miner density and mortality; however, the effects of shading vary from positive to negative among leaf miner species.

     

Does low nutritional quality act as a plant defence? An experimental test of the slow‐growth,high‐mortality hypothesis

Abstract.  1. The slow-growth, high-mortality hypothesis was experimentally tested in this study by investigating the effects of plant quality and natural enemies on leaf-miner growth, performance, and survivorship. Two leaf miners ( Acrocercops albinatella and Brachys tesselatus ) occurring on the turkey oak Quercus laevis were studied using a factorial design that manipulated predation/parasitism pressure and plant nutritional quality.
2. Forty trees were randomly divided into four treatments: (1) control plants (nutrients and natural enemies unaltered); (2) nutrients added, natural enemies unaltered; (3) nutrients unaltered, natural enemies reduced; and (4) nutrients added and natural enemies reduced. Water content, leaf toughness, tannin concentration, and foliar nitrogen were quantified monthly for each plant, and mine growth and survivorship were assessed by tracing mines on a 2–3-day interval and by following the fates of 50 mines per species per treatment combination.
3. Fertilised plants exhibited significantly higher amounts of nitrogen, but no significant differences among treatments were observed for water content, leaf toughness, and tannin concentration. These results only partially support the slow-growth, high-mortality hypothesis, as mines were significantly smaller and developed faster on fertilised plants, but neither fertilisation nor natural enemy exclusion significantly affected mine survivorship or mortality caused by natural enemies.     

Neither the devil nor the deep blue sea: larval mortality factors fail to explain the abundance and distribution of Tischeria ekebladella

Abstract.  1. The distribution, abundance and population dynamics of herbivorous insects may be affected by trophic interactions, by abiotic influences, or by intra-specific processes. Relatively little is known about how trophic influences vary across space. Here, we investigate spatial variation in mortality in the oak-feeding leaf miner Tischeria ekebladella as attributable to individual causal agents.
2. Leaf miners were experimentally introduced on 67 trees on an island 5 km² in area in south-western Finland. On each tree, some larvae were protected by a muslin bag, others by a glue barrier around the branch and some left exposed.
3. In the bagged transplants, 78.4% of larvae survived, compared with only 9.6% in the other two treatments. Most of the mortality was because of airborne agents: mortality on branches sheltered by a glue barrier was as high as on fully exposed branch tips.
4. We consider mortality caused by parasitoid wasps to be the main source of larval death and the primary factor driving general patterns of survival. The effects of bird predation and premature leaf abscission were negligible.
5. We detected spatial aggregation in larval survival and parasitism rates at the level of individual trees, but not across the landscape.
6. Spatial variation in overall leaf miner survival, parasitism and leaf abscission does not suffice to explain patterns of incidence and abundance of wild T. ekebladella on experimental trees. Rather, we identify metapopulation dynamics as a likely determinant of the spatial distribution of T. ekebladella in the landscape.     

The potential for interspecific competition between two abundant defoliators on oak: leaf damage and habitat quality

Abstract. 1. The pedunculate oak, Quercus robur L., suffers high annual levels of spring defoliation in Wytham Woods, Oxon. The two major defoliators, Tortrix viridana L. and Operophtera brumata L., sometimes reach high enough densities to completely defoliate trees.
2. Experiments with larvae of both species enclosed on oak trees in Wytham Woods suggest considerable potential for intra- and interspecific competition between T. viridana and O. brumata.
3. T. viridana and O. brumata consume equivalent amounts of leaf tissue per unit caterpillar density. Despite this, O. brumata is the superior competitor, measured by mortality during the larval feeding period.
4. Evidence is presented that T. viridana larvae depend upon the protected microclimate of leaf rolls to maintain water balance, and that leaf damage may disrupt the construction of, and microclimate within, leaf rolls. O. brumata may therefore be competitively superior because it indirectly interferes with the water balance of T. viridana.
5. The importance of interspecific competition in the dynamics of T. viridana and O. brumata populations is discussed. Stochastic mortality factors acting on first instar larvae of both species probably make intense competition a rare event.     

