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Premise: Life span varies greatly across plants, with some species being capable of extreme longevity. Yet even long-lived individuals are susceptible to climatic events, fire, and other challenges. We examined rare mortality events and their causes in two long-lived palmettos over four decades., Methods: We monitored the survival of the clonal Serenoa repens and non-clonal, Florida-endemic Sabal etonia from 1981 to 2022 in four habitats along an elevational gradient within the globally imperiled Florida scrub ecosystem. We considered several challenges to palmetto survival, including extreme fires, shading due to lack of fire, droughts, periods of high precipitation, and possible pathogens., Results: Survival of palmettos was remarkably high, and mortality was infrequent (Serenoa: cumulative, 5.7%; annualized, 0%-0.68%; Sabal: cumulative, 3.5%; annualized, 0%-0.43%). Mortality was highest in higher-elevation habitats with greater soil drainage, and smaller palmettos were more likely to die. When subjected to extreme fire, Serenoa suffered greater mortality than Sabal. Mortality in long-unburned habitats with increased shading rivaled that which occurred with extreme fire. There was no evidence of mortality due to lethal bronzing palm disease., Conclusions: Both palmettos had exceptionally low mortality rates, which, coupled with earlier work showing slow rates of transition from seedling to adult and remarkable adult longevity, suggest notably low rates of population turnover. Observed mortality in long-unburned habitats suggests the importance of fire-management planning with prescription burning. Lengthy age to reproduction and/or dependency on clonal propagation limits migration or genetic adaptation to altered conditions caused by climate change., (© 2023 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

Twenty nine new species of cynipid oak gall wasps from the Nearctic region (America north of Mexico) are described: Andricus archboldi Melika Abrahamson, sp. nov., A. catalinensis Melika, Nicholls Stone, sp. nov., A. chapmanii Melika Abrahamson, sp. nov., A. chiricahuensis Melika, Nicholls Stone, sp. nov., A. coconinoensis Melika, Nicholls Stone, sp. nov., A. columbiensis Melika, Nicholls Stone, sp. nov., A. cooki Melika, Nicholls Stone, sp. nov., A. fitzpatricki Melika Abrahamson, sp. nov., A. highlandensis Melika, Nicholls Stone, sp. nov., A. mellificus Nicholls, Stone Melika, sp. nov., A. menkei Melika Abrahamson, sp. nov., A. mogollonensis Melika, Nicholls Stone, sp. nov., A. nichollsi Melika Stone, sp. nov., A. schickae Nicholls, Melika Stone, sp. nov., A. torreyaensis Melika Abrahamson, sp. nov., A. williami Melika, Nicholls Stone, sp. nov., Antron lovellae Melika, Nicholls Stone, sp. nov., A.tomkursari Melika, Nicholls Stone, sp. nov., Dryocosmus archboldi Melika Abrahamson, sp. nov., Loxaulus virginianae Melika Buss, sp. nov., Neuroterus alexandrae Nicholls Melika, sp. nov., N. aliceae Melika, Nicholls Stone, sp. nov., N. bussae Melika Nicholls, sp. nov., N. oblongifoliae Nicholls, Stone Melika, sp. nov., N. quaili Melika, Nicholls Stone, sp. nov., N. rosieae Melika, Nicholls Stone, sp. nov., N. stonei Melika Nicholls, sp. nov., Zapatella abrahamsoni Melika, sp. nov., Z. brooksvillei Melika Abrahamson, sp. nov.. Alternate asexual and sexual generations are described for four species, Andricus archboldi Melika Abrahamson, sp. nov., A. fitzpatricki Melika Abrahamson, sp. nov., A. schickae Nicholls, Melika Stone, sp. nov., Neuroterus aliceae Melika, Nicholls Stone, sp. nov.. Descriptions, diagnoses, plus information on biology and host associations are given for all new species. All taxa are supported by morphological data; matching of generations is established using DNA sequence data. We also demonstrate that Neuroterus niger var. alimas Kinsey should be considered as a nomen dubium.

Plant populations frequently maintain submaximal levels of resistance to natural enemies, even in the presence of substantial genetic variation for resistance. Identifying constraints on the evolution of increased resistance has been a major goal of researchers of plant-herbivore interactions. In a glasshouse study, we measured relative costs and benefits of resistance of tall goldenrod (Solidago altissima) to the gall-inducing tephritid Eurosta solidaginis. We exposed multiple ramets of 26 goldenrod genets to nutrient or shade stress and to oviposition by E. solidaginis. The presence of a gall cost a ramet an average of 1743 seeds, but the cost differed 10-fold across environments. Plant genets varied widely for an induced 'hypersensitive' response in which meristem cells become necrotic and kill E. solidaginis hatchlings before gall induction. There was no evidence that this highly effective resistance trait carried an allocation cost. However, the response carried a risk of autotoxicity, as necrosis killed the apex of 37% of the ungalled ramets. On average, a damaged apex cost each ramet 5015 seeds. Autotoxicity may constrain the resistance of S. altissima to an intermediate level, and variation in environmental conditions may alter the relative costs and benefits of resistance and tolerance, thus maintaining genetic variation within goldenrod populations., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

