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Background and Aims Displacement of native plant species by non-native invaders may result from differences in their carbon economy, yet little is known regarding how variation in leaf traits influences native–invader dynamics across climate gradients. In Hawaii, one of the most heavily invaded biodiversity hotspots in the world, strong spatial variation in climate results from the complex topography, which underlies variation in traits that probably drives shifts in species interactions. Methods Using one of the most comprehensive trait data sets for Hawaii to date (91 species and four islands), we determined the extent and sources of variation (climate, species and species origin) in leaf traits, and used mixed models to examine differences between natives and non-native invasives. Key Results We detected significant differences in trait means, such that invasives were more resource acquisitive than natives over most of the climate gradients. However, we also detected trait convergence and a rank reversal (natives more resource acquisitive than invasives) in a sub-set of conditions. There was significant intraspecific trait variation (ITV) in leaf traits of natives and invasives, although invasives expressed significantly greater ITV than natives in water loss and photosynthesis. Species accounted for more trait variation than did climate for invasives, while the reverse was true for natives. Incorporating this climate-driven trait variation significantly improved the fit of models that compared natives and invasives. Lastly, in invasives, ITV was most strongly explained by spatial heterogeneity in moisture, whereas solar energy explains more ITV in natives. Conclusions Our results indicate that trait expression and ITV vary significantly between natives and invasives, and that this is mediated by climate. These findings suggest that although natives and invasives are functionally similar at the regional scale, invader success at local scales is contingent on climate.

Given that the rate of resource capture constrains plant growth and defence, understanding the linkage between the leaf economic spectrum (LES) and defence and how it contributes to growth is central to predicting species performance. In spite of the prevalence of spiny plants in many plant communities, little is known about how the LES relates to defence and growth rate across these species. We grew 42 spiny species, from diverse environments, under common garden conditions for 15 weeks and measured LES (leaf N, SLA and assimilation rate), defence and growth traits. We assessed general relationships between LES and growth rate and tested whether structural defences (spines, leaf fibre and lignin content) and quantitative chemical defences (condensed tannins) are linked to the LES and growth and if different spine types (i.e. leaf spines, stipular spines, prickles and thorns), with distinct anatomical origins, partition out across the LES. We observed two independent trait axes that together explained ~68% of trait variation across species. The first axis showed that structural defences (spines, leaf fibre and lignin content) trade off with leaf productivity along the LES. Axis 2 revealed that condensed tannins is orthogonal and less integrated with the LES‐structural defence axis. Bivariate trait analyses disclosed positive covariations between LES traits and sapling growth rate. All structural defence traits were negatively related to sapling growth. Across spine types, species with leaf spines were associated with the conservative end of the LES, characterized by high structural defences and lower leaf productivity relative to other spine types. Synthesis: Our study shows that the LES and structural defences are coupled in spiny species such that constitutive growth – defence strategies range from fast‐growing species with low allocation to defences to slow‐growing species that invest heavily in structural defences (dominated by leaf spiny species).

Biodiversity patterns emerge as a consequence of evolutionary and ecological processes. Their relative importance is frequently tested on model ecosystems such as oceanic islands that vary in both. However, the coarse-scale data typically used in biogeographic studies have limited inferential power to separate the effects of historical biogeographic factors (e.g., island age) from the effects of ecological ones (e.g., island area and habitat heterogeneity). Here, we describe local-scale biodiversity patterns of woody plants using a database of more than 500 forest plots from across the Hawaiian archipelago, where these volcanic islands differ in age by several million years. We show that, after controlling for factors such as island area and heterogeneity, the oldest islands (Kaua’i and O’ahu) have greater native species diversity per unit area than younger islands (Maui and Hawai’i), indicating an important role for macroevolutionary processes in driving not just whole-island differences in species diversity, but also local community assembly. Further, we find that older islands have a greater number of rare species that are more spatially clumped (i.e., higher within-island β-diversity) than younger islands. When we included alien species in our analyses, we found that the signal of macroevolutionary processes via island age was diluted. Our approach allows a more explicit test of the question of how macroevolutionary factors shape not just regional-scale biodiversity, but also local-scale community assembly patterns and processes in a model archipelago ecosystem, and it can be applied to disentangle biodiversity drivers in other systems.

Climate change models project an increase in the frequency and duration of drought globally. Changes in rainfall are expected to have particularly detrimental effects on seedlings due to their inability to reach deep water sources. We conducted a greenhouse experiment to test how the timing of watering affects seedling performance, physiology, and morphology in a keystone island endemic tree, Metrosideros polymorpha. Seedlings were grown for 60 days in one of five conditions: control treatment with daily watering to 100% field capacity (FC), a 28-day pulse drought with no water, and three groups maintained at 80% FC and watered every 2, 7, or 14 days. Seedling drought tolerance depended on the duration of drought; growth for plants watered to 80% FC every 2 or 14 days was comparable to 100% FC plants, but was reduced in the 28-day pulse drought and 7-day 80% FC. Survival was high in all treatments except the pulse drought. Stomatal conductance, chlorophyll content, and C:N ratio were positively related to height and biomass under drought. Traits associated with resource investment, fluorescence, and water use were plastic in their expression, shifting under drought compared to control conditions. M. polymorpha seedlings exhibit trait plasticity that allows them to tolerate drought, but its duration and intensity influence their survival and growth. Our results highlight the need for experimental studies manipulating temporal variation in water with explicit consideration of the role of trait plasticity in drought tolerance.