Seed predation by mammals and forest dominance by Quercus oleoides,a tropical lowland oak

Summary Quercus oleoides Cham. and Schlecht is an unusual tree in several respects: it is an oak found in neotropical lowland forests, its distribution is not continuous but ratherdivided into many patches of various sizes, and it is a dominant in all the forests in which it occurs, attaining densities far higher than most species of tropical trees. This density pattern is related to the vulnerability of Q. oleoides acorns to predation by mammals. Observations of agoutis, deer, peccaries, squirrels, pocket mice and other seed consumers in Santa Rosa National Park, Costa Rica, showed that these mammals act only as predators, not dispersers, of Q. oleoides acorns. Experiments which involved placing acorns in deciduous forest where Q. oleoides does not occur, demonstrated that, due to high predation rates, the number of acorns produced by an isolated tree is far too low for adults to replace themselves.In oak forest, on the other hand, where the combined acorn crops of many oaks satiate the seed predators, acorn survivorship until germination is high enough to maintain the population. Furthermore, acorn survivorship in oak forest areas is inversely proportional to the apparent mammal density in those areas. Thus the pattern of forest dominance and patchy distribution is related to positively density-dependent acorn survivorship: where Q. oleoides is the forest dominant, it will survive, but if its density falls to the level typical of tropical trees, it will go locally extinct.     

Oviposition site preference and larval mortality in a leaf-mining moth

Abstract. 1. The univoltine leaf-mining moth, Lithocolletis quercus Ams., is endemic to Israel, where it spends its 10.5 month larval period feeding only in the leaves of Quercus calliprinos Webb.
2. We compared patterns of egg deposition and sources of larval mortality to test whether oviposition patterns and site preferences confer an enhanced likelihood of larval survival.
3. Dominant sources of larval mortality were premature leaf abscission and death from unknown causes, whereas predation, parasitism and intraspecific interference accounted for relatively little larval mortality.
4. Eggs, and thus mines, were aggregated among leaves of host trees even though premature leaf abscission was positively correlated with density of mines per leaf. Interference competition among larvae was the only other density-dependent mortality factor.
5. Oviposition patterns within leaves mitigated the probability of death from larval interference, and probably also from early leaf abscission.
6. Despite these density-dependent mortality factors, overall probability of larval survival to pupation was independent of initial density of mines on a leaf.
7. The long larval period allows synchrony between oviposition flights and times of predictable resource availability.     

The clonal structure of Quercus geminata revealed by conserved microsatellite loci

The scrub oak communities of the southeastern USA may have existed at their present locations for thousands of years. These oaks form suckers, and excavations of root systems suggest that clones may occupy very large areas. Resolution of the clonal nature of scrub oaks is important both to manage the tracts of this ecosystem that remain, and in conducting long-term ecological studies, where the study area must substantially exceed the area occupied by any single clone. Microsatellites were used to determine the genetic diversity of a dominant oak species within a 2-ha long-term experimental site on Merritt Island at the Kennedy Space Center. This area contains a long-term study of the effects of elevated CO2 on the ecosystem. Conservation of seven microsatellite loci, previously identified in the sessile oak, Quercus petraea, was tested in two Florida scrub oak species, Q. geminata and Q. myrtifolia. Sequence analysis revealed that all seven microsatellite loci were conserved in Q. geminata and five loci were conserved in Q. myrtifolia. Six microsatellite loci were polymorphic in Q. geminata and these were subsequently used to investigate the clonal structure of the Q. geminata population. Twenty-one unique combinations of microsatellites, or haplotypes, occurred only once, whereas the remaining 26 individuals belonged to a total of seven different haplotypes. Trees with identical haplotypes were in close proximity, supporting the interpretation that they were clones. The results showed that there is significant genetic diversity within the 2-ha experimental site. Microsatellites provided a powerful and noninvasive tool for distinguishing individual genotypes and determining an adequate area for long-term ecosystem studies.     