The lack of effective tools has hampered out ability to assess the size, growth and ages of clonal plants. With Serenoa repens (saw palmetto) as a model, we introduce a novel analytical framework that integrates DNA fingerprinting and mathematical modelling to simulate growth and estimate ages of clonal plants. We also demonstrate the application of such life-history information of clonal plants to provide insight into management plans. Serenoa is an ecologically important foundation species in many Southeastern United States ecosystems; yet, many land managers consider Serenoa a troublesome invasive plant. Accordingly, management plans have been developed to reduce or eliminate Serenoa with little understanding of its life history. Using Amplified Fragment Length Polymorphisms, we genotyped 263 Serenoa and 134 Sabal etonia (a sympatric non-clonal palmetto) samples collected from a 20 × 20 m study plot in Florida scrub. Sabal samples were used to assign small field-unidentifiable palmettos to Serenoa or Sabal and also as a negative control for clone detection. We then mathematically modelled clonal networks to estimate genet ages. Our results suggest that Serenoa predominantly propagate via vegetative sprouts and 10,000-year-old genets may be common, while showing no evidence of clone formation by Sabal. The results of this and our previous studies suggest that: (i) Serenoa has been part of scrub associations for thousands of years, (ii) Serenoa invasion are unlikely and (ii) once Serenoa is eliminated from local communities, its restoration will be difficult. Reevaluation of the current management tools and plans is an urgent task., (© 2011 Blackwell Publishing Ltd.)

Herbivores are among the most pervasive selective forces acting on plants, and the number of plant chemicals that presumably evolved for defense against herbivory is immense. In contrast, biologists are only beginning to appreciate the important roles that architectural traits can play in antiherbivore defense. One putative architectural-resistance trait is the nodding stem apex of some goldenrods (Solidago; Asteraceae). Individuals of S. altissima genets that undergo temporary nodding in the late spring (i.e., "candy-cane" ramets) have been shown to be more resistant than individuals of erect-stemmed genets to certain apex-attacking herbivores. We tested the hypothesis that the greater resistance of candy-cane ramets is accomplished by the ramets' "ducking" from the herbivores. In a greenhouse experiment, nodding candy-cane ramets were significantly more resistant to oviposition by the gall-inducing fly Eurosta solidaginis than were ramets of the same genets that had been experimentally straightened. The straightened candy-cane stems were just as susceptible to ovipositions as were ramets of erect-stemmed genets. Thus, ducking indeed appears to confer a resistance advantage to candy-cane genets of S. altissima.

Host-race formation is promoted by genetic trade-offs in the ability of herbivores to use alternate hosts, including trade-offs due to differential timing of host-plant availability. We examined the role of phenology in limiting host-plant use in the goldenrod gall fly (Eurosta solidaginis) by determining: (1) whether phenology limits alternate host use, leading to a trade-off that could cause divergent selection on Eurosta emergence time and (2) whether Eurosta has the genetic capacity to respond to such selection in the face of existing environmental variation. Experiments demonstrated that oviposition and gall induction on the alternate host, Solidago canadensis, were the highest on young plants, whereas the highest levels of gall induction on the normal host, Solidago gigantea, occurred on intermediate-age plants. These findings indicate a phenological trade-off for host-plant use that sets up the possibility of divergent selection on emergence time. Heritability, estimated by parent-offspring regression, indicated that host-race formation is impeded by the amount of genetic variation, relative to environmental, for emergence time.

Host-associated differentiation (HAD) is considered a step towards ecological speciation and an important mechanism promoting diversification in phytophagous insects. Although the number of documented cases of HAD is increasing, these still represent only a small fraction of species and feeding guilds among phytophagous insects, and most reports are based on a single type of evidence. Here we employ a comprehensive approach to present behavioural, morphological, ecological and genetic evidence for the occurrence of HAD in the gall midge Dasineura folliculi (Diptera: Cecidomyiidae) on two sympatric species of goldenrods (Solidago rugosa and S. gigantea). Controlled experiments revealed assortative mating and strong oviposition fidelity for the natal-host species. Analysis of mitochondrial DNA showed an amount of genetic divergence between the two host-associated populations compatible with cryptic species rather than host races. Lower levels of within-host genetic divergence, gall development and natural-enemy attack in the S. gigantea population suggest this is the derived host.

Many populations of goldenrod show a peculiar, genetically controlled stem dimorphism. In Solidago altissima, for instance, while most stems are erect, a sizable minority (the "candy-cane" stems) nod at the apex during growth. We used data from three studies to test the hypothesis that this candy-cane growth form confers resistance to herbivory. In a controlled growth trial, we showed that nodding is a temporary phenomenon that coincides with the oviposition period of at least two common apex-attacking herbivores: the tephritid galler Eurosta solidaginis and the gall midge Rhopalomyia solidaginis. In a large field survey, stems of candy-cane genets were only half as likely to be ovipunctured by E. solidaginis. In a common-garden study, candy-cane stems were less likely to be ovipunctured by E. solidaginis, and they were galled only half as often by R. solidaginis as erect stems. These results suggest that the candy-cane stems of goldenrod possess a resistance strategy that allows them to essentially duck and hide from certain herbivores.

Apical meristem damage (AMD) is a common result of herbivory. AMD can have dramatically variable effects on plant architecture and fitness, ranging from a total loss of reproductive capacity to overcompensation. We explored the influence of environmental stresses and meristem limitation on tolerance of AMD by applying the limiting resource model (LRM) of plant tolerance to 17 previously published studies and a new empirical study on Solidago altissima. In the S. altissima experiment, AMD released axillary meristems from apical dominance, and fertilizer addition enabled plants to take full advantage of the lateral branches. AMD caused a 58% reduction in seed production in nutrient-stressed plants but only a 6% reduction in seed production in fertilized plants. In 12 of the 18 studies reviewed, tolerance was greater in the high-resource (or low-competition) treatment; in two, tolerance was greater in the low-resource treatment; and in four, resource level did not affect tolerance of AMD. The results of 15 studies (83%) were consistent with LRM predictions. Overcompensation was observed in six studies, and it occurred only in the high-resource treatments in five of these studies, as would be expected from applying the LRM.