Background: Climates are changing at a rate that exceeds the adaptive capacity of species, especially endangered species. Genetic variation and phenotypic plasticity are important for population persistence, yet few studies have linked traits to seedling performance under drought in endangered species. Aims: We assessed intraspecific variation and trait plasticity under drought, to understand how an endangered species would cope with increasingly severe droughts. Methods: Using greenhouse experiments, we assessed drought tolerance of a critically endangered Hawaiian shrub, Schiedea obovata. Seedlings from five maternal families (three selfed and two interpopulation crosses) were subjected to daily watering or a simulated drought, and we measured growth, traits linked to drought tolerance, and days until death under terminal drought. Results: Drought reduced growth but not days until death. We detected genetic variation in growth, but no trait plasticity except in carbon:nitrogen, which decreased under drought. We did not detect traits that enhanced performance under drought but identified four physiological traits whose effects on growth varied under control and drought. Conclusions: Our results indicate moderate drought tolerance of an endangered shrub, and low trait plasticity. Conservation of endangered species under shifting climates will benefit from studies of stress tolerance, particularly at the vulnerable seedling stage.

Background Investigating the causes and consequences of intraspecific trait variation (ITV) in plants is not novel, as it has long been recognized that such variation shapes biotic and abiotic interactions. While evolutionary and population biology have extensively investigated ITV, only in the last 10 years has interest in ITV surged within community and comparative ecology. Scope Despite this recent interest, still lacking are thorough descriptions of ITV’s extent, the spatial and temporal structure of ITV, and stronger connections between ITV and community and ecosystem properties. Our primary aim in this review is to synthesize the recent literature and ask: (1) How extensive is intraspecific variation in traits across scales, and what underlying mechanisms drive this variation? (2) How does this variation impact higher-order ecological processes (e.g. population dynamics, community assembly, invasion, ecosystem productivity)? (3) What are the consequences of ignoring ITV and how can these be mitigated? and (4) What are the most pressing research questions, and how can current practices be modified to suit our research needs? Our secondary aim is to target diverse and underrepresented traits and plant organs, including anatomy, wood, roots, hydraulics, reproduction and secondary chemistry. In addressing these aims, we showcase papers from the Special Issue. Conclusions Plant ITV plays a key role in determining individual and population performance, species interactions, community structure and assembly, and ecosystem properties. Its extent varies widely across species, traits and environments, and it remains difficult to develop a predictive model for ITV that is broadly applicable. Systematically characterizing the sources (e.g. ontogeny, population differences) of ITV will be a vital step forward towards identifying generalities and the underlying mechanisms that shape ITV. While the use of species means to link traits to higher-order processes may be appropriate in many cases, such approaches can obscure potentially meaningful variation. We urge the reporting of individual replicates and population means in online data repositories, a greater consideration of the mechanisms that enhance and constrain ITV’s extent, and studies that span sub-disciplines.

Background: Prickly poppies (genus Argemone , Papaveraceae) epitomize well-defended plants. With high prickle densities and exudation of bright yellow latex from glaucous leaves, there are few reports of herbivore damage on these plants. Yet, little ecological work has examined within-plant or among-species variation in levels of anti-herbivore defenses in prickly poppies. Questions: Are prickly poppies well defended against generalist herbivores? Does chemical defense vary within prickly poppy plants in a pattern consistent with optimal defense theory? Species study: Argemone glauca, A. mexicana , A. ochroleuca, A. platyceras were examined in bioassays using generalist caterpillars, Agrotis ipsilon Hufnagel and Chrysodeixis eriosoma Doubleday (Noctuidae). Study sites and dates: Seeds were collected from field populations in Mexico (2015) and Hawaii (2011); experiment was conducted at the University of Hawaii at Manoa, Fall 2015. Methods: Using a pair of no-choice bioassays, variation in the quality of whole-leaf and chemical extract infused artificial diets for caterpillar development was assessed. Results: Survivorship was lowest on the control lettuce diet, and varied little between poppy diets although pupal mass was marginally lower for caterpillars reared on A. platyceras than A. ochroleuca . Isolating the effects of plant chemistry, C. eriosoma performance was most strongly reduced by seed extracts, with an extension in pupal development time by one week compared to herbivores reared on other diets, as well as a significant reduction in mean adult mass. Conclusions: These results are consistent with optimal defense theory, reflecting greater allocation of defense in tissues of high fitness value ( i.e., seeds). Future work with native herbivores, chemical analyses, and investigations into alternative functions for these plant traits would shed new light on this interesting group of plants.