Influence of density and salinity on larval development of salt‐adapted and salt‐naïve frog populations

Environmental change and habitat fragmentation will affect population densities for many species. For those species that have locally adapted to persist in changed or stressful habitats, it is uncertain how density dependence will affect adaptive responses. Anurans (frogs and toads) are typically freshwater organisms, but some coastal populations of green treefrogs (Hyla cinerea) have adapted to brackish, coastal wetlands. Tadpoles from coastal populations metamorphose sooner and demonstrate faster growth rates than inland populations when reared solitarily. Although saltwater exposure has adaptively reduced the duration of the larval period for coastal populations, increases in densities during larval development typically increase time to metamorphosis and reduce rates of growth and survival. We test how combined stressors of density and salinity affect larval development between salt‐adapted (“coastal”) and nonsalt‐adapted (“inland”) populations by measuring various developmental and metamorphic phenotypes. We found that increased tadpole density strongly affected coastal and inland tadpole populations similarly. In high‐density treatments, both coastal and inland populations had reduced growth rates, greater exponential decay of growth, a smaller size at metamorphosis, took longer to reach metamorphosis, and had lower survivorship at metamorphosis. Salinity only exaggerated the effects of density on the time to reach metamorphosis and exponential decay of growth. Location of origin affected length at metamorphosis, with coastal tadpoles metamorphosing slightly longer than inland tadpoles across densities and salinities. These findings confirm that density has a strong and central influence on larval development even across divergent populations and habitat types and may mitigate the expression (and therefore detection) of locally adapted phenotypes.     

Leaf ties as colonization sites for forest arthropods: an experimental study

Abstract.  1. Secondary colonization of leaf shelters constructed by caterpillars has been reported from a number of systems. Both the mechanism (larval or adult movement vs. oviposition) and the cues used by arthropods in locating leaf shelters, however, have received little attention.
2. Artificial leaf shelters (i.e. leaf ties or pairs of leaves clipped together to form sandwiches) were constructed on understorey white oak ( Quercus alba L.) trees and the abundance and species composition of arthropods colonizing and ovipositing on leaf pairs was examined in three treatments: occupied leaf ties (containing a leaf-tying caterpillar), unoccupied leaf ties, and non-tied control leaves.
3. The density of arthropods present in the occupied and unoccupied leaf ties after two weeks was seven and four times greater respectively, than non-tied controls. The guild composition of these early colonists differed among treatments, with the highest densities of leaf chewers, scavengers, and predators in occupied ties and the lowest densities in non-tied controls.
4. The densities of all arthropods ovipositing on leaf pairs in the occupied and unoccupied leaf tie treatments were four and three times greater than non-tied controls. Leaf-chewing insects (including leaf-tiers and non-tying inquiline species) and scavengers showed strong oviposition preferences for tied leaves. However, most species of leaf-tying caterpillars and the psocids (Psocoptera) did not distinguish between occupied and unoccupied ties, suggesting that these groups do not use occupancy-related cues in selecting oviposition sites.     

An admixture of Quercus dentata in the coastal ecotype of Q. mongolica var. crispula in northern Hokkaido and genetic and environmental effects on their traits

Journal of Plant Research - In northern Japan, coastal oak forests consist of Quercus dentata (Qd) on the coastal side and Q. mongolica var. crispula (Qc) on the inland side. In the forests of...     

Foliar nitrogen and larval parasitism as determinants of leafminer distribution patterns on Spartina alterniflora

ABSTRACT, 1. In northwest Florida, Hydrellia valida hew (Diptera: Ephydridae) frequently mines the leaves of salt marsh cord, grass, Spartina alterniflora . Larvae and pupae are more commonly found in shoreline plants within 3 m of the sea than in inland plants.
2. Shoreline plants contain over twice as much total foliar nitrogen as inland plants. In transects from inland to shoreline plants, leafminer density is positively correlated with total leaf nitrogen content.
3. We experimentally increased foliar nitrogen content on inland Spartina patches to levels above those of shoreline plants. Leafminer densities on experimental centres remained far below those on control edges.
4. Larval parasitism by Opius sp. (Hymenoptera: Braconidae) and Pteromalus sp. (Hymenoptera: Pteromalidae) reaches over 90%, with parasitism increasing from shoreline to inland plants. We suggest that gradients in leafminer densities are caused by differences in parasitoid abundance.     