A main tenet of sex-allocation theory is that environmental stress should lead to increased maleness because reproducing through pollen is generally cheaper than producing fruits and seeds. Though this prediction has held for many species, it has been little tested for gynomonoecious plants, in which individuals produce both female and perfect flowers. We exposed eight ramets of each of 22 genets of a gynomonoecious goldenrod, Solidago altissima (Asteraceae), to a factorial combination of nutrient stress and herbivory by the gall-inducer Eurosta solidaginis (Tephritidae). Nutrient stress alone increased relative femaleness: Stressed ramets produced fewer flowers total and a higher ratio of ray (female) flowers to disk (perfect) flowers. Galling caused no change in fertilized ramets, but the combination of nutrient stress and galling caused an increase in relative maleness: Nutrient-stressed, galled ramets produced fewer flowers total and had a higher disk to ray ratio. In addition to being phenotypically plastic, floral-sex ratio had a great deal of genetic variation, with a broad-sense heritability of 0.68. While the floral-sex ratio responses of gynomonoecious plants may be more complicated than for plants of other breeding systems, they offer the potential to test and refine the already rich body of sex-allocation theory.

Evidence of poor correspondence between an insect herbivore's oviposition preferences and the performance of its offspring has generally been attributed either to maladaptive behavior of the insect mother or inadequate measurement by the researcher. In contrast, we hypothesize that many cases of "bad mothers" in herbivores may be a byproduct of the hierarchical way natural selection works on resistance in host plants. Epistatic selection on the components of resistance (i.e., antixenosis and antibiosis) may generate negative genetic correlations between the resistance components, which could counteract the efforts of herbivores to oviposit on the best hosts for the performance of their offspring. In common garden and greenhouse experiments, we measured aspects of antixenosis and antibiosis resistance in 26 genets of tall goldenrod, Solidago altissima, against two common herbivores: the gall-inducing fly Eurosta solidaginis and the spittlebug Philaenus spumarius. Goldenrod antixenosis and antibiosis were positively correlated against E. solidaginis and negatively correlated against P. spumarius. Analogously, population-wide preference-performance correlations were positive for the gall flies and negative for the spittlebugs. Several natural history differences between the two insects could make gall flies better mothers, including better synchrony of the phenologies of the flies and the host plant, the much narrower host range of the gall flies than the spittlebugs, and the more sedentary lifestyle of the gall fly larvae than the spittlebug nymphs. If these results are typical in nature, then negative genetic correlations in antixenosis and antibiosis in plants may often result in zero or negative population-wide correlations between preference and performance in herbivores, and thus may be an important reason why herbivorous insects often appear to be bad mothers.

Parasitic species can dramatically alter host traits. Some of these parasite-induced changes can be considered adaptive manipulations that benefit the parasites. Gall-inducing insects are parasites well known for their ability to alter host-plant morphology and physiology, including the distribution of plant defensive compounds. Here it was investigated whether gall-inducing species alter indirect plant defenses, involving the release of volatile compounds that are attractive to foraging natural enemies. Using field and factorial laboratory experiments, volatile production by goldenrod (Solidago altissima) plants was examined in response to attack by two gall-inducing species, the tephritid fly Eurosta solidaginis and the gelechiid moth Gnorimoschema gallaesolidaginis, as well as the meadow spittlebug, Philaenus spumarius, and the generalist caterpillar Heliothis virescens. Heliothis virescens elicited strong indirect defensive responses from S. altissima, but the gall-inducing species and spittlebugs did not. More significantly, infestation by E. solidaginis appeared to suppress volatile responses to subsequent attack by the generalist caterpillar. The extensive control that E. solidaginis apparently exerts over host-plant defense responses may reduce the predation risk for the gall inducer and the subsequent herbivore, and could influence community-level dynamics, including the distribution of herbivorous insect species associated with S. altissima parasitized by E. solidaginis.

Plants of nutrient-poor, arid environments often have leaf traits that include small size, sclerophylly, long life span, low nutrient concentration, and low photosynthetic rate. Hence, the success of two large-leaved palmettos in peninsular Florida's seasonally xeric, nutrient-impoverished uplands seems anomalous, given that their leaves are orders of magnitude larger than the leaves of sympatric species. An examination of a 16-yr data set of leaf traits and leaf life spans across four vegetative associations differing in available light showed that Serenoa repens and Sabal etonia had low rates of leaf production coupled with long leaf life spans reaching 3.5 yr in heavily shaded plants. The adaptation of these palmettos to xeric, nutrient-poor habitats has generated dwarf statures, diminished leaf sizes and numbers, increased leaf life spans, and reduced rates of leaf production relative to other palms and congeners of more mesic sites. Leaf and petiole size, plant leaf canopy area, and leaf life span increased in both palmettos with decreasing available light, helping to compensate for reduced photosynthetic rates under shaded conditions and for the high leaf construction costs of the large, thick palmetto leaves. Large leaf size in these palmettos, likely due to phylogenetic conservatism, is compensated by other leaf traits (e.g., heavily cutinized epidermises, thick laminas) that increase survival in seasonally xeric, nutrient-impoverished environments.