As the most remote archipelago in the world, the Hawaiian Islands are home to a highly endemic and disharmonic biota that has fascinated biologists for centuries. Forests are the dominant terrestrial biome in Hawai‘i, spanning complex, heterogeneous climates across substrates that vary tremendously in age, soil structure, and nutrient availability. Species richness is low in Hawaiian forests compared to other tropical forests, as a consequence of dispersal limitation from continents and adaptive radiations in only some lineages, and forests are dominated by the widespread Metrosideros species complex. Low species richness provides a relatively tractable model system for studies of community assembly, local adaptation, and species interactions. Moreover, Hawaiian forests provide insights into predicted patterns of evolution on islands, revealing that while some evidence supports “island syndromes,” there are exceptions to them all. For example, Hawaiian plants are not as a whole less defended against herbivores, less dispersible, more conservative in resource use, or more slow-growing than their continental relatives. Clearly, more work is needed to understand the drivers, sources, and constraints on phenotypic variation among Hawaiian species, including both widespread and rare species, and to understand the role of this variation for ecological and evolutionary processes, which will further contribute to conservation of this unique biota. Today, Hawaiian forests are among the most threatened globally. Resource management failures – the proliferation of non-native species in particular – have led to devastating declines in native taxa and resulted in dominance by novel species assemblages. Conservation and restoration of Hawaiian forests now rely on managing threats including climate change, ongoing species introductions, novel pathogens, lost mutualists, and altered ecosystem dynamics through the use of diverse tools and strategies grounded in basic ecological, evolutionary, and biocultural principles. The future of Hawaiian forests thus depends on the synthesis of ecological and evolutionary research, which will continue to inform future conservation and restoration practices.

Background and Aims Herbivory by large mammals imposes a critical recruitment bottleneck on plants in many systems. Spines defend plants against large herbivores, and how early they emerge in saplings may be one of the strongest predictors of sapling survival in herbivore-rich environments. Yet little effort has been directed at understanding the variability in spine emergence across saplings. Methods We present a multispecies study examining whether and how sapling size, spine type and species' environmental niche (light and precipitation environment) influence early emergence and biomass investment in spines. A phylogenetically diverse pool of 45 species possessing different spine types (spines, prickles and thorns; that are derived from distinct plant organs: leaf, epidermis or cortex, and branch, respectively), were grown under common-garden conditions, and patterns of spine emergence and biomass allocation to spines at 5 and 15 weeks after transplanting were characterized. Key Results Spine type and species' resource niche were the main factors driving early emergence and investment patterns. Spines emerged earliest in leaf spine-bearing species, and latest in thorn-bearing species. The probability of early spine emergence increased with decreasing precipitation, and was greater in species from open than from closed habitats. Sapling investment in spines changed with plant mass but was contingent on spine type and habitat type. Conclusions Different spine types have strikingly different timing of expression, suggesting that developmental origins of spines play a critical role in sapling defences. Furthermore, species from different precipitation and light environments (open vs. closed habitats) showed contrasting patterns of early spine expression, suggesting that resource limitation in their native range may have driven divergent evolution of early defence expression.

Background and AimsGlobal climate change includes shifts in temperature and precipitation, increases in the frequency and intensity of extreme weather events and sea level rise, which will drastically impact coastal ecosystems. The aim of this study is to quantify salinity tolerance and to identify physiological mechanisms underlying tolerance across wholeplant ontogeny in two widespread native coastal plant species in Hawaiʻi, Jacquemontia sandwicensis (Convolvulaceae) and Sida fallax (Malvaceae).MethodsAt the seed, seedling, juvenile and mature ontogenetic stages, plants were exposed to high salinity watering treatments. Tolerance was assayed as the performance of stressed compared with control plants using multiple fitness metrics, including germination, survival, growth and reproduction. Potential physiological mechanisms underlying salinity tolerance were measured at each ontogenetic stage, including: photosynthesis and stomatal conductance rates, leaf thickness, leaf mass per area and biomass allocation.Key ResultsSalinity tolerance varied between species and across ontogeny but, overall, salinity tolerance increased across ontogeny. For both species, salinity exposure delayed flowering. Physiological and morphological leaf traits shifted across plant ontogeny and were highly plastic in response to salinity. Traits enhancing performance under high salinity varied across ontogeny and between species. For J. sandwicensis, water use efficiency enhanced growth for juvenile plants exposed to high salinity, while chlorophyll content positively influenced plant growth under salinity in the mature stage. For S. fallax, transpiration enhanced plant growth only under low salinity early in ontogeny; high transpiration constrained growth under high salinity across all ontogenetic stages.ConclusionsThat salinity effects vary across ontogenetic stages indicates that demographic consequences of sea level rise and coastal flooding will influence population dynamics in complex ways. Furthermore, even coastal dune plants presumably adapted to tolerate salinity demonstrate reduced ecophysiological performance, growth and reproduction under increased salinity, highlighting the conservation importance of experimental work to better project climate change effects on plants.