The effects of substrate and vertebral number on locomotion in the garter snake Thamnophis elegans

1. Locomotor performance of limbless vertebrates depends on the substrate through which individuals move and may result in selection on vertebral number in different habitats. To evaluate the effect of push-point density on snake locomotion, the density of vegetation and other potential push-points was quantified at two sites in California (coastal and inland), where conspecific snakes differed greatly in vertebral number (230 and 256 average total vertebrae, respectively; Arnold 1988). The coastal site had significantly higher push-point densities than the inland site.
2. Five experimental push-point densities that fell within the natural range of push-point densities were employed in laboratory trials of juvenile snake locomotion. Density of push-points significantly affected both crawling speed and head-to-tail distance (HTD), an indirect measure of lateral bending. The fastest speed was achieved at an intermediate push-point density. The shortest HTD occurred when snakes moved through the lowest push-point density.
3. Sex, total number of vertebrae and total length significantly affected HTD, regardless of push-point density. Snakes with relatively more vertebrae had a shorter HTD, suggesting they were able to achieve greater lateral bending than snakes with fewer vertebrae. Coastal and inland populations did not differ in HTD during locomotion.
4. Numbers of body and tail vertebrae significantly influenced speed at different push-point densities. In general, snakes with more body vertebrae were slower than those with fewer, while snakes with more tail vertebrae were faster than those with fewer. Snakes of greater total length were faster at all densities. Coastal snakes crawled faster than inland snakes at all push-point densities.     

Relation of ramet size to acorn production in five oak species of xeric upland habitats in south-central Florida

This study examined variation in two components of acorn production. Percentage of bearing ramets (stems) and number of acorns per bearing ramet were examined in five clonal oaks in three xeric habitats of south-central peninsular Florida in relation to ramet size within and between species and vegetative associations. Counts of acorns on two white oaks (Quercus chapmanii and Q. geminata) and three red oaks (Q. inopina, Q. laevis, and Q. myrtifolia) were conducted annually from 1969 to 1996 (except in 1991) on permanent grids in southern ridge sandhill, sand pine scrub, and scrubby flatwoods associations at the Archbold Biological Station, Florida, USA. Percentage of bearing individuals and mean number of acorns per bearing individual increased with increasing ramet size for all species across all vegetation associations. However, in Q. geminata and Q. myrtifolia, acorn production declined in the largest size class (>3.2 m), implying that larger individuals of these clonal species may become senescent. All oak species in sand pine scrub, which had a nearly closed overstory, had lower frequencies of bearing oaks and mean numbers of acorns compared with similar-sized individuals of the same species in the more open-canopied southern ridge sandhill and scrubby flatwoods associations, suggesting light limitation. The annual production of acorns by a given oak species was correlated across vegetative associations and annual acorn production of oak species was correlated for species within the same section. Intermediate-size class oaks contributed the most acorns per unit area, suggesting that stands managed with short fire-return times will provide fewer acorns to wildlife than stands managed to produce more variable distributions of oak size classes. However, our study suggests that long-unburned stands, such as those studied here, will maintain relatively constant levels of acorn production as a consequence of ramet replacement within the clones of these shrubby oaks to create a variable distribution of size classes. Of the oak species studied, Q. myrtifolia had the highest acorn production and the smallest acorns, while Q. laevis had the lowest acorn production and the largest acorns, suggesting an allocation trade-off between acorn numbers and size.     

The causes of over wintering mortality of Phyllonorycter on Quercus robur

Abstract. 1. An experiment was performed to examine the relationship between overwintering mortality rates and physical conditions in the leaf litter for Phyllonorycter spp. on Quercus robur .
2. Analyses also were performed to determine if survival and mortality rates vary with leaf size and the position of a leaf mine within a leaf.
3. The results indicate that survival rates are independent of variation in physical conditions in the leaf litter, as well as leaf size and the position of a mine on a leaf.
4. Mortality from leaf decomposition was higher at the bottom of the litter pack, and it may compensate for reduced mortality from predation, crushing, desiccation, and fungal attack at the level.
5. Rates of parasitism were significantly higher on small than on large leaves.