Although it is widely acknowledged that a plant's tolerance of herbivore damage depends on resource availability in the plant's environment, there is no consensus on whether higher resource levels lead to greater or to lower tolerance. The prevailing model, the compensatory continuum hypothesis (CCH), predicts that tolerance of herbivory should be greater in high-resource or low-competition conditions. The main rival hypothesis, the growth rate model (GRM), makes the opposite prediction: tolerance of herbivory should be greater in more stressful conditions. The tolerance predictions of a recently introduced model, the limiting resource model (LRM), are more flexible and depend on the type of resource and herbivore under consideration. We reviewed 48 studies (from 40 published articles) of plant tolerance of leaf damage in conditions differing in levels of light, inorganic nutrients, water stress, or competition. The results of 31%, 48%, and 95% of the studies were consistent with the predictions of the CCH, GRM, and LRM, respectively. Thus, by considering which resource is primarily affected by herbivory and which resource is limiting a plant's fitness, the LRM offers a substantial advance in predicting how tolerance will be affected by environmental differences in resource availability.

Host-race formation remains controversial as a source of herbivorous insect diversity, and examples of host races are still fairly scarce. In this study, analysis of five enzyme loci in the ostensibly generalist tumbling flower beetle Mordellistena convicta (Coleoptera: Mordellidae) revealed hidden host-plant and plant-organ related genetic differentiation. Mordellistena convicta turned out to be a complex of cryptomorphic species, each with fewer hosts than the nominal species. These cryptic species, in turn, were divided into taxa that showed host-race characteristics: samples from different host plants and organs exhibited (1) genetic indications of partial reproductive isolation, (2) differences in size and emergence timing that suggested divergent host-related selection, and (3) among-host selective differences in mortality from parasitoids. Host-race formation in M. convicta, which has a somewhat different life history from the well-studied host races, enlarges the group of insects considered likely to undergo this process. The widespread sympatry of the M. convicta species complex, along with its spectrum of host-correlated genetic differentiation, suggests that these host specialist taxa developed in sympatry.

Reproductive output by the Florida-endemic scrub hickory (Carya floridana Sargent) was studied over a 28-yr period in three south-central Florida vegetation associations: southern ridge sandhill, sand pine scrub, and scrubby flatwoods. The objectives were to describe multi-annual patterns of variation in nut production, identify factors involved in this variation, and investigate differences in patterns among associations. Peaks (higher values bracketed by lower values) in nut production occurred in 7 yr in sandhill and scrub and 8 yr in scrubby flatwoods during the 22-yr period for which we had continuous data. Of the total of 22 peaks in the three associations combined, 17 occurred at intervals of 2 or 3 yr, and peaks occurred in the same years in 18 of the 22 cases. Periodicities of nut production generated by spectral analyses (Fourier transforms) generally agreed with the observed peaks. Numbers of nuts per bearing ramet, proportion of ramets bearing nuts, ramet height, and light availability were positively correlated with nut production. Weather variables, specifically winter rainfall and minimum spring temperatures, accounted for a total of about one-quarter to one-half of the variance in nut production depending on the vegetation association. Following a prescribed fire in a sandhill plot, scrub hickory quickly regained fruit production, but over a 5-yr period following the fire nut production by ramets in the largest size class was reduced compared with the unburned control plot.

Host shifts and the formation of insect-host races are likely common processes in the speciation of herbivorous insects. The interactions of goldenrods Solidago (Compositae), the gall fly Eurosta solidaginis (Diptera: Tephritidae) and the beetle Mordellistena convicta (Coleoptera: Mordellidae) provide behavioural, ecological and genetic evidence of host races that may represent incipient species forming via sympatric speciation. We summarize evidence for Eurosta host races and show that M. convicta has radiated from goldenrod stems to Eurosta galls to form host-part races and, having exploited the galler's host shift, has begun to differentiate into host races within galls. Thus, host-race formation has occurred in two interacting, but unrelated organisms representing two trophic levels, resulting in 'sequential radiation' (escalation of biodiversity up the trophic system). Distributions of host races and their behavioural isolating mechanisms suggest sympatric differentiation. Such differentiation suggests host-race formation and subsequent speciation may be an important source of biodiversity.

Host-plant association data, gathered from field surveys conducted throughout Florida and from the literature, were used to identify the specificity of cynipid gall inducers to one or more of six Quercus species that occur at Archbold Biological Station, Lake Placid, Florida, USA, including the red oaks Q. laevis, Q. myrtifolia, and Q. inopina, and the white oaks Q. chapmanii, Q. geminata, and Q. minima. Quercus myrtifolia had the highest cynipid richness and diversity (37 cynipid species, Shannon H' = 3.61, Simpson's D = 0.97), followed by Q. chapmanii, Q. laevis, Q. inopina, Q. geminata, and finally Q. minima (10 species, H' = 2.30, D = 0.90). All cynipid species showed strong fidelity to a particular host plant or a restricted set of host plants. An ordination of gall-wasp host associations indicated that the cynipid communities of each oak species were distinct and specific to a given oak species. Leaf samples taken from each oak species were analyzed for condensed and hydrolyzable tannins, total phenolics, lignin, cellulose and hemicellulose, nitrogen, and carbon. All of these chemical traits, with the exception of carbon, differed by oak species, and the differences were strongly correlated with the axes of the cynipid-species ordination. These results suggest that gall-wasp occurrence is influenced by oak chemistry and imply that experimental studies of cynipid gall inducers that examine host-plant chemistry and female oviposition choice and larval performance will yield useful insights.