This data paper provides a description of OpenNahele, the open Hawaiian forest plot database. OpenNahele includes 530 forest plots across the Hawaiian archipelago containing 43,590 individuals of 185 native and alien tree, shrub and tree fern species across six islands. We include estimates of maximum plant size (D950.1 and Dmax3) for 58 woody plant species, a key functional trait associated with dispersal distance and competition for light. OpenNahele can serve as a platform to test key ecological, evolutionary and conservation questions in a hotspot archipelago. OpenNahele is the first database that compiles data from a large number of forest plots across the Hawaiian archipelago to allow broad and high resolution studies of biodiversity patterns. Keywords: Hawaii, forests, islands, biodiversity, community ecology, evolutionary ecology

Summary Most plants rely at least in part on physical defence traits to deter herbivores, and yet, very little is known about the inducibility of these traits. A meta-analysis was conducted on data extracted from 57 publications from 1982 to 2014, including 53 species in 20 plant families in which the induction of physical defence traits was examined. Log response ratio effect sizes were calculated for 112 separate responses and analysed to characterize general patterns and test whether induction differed between physical and chemical defence traits (in the same study), among defence traits, and in response to the damage type (artificial, herbivore, jasmonates). Physical defence traits are generally inducible, showing a mean increase of 52% in damaged compared to control plants. The magnitude of increase did not differ between physical and chemical defence traits measured within the same experiments, and nutrients showed no overall response. Physical defence traits varied significantly in their responses, with non-glandular trichomes showing the greatest magnitude of increase, and no induction detected for leaf toughness. Responses induced by real herbivores or by jasmonate application were significantly greater than responses elicited by mechanical damage alone. This suggests that specificity in physical trait induction in response to different kinds of damage may occur. Induction of physical defence traits is common and widespread, but the ecological and evolutionary consequences of this response remain unknown. Future studies examining genetic variation in the inducibility of physical defence traits and their fitness consequences for plants would be particularly informative.

Herbivores are important drivers of plant population dynamics and community composition in natural and managed systems. Intraspecific genetic diversity of long-lived plants like trees might shape patterns of herbivory by different guilds of herbivores that trees experience through time. However, previous studies on plant genetic diversity effects on herbivores have been largely short-term. We investigated how tree genotypic variation and diversity influence herbivory of silver birch Betula pendula in a long-term field experiment. Using clones of eight genotypes, we constructed experimental plots consisting of one, two, four or eight genotypes, and measured damage by five guilds of arthropod herbivores twice a year over three different years (four, six and nine years after the experiment was established). Genotypes varied significantly for most types of herbivore damage, but genotype resistance rankings often shifted over time, and none of the clones was more resistant than all others to all types of herbivores. At the plot level, birch genotypic diversity had significant positive additive effect on leaf rollers and negative non-additive effects on chewing herbivores and gall makers. In contrast, leaf-mining and leaf-tying damage was not influenced by birch genotypic diversity. Within diverse plots, the direction of genotypic diversity effects varied depending on birch genotype, some having lower and some having higher herbivory in mixed stands. This research highlights the importance of long-term studies including different feeding guilds of herbivores to understand the effects of plant genetic diversity on arthropod communities. Different responses of various feeding guilds to genotypic diversity and shifts in resistance of individual genotypes over time indicate that genotypic mixtures are unlikely to result in overall reduction in herbivory over time.

Ontogeny has been identified as a main source of variation in the expression of plant phenotypes. However, there is limited information on the mechanisms behind the evolution of ontogenetic trajectories in plant defense. We explored if risk of attack, herbivore damage, heritability, and phenotypic plasticity can promote or constrain the evolutionary potential of ontogenetic trajectories in three defensive traits. We exposed 20 genotypes of Turnera velutina to contrasting environments (shadehouse and field plots), and measured the cyanogenic potential, trichome density, and sugar content in extrafloral nectar in seedlings, juveniles and reproductive plants. We also assessed risk of attack through oviposition preferences, and quantified herbivore damage in the field. We estimated genetic variance, broad sense heritability, and evolvability of the defensive traits at each ontogenetic stage, and of the ontogenetic trajectories themselves. For plants growing in the shadehouse, we found genetic variation and broad sense heritability for cyanogenic potential in seedlings, and for trichome density at all ontogenetic stages. Genetic variation and heritability of ontogenetic trajectories was detected for trichome density only. These genetic pre-requisites for evolution, however, were not detected in the field, suggesting that environmental variation and phenotypic plastic responses mask any heritable variation. Finally, ontogenetic trajectories were found to be plastic, differing between shadehouse and field conditions for the same genetic families. Overall, we provide support for the idea that changes in herbivore pressure can be a mechanism behind the evolution of ontogenetic trajectories. This evolutionary potential, however, can be constrained by phenotypic plasticity expressed in heterogeneous environments.

Plant-herbivore interactions have been predicted to play a fundamental role in plant invasions, although support for this assertion from previous research is mixed. While plants may escape from specialist herbivores in their introduced ranges, herbivory from generalists is common. Tolerance traits may allow non-native plants to mitigate the negative consequences of generalist herbivory that they cannot avoid in their introduced range. Here we address whether tolerance to herbivory, quantified as survival and compensatory growth, is associated with plant invasion success in Hawaii and investigate traits that may enhance tolerance in seedlings, the life stage most susceptible to herbivory. In a greenhouse experiment, we measured seedling tolerance to simulated herbivory through mechanical damage (50% leaf removal) of 16 non-native woody plant species differing in invasion status (invasive vs. non-invasive). Seedlings were grown for 2 weeks following damage and analyzed for biomass to determine whether damaged plants could fully compensate for the lost leaf tissue. Over 99% of all seedlings survived defoliation. Although species varied significantly in their levels of compensation, there was no consistent difference between invasive and non-invasive species. Seedlings of 11 species undercompensated and remained substantially smaller than control seedlings 2 weeks after damage; four species were close to compensating, while one species overcompensated. Across species, compensation was positively associated with an increased investment in potential storage reserves, specifically cotyledons and roots, suggesting that these organs provide resources that help seedlings re-grow following damage. Our results add to a growing consensus that pre-damage growth patterns determine tolerance to damage, even in young seedlings which have relatively low biomass. The lack of higher tolerance in highly invasive species may suggest that invaders overcome herbivory barriers to invasion in other ways, such as resistance traits, or that herbivory does not play an important role in the seedling invasion dynamics of these woody species in Hawaii.

BACKGROUND AND AIMS Herbivory and plant defence differ markedly among seedlings and juvenile and mature plants in most species. While ontogenetic patterns of chemical resistance have been the focus of much research, comparatively little is known about how tolerance to damage changes across ontogeny. Due to dramatic shifts in plant size, resource acquisition, stored reserves and growth, it was predicted that tolerance and related underlying mechanisms would differ among ontogenetic stages. METHODS Ontogenetic patterns in the mechanisms of tolerance were investigated in Plantago lanceolata and P. major (Plantaginaceae) using the genetic sib-ship approach. Pot-grown plants were subjected to 50 % defoliation at the seedling, juvenile and mature stages and either harvested in the short-term to look at plasticity in growth and photosynthesis in response to damage or allowed to grow through seed maturation to measure phenology, shoot compensation and reproductive fitness. KEY RESULTS Tolerance to defoliation was high in P. lanceolata, but low in P. major, and did not vary among ontogenetic stages in either species. Mechanisms underlying tolerance did vary across ontogeny. In P. lanceolata, tolerance was significantly related to flowering (juveniles) and pre-damage shoot biomass (mature plants). In P. major, tolerance was significantly related to pre-damage root biomass (seedlings) and induction of non-photochemical quenching, a photosynthetic parameter (juveniles). CONCLUSIONS Biomass partitioning was very plastic in response to damage and showed associations with tolerance in both species, indicating a strong role in plant defence. In contrast, photosynthesis and phenology showed weaker responses to damage and were related to tolerance only in certain ontogenetic stages. This study highlights the pivotal role of ontogeny in plant defence and herbivory. Additional studies in more species are needed to determine how seedlings tolerate herbivory in general and whether mechanisms vary across ontogeny in consistent patterns.

Plant defence often varies by orders of magnitude as plants develop from the seedling to juvenile to mature and senescent stages. Ontogenetic trajectories can involve switches among defence traits, leading to complex shifting phenotypes across plant lifetimes. While considerable research has characterised ontogenetic trajectories for now hundreds of plant species, we still lack a clear understanding of the molecular, ecological and evolutionary factors driving these patterns. In this study, we identify several non-mutually exclusive factors that may have led to the evolution of ontogenetic trajectories in plant defence, including developmental constraints, resource allocation costs, multi-functionality of defence traits, and herbivore selection pressure. Evidence from recent physiological studies is highlighted to shed light on the underlying molecular mechanisms involved in the regulation and activation of these developmental changes. Overall, our goal is to promote new research avenues that would provide evidence for the factors that have promoted the evolution of this complex lifetime phenotype. Future research focusing on the questions and approaches identified here will advance the field and shed light on why defence traits shift so dramatically across plant ontogeny, a widespread but poorly understood ecological pattern.