We show that a predator, the tumbling flower beetle Mordellistena convicta (Coleoptera: Mordellidae), has formed host races in response to a host-plant shift and subsequent host-race formation by its prey, the gall-inducing fly Eurosta solidaginis (Diptera: Tephritidae). This fly has formed two host races, one that induces stem galls on the ancestral host plant, Solidago altissima (Compositae), and another that induces stem galls on the closely related S. gigantea. We found that subpopulations of M. convicta that attack E. solidaginis galls on the different host plants have significantly different emergence times and, although slight, these allochronic differences are consistent across a range of temperatures. More importantly, we found that beetles assortatively mate according to their natal host plants, and female M. convicta preferentially attack and/or their offspring have higher survival in galls on natal host plants. Our data suggest that subpopulations of M. convicta that attack E. solidaginis galls on S. altissima and S. gigantea have formed host races. This is one of the first studies to demonstrate that a host shift and subsequent host-race formation by an herbivorous insect may have resulted in subsequent diversification by one of its natural enemies.

We examined post-fire recovery of two components of acorn production (percentage of bearing ramets [stems] and number of acorns per bearing ramet) for four species of oaks in southern ridge sandhill vegetation in south-central peninsular Florida. Annual counts of acorns on two white oaks (Quercus chapmanii and Q. geminata) and two red oaks (Q. laevis and Q. myrtifolia) were conducted annually (except in 1991) on two 2.7-ha grids from 1969 to 1998. A prescribed burn was conducted on one of the grids in May 1993. Newly sprouted ramets of both white oaks produced acorns during the first year following the fire, whereas red oaks required 3 yr (Q. myrtifolia) or 4 yr (Q. laevis) to produce acorns. The difference in the timing of post-fire acorn production between the white and red oak species reflected the difference in the number of years from flower bud initiation to mature acorns in the two groups, with the additional year-long lag in Q. laevis probably attributable to the fact that it is typically a tree rather than a shrub species. The data suggested that percentage of bearing ramets in the smallest size class of the two white oak species was markedly lower in the burned than unburned grid in the first year of post-fire acorn production and higher in the fifth year, but these trends were not evident for the red oaks. Among all four species, differences between mean number of acorns in burned and unburned grids were significant in only two cases (the largest size class of both white oak species in the fifth year). There was no evidence of recruitment from acorns on the burned grid, possibly due to the rapid redevelopment of the shrub layer because of low mortality of the extensive clonal root systems. Rapid post-fire recovery of acorn production in xeric fire-prone habitats is presumably the result of selection to increase the probability of recovery and persistence following sufficiently intense fires that result in high oak mortality. The timing and magnitude of post-fire acorn production in sandhill and other xeric Florida associations has a potential impact on a wide variety of insects, birds, and mammals that feed on acorns, as well as on the species with which they interact.

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.

Few studies have examined how the preference-performance relationship of an herbivore for different genotypes of its host plant is affected by the presence and/or feeding activity of other members of the herbivore assemblage. In an outdoor garden, we manipulated the abundance of three common herbivores, the meadow spittlebug, a leaf beetle, and an aphid, on replicate 1-m 2 plots of 16 different genotypes of tall goldenrod, Solidago altissima. Adults of the goldenrod stem galler, Eurosta solidaginis, were subsequently released into the garden to oviposit among the host plants. Oviposition preference was strongly influenced by plant genotype and the presence of two of the herbivores, spittlebugs and leaf beetles. The effects of the herbivores were additive: the presence of leaf beetles reduced preference by 6%, spittlebugs by 18%, and both herbivores combined by 25%. Plant genotype-herbivore species interaction effects on stem-galler preference, which would indicate the presence of genetic variation among goldenrod genotypes in their norms of reaction for their acceptability as a host to the stem galler, were absent in this study. The performance of the stem galler was also significantly affected by goldenrod genotype, but in general was not affected by the presence of herbivores early in the season (the exception was a positive correlation between the proportion of ramets infested by all herbivores and gall size). Overall, we could find no correlation between preference and performance. This is in accord with results from previous studies on this system that were performed in the absence of herbivores, suggesting that the presence of herbivores in this study did not qualitatively alter the preference-performance relationship. We suggest that the lack of a positive correlation between host-plant preference and larval performance may reflect a constraint on the discriminatory ability of female stem gallers preventing them from selecting the best hosts among plants that differ in genotype and level of environmental stress (e.g., presence of interspecific herbivores).

Environmental catastrophes, such as severe drought, can reduce host-plant quality and/or abundance, which in turn decrease levels of herbivore populations. Such changes in herbivore populations affect populations of their natural enemies. As part of a long-term field experiment (1983-1991), galls of Eurosta solidaginis from 16 fields in central Pennsylvania were systematically collected from goldenrod ramets. Galls were dissected to compare the occurrence of E. solidaginis mortality caused by its natural enemies in 2 drought years (1988, 1991) with 5 pre-drought years (1983-1987) and 2 post-drought years (1989-1990). Gall diameters were significantly smaller in both drought years and early larval death significantly decreased E. solidaginis survivorship in the first drought year. Of the natural enemies, the parasitoid wasp Eurytoma gigantea caused significant selection for larger gall size in all pre-drought years, the 1991 drought, and both post-drought years, due to its differential attack of smaller galls. In spite of drought-induced small gall size in 1988, there was negligible selection on gall size by natural enemies. However, populations of E. solidaginis did suffer local extirpations at nine of the 16 fields during the first drought year and population recoveries of the gall inducer and natural enemies varied among fields in the post-drought years. As a consequence of reduced herbivore abundance in drought and post-drought years, some natural-enemy populations were absent. Drought therefore drastically reduced the abundance of E. solidaginis and natural enemies resulting in slow recoveries to pre-drought numbers.