BACKGROUND AND AIMS Seedling herbivory is an important factor underlying plant community diversity and structure. While considerable research has characterized seedling defence in terms of resistance, very little is known about seedling tolerance of herbivory. Moreover, few studies have attempted to identify mechanisms of tolerance across a range of plant species. METHODS Seedling tolerance of simulated herbivory was tested in a diverse pool of ten Hawaiian plant species, including several lobeliad species (family Campanulaceae), a grass, a herb and common woody trees and shrubs. Tolerance was measured as the relative survival and growth of damaged plants receiving 50 % defoliation with simultaneous jasmonic acid application compared with undamaged control plants, assessed 1·5 and 5 weeks after damage. Putative mechanisms of tolerance were measured, including photosynthetic parameters, light use efficiency, and biomass allocation reflecting growth priorities, and analysed using species-level regression analyses on tolerance indices. KEY RESULTS No species fully tolerated 50 % defoliation at either harvest date, and simulated herbivory significantly reduced shoot as well as root biomass. Lobeliad species had particularly low tolerance. Species varied considerably in size, biomass allocation parameters and their constitutive (pre-damage) and induced (post-damage) photosynthetic parameters. However, only constitutive levels of non-photochemical quenching were significantly related to tolerance, indicating that species with more efficient light use (and less heat dissipation) are better at tolerating damage than species with high levels of heat dissipation. CONCLUSIONS Native Hawaiian plants expressed low tolerance to a conservative level of simulated herbivory. Root growth decreased in response to damage, but this was not associated with greater tolerance, suggesting this response may be due to allocation constraints following defoliation and not due to adaptive plasticity. Conservation of native island plants threatened by invasive herbivores should prioritize protection for seedlings for improved regeneration and the persistence of native plants in disturbed habitats.

Phenotypic plasticity in growth (leading to compensation) and secondary chemical production (leading to induction) in response to herbivory are key defense strategies in adult plants, but their role in seedling defense remains unclear. A pair of greenhouse studies was conducted to investigate compensation and induction in seedlings and juvenile plants, using Plantago lanceolata (Plantaginaceae) and the specialist buckeye caterpillar Junonia coenia (Nymphalidae) as a model system. Plants received 50% defoliation at two and four weeks of age, and groups of plants were harvested one week after herbivory and six to eight weeks after herbivory to investigate the duration of the responses. Plants damaged at two weeks showed no chemical induction and fully compensated for the lost leaf tissue by ten weeks of age. Plants damaged at four weeks showed a significant reduction in iridoid glycosides one week after herbivory and achieved full shoot compensation by ten weeks of age at the expense of root biomass. These results indicate that P. lanceolata seedlings use compensation, but not chemical induction, as a defense strategy. This research highlights the importance of considering ontogeny in studies of plant-herbivore interactions and suggests that seedling defense may differ markedly from adult plant defense.

We examined variation in leaf size and specific leaf area (SLA) in relation to the distribution of 22 chaparral shrub species on small-scale gradients of aspect and elevation. Potential incident solar radiation (insolation) was estimated from a geographic information system to quantify microclimate affinities of these species across north- and south-facing slopes. At the community level, leaf size and SLA both declined with increasing insolation, based on average trait values for the species found in plots along the gradient. However, leaf size and SLA were not significantly correlated across species, suggesting that these two traits are decoupled and associated with different aspects of performance along this environmental gradient. For individual species, SLA was negatively correlated with species distributions along the insolation gradient, and was significantly lower in evergreen versus deciduous species. Leaf size exhibited a negative but non-significant trend in relation to insolation distribution of individual species. At the community level, variance in leaf size increased with increasing insolation. For individual species, there was a greater range of leaf size on south-facing slopes, while there was an absence of small-leaved species on north-facing slopes. These results demonstrate that analyses of plant functional traits along environmental gradients based on community level averages may obscure important aspects of trait variation and distribution among the constituent species.

Plants frequently attract natural enemies of their herbivores, resulting in a reduction in tissue damage and often in enhanced plant fitness. While such indirect defenses can dramatically change as plants develop, only recently have ecologists begun to explore such changes and evaluate their role in mediating plant-herbivore-natural enemy interactions. Here we review the literature documenting ontogenetic patterns in plant rewards (i.e. extrafloral nectaries (EFNs), food bodies (FBs) and domatia) and volatile organic compounds (VOCs), and identify links between ontogenetic patterns in such traits and the attraction of natural enemies (ants). In the case of reward traits we concentrate in ant-plant studies, which are the most numerous. We report that all indirect defensive traits commonly vary with plant age but ontogenetic trajectories differ among them. Myrmecophytic species, which provide both food and shelter to their defenders, do not produce rewarding traits until a minimum size is reached. Then, a pronounced increase in the abundance of food rewards and domatia often occurs as plants develop, which explains the temporal succession or colony size increase of mutualistic ant species and, in some cases, leads to a reduction in herbivore damage and enhanced fitness as plants age. In contrast, ontogenetic patterns were less consistent in plant species that rely on VOC emissions to attract natural enemies or those that provide only food rewards (EFNs) but not nesting sites to their associated ants, showing an overall decline or lack of trend with plant development, respectively. Future research should focus on uncovering: (i) the costs and mechanisms underlying ontogenetic variation in indirect defenses, (ii) the relative importance of environmental and genetic components shaping these ontogenetic trajectories, and (iii) the consequences of these ontogenetic trajectories on plant fitness. Advances in this area will shed light on the context dependency of bottom-up and top-down controls of herbivore populations and on how natural selection actually shapes the ontogenetic trajectories of these traits. Fil: Quintero, Carolina. State University Of Colorado-boulder; Estados Unidos. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Patagonia Norte. Instituto de Investigación en Biodiversidad y Medioambiente; Argentina Fil: Barton, Kasey E.. University Of Hawaii At Manoa; Estados Unidos Fil: Boege, Karina. Universidad Nacional Autónoma de México; México

Covariance among plant defense traits is predicted to occur both within and among plant species, potentially leading to characteristic defense syndromes. I examined patterns of variation in prickle density, latex exudation, and tolerance in order to assess whether traits varied between populations, across plant ontogeny, and as phenotypic plasticity in response to water and light limitation and physical damage using the endemic Hawaiian prickly poppy, Argemone glauca, as a model system. Plants produced copious latex, had extremely variable prickle densities, and were generally tolerant of 50 % defoliation. However, expression patterns differed among defense traits. Prickle density was consistent across ontogeny and was not induced by either water limitation or mechanical damage, but was significantly induced under high light conditions. In contrast, latex exudation increased significantly across ontogeny and was reduced by water limitation, but had no response to mechanical damage or light. Prickles, latex, and tolerance differed considerably between populations, suggesting different evolutionary histories for these populations. These disparate patterns indicate that latex and prickles are unlinked within A. glauca, potentially as a result of differences in their function, and providing little evidence that they jointly function as a defense syndrome. Moreover, this study provides the first description patterns of variation for multiple defense traits in an island endemic, and high levels of prickles, latex, and tolerance suggest that A. glauca is well defended against herbivores. Future research in the field will provide additional insights into the functional ecology of these traits in A. glauca.

Predictions based on the plant age and growth-differentiation balance hypotheses of defense were tested in two congeneric species, Plantago lanceolata and P. major, by quantifying iridoid glycosides, defensive chemicals, in seeds and leaves during the first 6 wk of growth. Concentrations decreased from the seed to 2-wk-old seedling stage in P. lanceolata, but increased during this period in P. major. In both species, levels were similar for 2- and 4-wk-old plants, then significantly increased from 4 to 6 wk. Genetic variation in the ontogeny of iridoid glycoside production was significant in both species at the maternal family level and at the population level. To examine whether allocation costs could explain the low production of iridoid glycosides in seedlings, relationships between growth and defense (iridoid glycosides) were characterized. Growth and defense had a positive or null relationship in all age groups, indicating that there was no trade-off in these plants at any age. This study provides some support for the growth-differentiation balance hypothesis, but offers no support for the plant age hypothesis. Measuring how herbivory affects plant fitness at different ontogenetic stages may shed light on these patterns in Plantago and on the evolution of the ontogeny of defense.

As trees develop, they undergo morphological and physiological changes that can influence not only their performance, but also their interactions with herbivores. The expression of their defenses is influenced by changes in the selective pressures exerted by herbivores and by the plant’s tradeoffs in resource allocation, and can result in ontogenetic trajectories that show increases, decreases, or mixed trends in the expression of anti-herbivory traits. In some species, these trajectories occur as gradual transitions among ontogenetic stages, but in other species there are pronounced phase changes marked by heterophylly or by abrupt changes in chemical, physical, or biotic defenses. This chapter discusses the patterns of such trajectories and the multiple factors that can influence them, including the specific herbivores feeding on trees, the activities of herbivores’ natural enemies, the switches among the different defensive mechanisms that trees can express to reduce herbivory, the particular ecosystem in which they grow, and the influence of phylogenetic constraints that restrict or allow the evolution of onto­genetic trajectories in plant defense. Studies that integrate the role of ontogeny into evolutionary ecology theory will advance our understanding of how natural selection can target the ontogenetic trajectories of plant defense. Such research will also have application for targeting pest control onto vulnerable ontogenetic stages, and for selection of lines with improved defensive mechanisms to protect rare and endangered species as well as promote productivity in commercial stands.

Defense against herbivores often changes dramatically as plants develop. Hypotheses based on allocation theory and herbivore selection patterns predict that defense should increase or decrease, respectively, across ontogeny, and previous research partly supports both predictions. Thus, it remains unclear which pattern is more common and what factors contribute to variability among studies. We conducted a meta-analysis of 116 published studies reporting ontogenetic patterns in plant defense traits and herbivory. Patterns varied depending on plant life form (woody, herbaceous, grass), type of herbivore (insect, mollusk, mammal), and type of defense trait (secondary chemistry, physical defense, tolerance). In woody plants, chemical defense increased during the seedling stage, followed by an increase in physical defenses during the vegetative juvenile stage. Mammalian herbivores showed a strong preference for mature compared to juvenile tissues in woody plants. Herbs experienced a significant increase in secondary chemistry across the entire ontogenetic trajectory, although the magnitude of increase was greatest during the seedling stage. Correspondingly, mollusks preferred young compared to older herbs. Future research investigating growth/defense trade-offs, allometry, herbivore selection patterns, and ecological costs would shed light on the mechanisms driving the ontogenetic patterns observed.