Ecologists have labored to find an explanation for the lack of a positive correlation between host preference and offspring performance in herbivorous insects. This study focuses on how one herbivore species can influence another herbivore species' ability to accurately assess the suitability of different host-plant genotypes for larval development. In particular, we examined the role that an early season xylem-feeding homopteran (meadow spittlebug, Philaenus spumarius) has on the preference-performance correlation of a late-season dipteran stem galler (Eurosta solidaginis) among different goldenrod genotypes. In a greenhouse, we released adult stem gallers into replicate cages that contained ramets from four different goldenrod genotypes crossed with three densities of spittlebugs (0, 1, or 8 nymphs placed 2 weeks previously on each ramet). Spittlebug feeding caused a density-dependent decline in ramet growth rates, which in turn caused a corresponding decrease in host-plant preference by the stem gallers (number of ovipunctures per bud or proportion of ramets attacked). Goldenrod genotype and the interaction between spittlebugs and genotypes also influenced host-plant preference by the stem galler. Goldenrod genotype had the greatest impact on stem galler offspring performance (gall size or survivorship). Spittlebug density also affected performance, but only through its interaction with goldenrod genotype. On some genotypes, the survivorship of stem-galler larvae decreased with increasing spittlebug density, while on other genotypes, survivorship remained unchanged, or actually increased, with increasing spittlebug density. This suggests that there was genetic variance among goldenrod genotypes in their norms of reaction for their suitability as a host to the stem gallers. One possible explanation for why spittlebugs caused a significant reduction in preference, but not in performance, was that spittlebugs had very few long-term effects on the host plant. Flower number, flowering phenology, and the allocation of the ramet's biomass to different structures (below-ground organs, stems, leaves, and flowers) were unchanged with respect to spittlebug density. The only effect of spittlebugs was a 3-4% decrease in ramet height at the end of the growing season. We argue that the lack of a positive correlation between host-plant preference and larval performance may reflect a constraint on the discriminatory ability of female stem gallers. The damage to goldenrods caused by spittlebugs prior to attack by the stem gallers is similar in effect to potentially innumerable other causes of goldenrod stress (e.g., reduction in ramet growth rates). As a consequence, stem gallers may not be able to discern the subtle differences among stresses that identify those that will negatively affect the fitness of stem-galler offspring. The fact that goldenrod genotypes differ in their response to stresses would only further complicate the host-selection process. We propose that the stem gallers may have evolved a strategy that uses simple cues as the basis for rejecting similarly stressed plants, whether all of those plant genotype-stress combinations reduce performance or not.

We determined the phylogenetic relationships and geographic distribution of mitochondrial haplotypes of two host races of the tephritid fly Eurosta solidaginis, a gallmaker that attacks species of goldenrod (Solidago). We performed a preliminary survey by sequencing 492 bp from the 3' ends of the mitochondrial cytochrome oxidase I and II subunits from a single individual from eight S. gigantea- and 10 S. altissima-associated populations across their range in eastern North America and from two outgroup species, Eurosta comma (two populations) and E. cribrata. Eurosta solidaginis haplotypes fell into two groups ("E" and "W" clades), which differed by four substitutions, one of which occurred within the recognition site of the DdeI restriction enzyme. We used the presence or absence of the restriction site to survey a total of 11 S. gigantea (20 individuals) and 20 S. altissima (43 individuals) host-race populations. All gigantea-fly haplotypes regardless of geographic origin carried the E-clade haplotype, whereas altissima-fly haplotypes were geographically partitioned. Altissima flies east of Michigan were of haplotype E, whereas those west of Michigan were of haplotype W, with mixed populations found in lower Michigan. These patterns confirm an earlier allozyme survey that suggested that S. altissima is the ancestral host for the gallmaker, but also suggest that the gigantea fly populations were derived from eastern U.S. altissima fly populations. The data support the conclusions of behavioral and ecological studies indicating that the shift to the derived host was facilitated by escape from natural enemies., (© 1996 The Society for the Study of Evolution.)

The successful colonization of novel host-plant species by herbivorous insects may be facilitated by a reduction in natural-enemy attack on insect populations associated with the novel (derived) host plant. This is particularly true if natural enemies use host-plant or habitat cues in searching for their herbivore prey. In order to test whether the acquisition of enemy-free space could have influenced the host shift in the goldenrod ball gallmaker, Eurosta solidaginis, we estimated levels of natural-enemy attack in 25 host-race populations associated with Solidago altissima and S. gigantea (Compositae) spanning the zone of sympatry between S. altissima and S. gigantea host races in New England. Mortality due to attack by the parasitoid wasp Eurytoma obtusiventris was significantly higher for the ancestral than for the derived host race (30.5% versus 0.4%) across the transect, which is consistent with the enemy escape hypothesis. Contrary to this hypothesis, mordellid beetles caused significantly higher mortality on the derived than ancestral host race (17.1% versus 2.6%). Mortality by a second parasitoid wasp and birds showed no significant differences between the two host races. Overall, the derived host race had significantly higher survivorship across the transect (36.6% versus 20.8%). An analysis of survivorship and parasitoid mortality levels from sympatric sites in this study and previous studies showed a highly significant correlation between the levels of Eurytoma obtusiventris attack and the survivorship advantage of the derived host race. Observations of this parasitoid's searching behavior confirmed that it preferentially searches the ancestral host for fly larvae. Current patterns of host-race mortality and naturalenemy behavior and abundance are consistent with the facilitation of the host shift by escape from a specialist parasitoid.

We report behavioral evidence that Eurosta solidaginis, a stem-galling tephritid fly, has formed host races on its two goldenrod hosts, Solidago altissima and S. gigantea. Previous work has shown that flies from each host plant differ electrophoretically at the level of host races. The two host-associated populations were truly sympatric and were frequently found on host plants of the two species growing interdigitated with each other. Each host-associated population demonstrated a strong preference for ovipuncturing its own host. The S. gigantea-associated population emerged 10 to 14 d earlier than the S. altissima-associated population, contributing to the reproductive isolation between populations. Partial reproductive isolation is also maintained by a preference for mating on the host from which the fly emerged. The populations meet the criteria established for host races, suggesting that they may be in an intermediate stage of sympatric speciation., (© 1993 The Society for the Study of Evolution.)

Natural fluctuations in environmental conditions are likely to induce variation in the intensity or direction of natural selection. A long-term study of the insect, Eurosta solidaginins Fitch (Diptera; Tephritidae), which induces stem galls on the perennial herb Solidago altissima (Asteraceae) was performed to explore the patterns of variation in phenotypic selection. The intensity of selection imposed by parasitoids and predators on gallmaking larvae, for gall size, was measured across 16 populations over the course of 4 generations, for a total of 64 population-generations. Directional selection was quantified by i, the selection intensity, and variance selection by j', a measure of the intensity of selection on phenotypic variance. Size-dependent attack by parasitoids caused upward directional selection (mean i p = 0.42; SE = 0.023), while size-dependent bird attack favored larvae that induced smaller galls (mean i b = -0.07; SE = 0.013. The mean net directional selection intensity was 0.35 (SE = 0.030), which indicates that insects inducing larger galls are generally favored by selection. The opposing patterns of size-dependent attack resulted in stabilizing selection in half the population generations, with an overall average. j' of -0.11 (SE = 0.078). The magnitude of directional selection was strongly influenced by the population mean gall size and weakly by the optimal gall size. The intensity of variance selection was strongly influenced by the shape of the fitness function, with sigmoidal and Gaussian-like shapes causing greater depletion of phenotypic variance., (© 1992 The Society for the Study of Evolution.)

Plant resistance to insect herbivores may derive from traits influencing herbivore preference, traits influencing the suitability of the plant as a host, or both. However, the plant traits influencing host-plant selection by ovipositing insect herbivores may not completely overlap those traits that affect larval survival, and distinct traits may exhibit different levels of genetic vs. environmental control. Therefore, resource supply to the host plant could affect oviposition preference and larval performance differently in different plant genotypes. To test this hypothesis, the effects of resistance level, plant genotype, and resource supply to the host plant on oviposition preference and larval performance of a gallmaking herbivore, and on various plant traits that could influence these, were examined. Replicates of four genotypes of Solidago altissima, grown under low, medium, or high levels of nutrient supply in full sun or with medium levels of nutrients in shade, were exposed to mass-released Eurosta solidaginis. The number of plants ovipunctured was significantly affected by plant genotype and the interaction between genotype and nutrient supply to the host plant: one susceptible and one resistant genotype were more preferred, and preference tended to increase with nutrient supply in the more-preferred genotypes. The growth rate of ovipunctured plants during the oviposition period was significantly greater than that of unpunctured plants. Bud diameter (which was strongly correlated with plant growth rate), leaf area, and leaf water content were significant determinants of the percentage of plants ovipunctured, explaining 74% of the variance. The number of surviving larvae was significantly affected by plant genotype, but no effect of nutrient or light supply to the host plant was detected. The ratio of bud diameter to bud length was positively related to the percentage of ovipunctured plants that formed galls, suggesting that the accurate placement of eggs near the apical meristem by ovipositing females may be easier in short, thick buds. No significant correlation was observed between oviposition preference and larval survival at the population level. These results suggest that the plant traits affecting oviposition preference may exhibit different magnitudes of phenotypic plasticity than those affecting larval survival, and that the degree of phenotypic plasticity in plant traits affecting oviposition preference may differ among genotypes within a species.

Eurosta solidaginis, a gallmaking tephritid (Diptera), infests at least two species of Solidago in the eastern United States. We used horizontal starch gel electrophoresis of enzymes to examine whether populations on the two principal host plant species, Solidago altissima and S. gigantea, have diverged genetically. At the d-β-hydroxybutyrate dehydrogenase (HBDH) locus, the predominant allele in all nine populations from S. altissima (HBDH 1.00 ) was absent or uncommon in all but 1 of the 12 populations from S. gigantea. At the phosphoglucomutase locus (PGM), the most common allele in all populations was PGM 1.00 , but the frequency of this allele was, with one exception, higher in populations from S. gigantea than in populations from S. altissima. Genetic heterozygosity was usually greater in populations from S. altissima (H̄ = 0.028) than in populations from S. gigantea (H̄ = 0.009). A phylogenetic tree derived from a genetic distance matrix clustered gallmaker populations from the same host plant together., (© 1990 The Society for the Study of Evolution.)