Stressful parental environments can influence offspring size and development either adaptively or maladaptively, yet little is known about species' differences in this complex aspect of phenotypic plasticity. We performed a reciprocal split-brood experiment to compare transgenerational plasticity in response to drought stress in two closely related annual plant species. We raised inbred replicate parent plants of eight genotypes per species in dry vs. moist soil to generate offspring of each genetic line that differed only in parental environment, then monitored seedling development in both dry and moist conditions. Individuals of the two species expressed contrasting patterns of transgenerational plasticity for traits important to seedling drought tolerance. In Polygonum persicaria, a weedy generalist found in moist, dry, and variably dry sites, drought-stressed plants produced offspring with longer and more rapidly extending root systems and greater biomass when growing in dry soil. In contrast, in P. hydropiper, a non-weedy congener restricted to moist habitats, the offspring of drought-stressed parents had reduced root system development and seedling biomass in dry soil. In P. persicaria, transgenerational and immediate adaptive plasticity combined to produce drought-adapted seedling phenotypes. These results make clear that characteristic patterns of transgenerational plasticity can contribute to ecological diversity among species.

Plant ontogeny is a common source of variation in defense and herbivory. Yet, few studies have investigated how the induction of physical defense traits changes across plant ontogeny. Physical defense traits are costly to produce, and thus, it was predicted that induction as a cost-saving strategy would be particularly favorable for seedlings, leading to ontogenetic declines in the inducibility of these traits. We tested for induction of three different physical defense traits (prickles, latex and leaf toughness) in response to mechanical defoliation and jasmonic acid application using prickly poppies (Argemone glauca and A. mexicana, Papaveraceae) as a model system. Genetic variation in the induction of physical defenses was tested using maternal sib-ships sampled from multiple populations. Both species induced higher densities of laminar prickles, although the magnitude of induction was much higher in the endemic Hawaiian prickly poppy, A. glauca, than in the cosmopolitan A. mexicana. The magnitude of prickle induction was also higher in young compared to older juvenile plant stages in A. glauca, demonstrating a strong role of ontogeny. Neither latex exudation nor leaf toughness was induced in either species. Although significant genetic variation was detected within and among populations for constitutive expression of physical defense traits in Argemone, there was no evidence for genetic variation in the induction of these traits. This study provides the first evidence for the induction of physical defenses in prickly poppies, emphasizing how an ontogenetically explicit framework can reveal new insights into plant defense. Moreover, this study illustrates how sister species comparisons between island vs. continental plants can provide new insights into plant functional and evolutionary ecology, highlighting a fruitful area for future research on more species pairs.

Invasive species, where successful, can devastate native communities. We studied the dynamics of the invasive Argentine ant, Linepithema humile, for 7 years in Jasper Ridge, a biological preserve in northern California. We monitored the distributions at the hectare scale of native ant taxa and L. humile in the spring and fall from 1993 to 1999. We also studied the invasion dynamics at a finer resolution by searching for ants in 1-m2 plots. Our results are similar at both scales. The distributions of several native species are not random with regard to L. humile; the distributions of several epigeic species with similar habitat affinities overlap much less frequently than expected with the distribution of L. humile. We found that season had a significant influence on the distributions of L. humile and several native taxa. Over the 7-year period, L. humile has increased its range size in Jasper Ridge largely at the expense of native taxa, but there is seasonal and yearly variation in this rate of increase. Studies of invasions in progress which sample across seasons and years may help to predict the spread and effects of invasive species.

The ants Aphaenogaster cockerelli and Pogonomyrmex barbatus compete for seed resources in the Chihuahuan desert. Previous work showed that intraspecific competition in P. barbatus is more intense between near neighbors with overlapping foraging ranges and depends on colony age. Just before reaching reproductive maturity (3–4 y), colonies are more aggressive and persistent in intraspecific competition for foraging area than younger or older colonies. In this study we examine how interspecific interference behavior by A. cockerelli towards P. barbatus depends on the age and proximity of P. barbatus colonies. Before sunrise when P. barbatus colonies become active, A. cockerelli colonies completely plug the nest entrances of some P. barbatus colonies, thereby delaying the onset of P. barbatus foraging behavior. Pogonomyrmex barbatus colonies closer to A. cockerelli were plugged more frequently than more distant colonies. As distance from A. cockerelli nests increased, older P. barbatus colonies were ...