Larvae of the tephritid fly Eurosta solidaginis induce ball-shaped galls on the stem of tall goldenrod, Solidago altissima. Survival probability depends on gall size; in small galls the larva is vulnerable to parasitoid oviposition, whereas larvae in large galls are more frequently eaten by avian predators. Fly populations from 20 natural old fields in central Pennsylvania were monitored in 1983 and 1984 to examine the distribution of the selection intensity imposed by natural enemies, the parasitoids Eurytoma gigantea and E. obtusiventris, the inquiline Mordellistena unicolor, and the predatory birds Dendrocopus pubescens and Parus atricapillus. Mordellistena and E. obtusiventris are able to attack galls of all diameters while E. gigantea and the predatory birds preferentially assaulted small and large diameter galls, respectively. Eurosta in intermediate sized galls had the highest survivorship, hence selection had a stabilizing component. However, parasitoid attack was more frequent than bird attack, and the two did not exactly balance, thus there was also a directional component. The mean directional selection intensity on gall size was 0.21 standard deviations of the mean, indicating that larger gall size was favored. Interactions among the insect members of the Eurosta natural enemy guild are complex and frequent.

Ramets from six Solidago altissima clones of known resistance to the stem gallmaker Eurosta solidaginis were grown with and without supplemental nutrients. In a greenhouse experiment, mated female Eurosta were allowed to oviposit in ramets that were subsequently grown through flowering and then harvested to determine biomass allocation. Supplemental nutrients increased plant biomass but did not affect resistance to this gallmaking herbivore. This result does not conform to the plant carbon/nutrient balance hypothesis which predicts that enhanced mineral nutrition will indirectly cause a reduction in carbon-based defensive chemistry. Our results indicate a strong genetic basis to ball gallmaker resistance since modification of host phenotype did not influence susceptibility. We suggest that evaluating the degree of genotypic or environmental control of plant resistance will be especially helpful in clarifying the patterns of defensive chemical responses.

The field site conditions (soil pH, soil moisture, soil nutrient availability, etc.) and abundances of Solidago altissima (often included in S. canadensis sensu lato), three S. altissima specific stem gall formers, and the parasitepredator guilds for two of the three gall insects were investigated. It was found that S. altissima is tolerant of a wide range of site conditions. Herbivore (stem gall insects) occurrences were positively correlated with plant occurrence, in a linear fashion. However, there was no disproportionate increase in stem gall insect densities with plant density as might be predicted by the resource concentration hypothesis. Parasitoid guilds were exploiting stem gall insect populations over a wide range of occurrence, but were under-utilizing fields of higher herbivore occurrences. Path analysis showed a high degree of predictability in the causal models, with all but 14% of the ball gall parasitoid guild and all but 43% of the elliptical gall parasitoid guild occurrences explained by the direct influences of stem gall insect occurrence and the indirect influences of goldenrod occurrence and site conditions. The numerical relations of this three trophic level system suggest a well-integrated and well-controlled food chain.

Armstrong (1982, 1983) predicted that all ramets within a clone should have the same ratio of biomass allocation to sexual reproduction versus vegetative growth. He presented data (1984) that he interpreted as showing that Solidago altissima ramets in a clone do have the predicted constant allocation ratio. Reanalysis of his methods shows that this conclusion was an artifact of his analysis. A simulation using random numbers and Armstrong's analysis showed the same pattern as his data. Data from S. altissima ramets of a single clone grown in a greenhouse experiment, using a different analysis, illustrated that the allocation ratios within a clone can be highly variable.

The allocations of biomass, N, P, and K were determined by standard methods in goldenrod ramets (1) parasitized by dipteran and lepidopterous gallmakers, (2) from fertilized and unfertilized plots, and (3) whose rhizome connections to their parental clone were severed. The presence of ball galls and their larvae increased allocation to stem but decreased allocation to leaves and seed production, and reduced the number of new rhizomes. There was a marked magnification of N and P concentrations going up the food chains; from goldenrods to gallmakers to the gallmaker's parasitoid/inquiline guild. Nutrient budgets expressed as flow diagrams indicated that N and P costs of gall presence were similar to energy costs under conditions where nutrients did not limit plant growth. Our data do not indicate that the growth of the galls of these gallmakers is limited by either N or P. Ramets from fertilized plots contained more N and P than controls but decreased the percentage of biomass allocated to leaves and inflorescences; ramets isolated by rhizome-cutting compensated their loss by increased allocation to roots, current rhizomes, and new rhizomes but at a cost of lower allocation to seed production and leaves. Gallmakers have a negative impact on host plant fitness characteristics. This may be especially important to establishing perennial hosts, given that herbivore effects would reduce clonal expansion and hence the ultimate clone size, thereby decreasing lifetime plant fitness.

Despite the widespread occurrence of plant-gallmaker interactions, little is known about the actual mechanisms of gall formation. To further characterize this type of parasite-host interaction, the mechanism of gall formation in Solidago altissima, tall goldenrod, by the larva of the tephritid fly Eurosta solidaginis was studied. Proteins produced by galled and ungalled tissues were examined, and the hyperinduction of a 58 kilodalton protein was observed in galled tissues for the second and third week of gall growth. The presence of this protein suggests that a substance secreted by the larva may function as a trans-acting gene regulator.

Three ramet clones of Solidago altissima were grown under greenhouse conditions to determine the effects of varying levels of attack by the goldenrod ball gallmaker (Eurosta solidaginis) on biomass allocation, leaf senescence rate and rhizome connections among ramets. The results, examined at both the individual ramet level and the clone level, showed that galled ramets became isolated from their clone through deterioration of rhizome connections. Gall effects were only observed at the ramet level although rhizome connection effects were detected at both the ramet and clone levels. The goldenrod ball gallmaker may therefore have little evolutionary impact on large clones but could appreciably affect newly established clones.