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Ontogenetic changes in anti-herbivore defences are common and result from variation in resource availability and herbivore damage throughout plant development. However, little is known about the simultaneous changes of multiple defences across the entire development of plants, and how such changes affect plant damage in the field. The aim of this study was to assess if changes in the major types of plant resistance and tolerance can explain natural herbivore damage throughout plant ontogeny. An assessment was made of how six defensive traits, including physical, chemical and biotic resistance, simultaneously change across the major transitions of plant development, from seedlings to reproductive stages of Turnera velutina growing in the greenhouse. In addition, an experiment was performed to assess how plant tolerance to artificial damage to leaves changed throughout ontogeny. Finally, leaf damage by herbivores was evaluated in a natural population. The observed ontogenetic trajectories of all defences were significantly different, sometimes showing opposite directions of change. Whereas trichome density, leaf toughness, extrafloral nectary abundance and nectar production increased, hydrogen cyanide and compensatory responses decreased throughout plant development, from seedlings to reproductive plants. Only water content was higher at the intermediate juvenile ontogenetic stages. Surveys in a natural population over 3 years showed that herbivores consumed more tissue from juvenile plants than from younger seedlings or older reproductive plants. This is consistent with the fact that juvenile plants were the least defended stage. The results suggest that defensive trajectories are a mixed result of predictions by the Optimal Defence Theory and the Growth-Differentiation Balance Hypothesis. The study emphasizes the importance of incorporating multiple defences and plant ontogeny into further studies for a more comprehensive understanding of plant defence evolution.

Plant functional strategies are usually accomplished through the simultaneous expression of different traits, and hence their correlations should be promoted by natural selection. The adaptive value of correlations among leaf functional traits, however, has not been assessed in natural populations. We estimated intraspecific variation in leaf functional traits related to the primary metabolism and anti-herbivore defence in a population of Turnera velutina. We analysed whether natural selection favoured the expression of individual traits, particular combinations of traits or leaf phenotypic integration Patterns of covariation among traits were related to water and nitrogen economy, and were similar among genotypes, but the magnitude of their phenotypic integration differed by 10-fold. Although families did not differ in the mean values of leaf functional traits, directional selection favoured low nitrogen content and low chemical defence, high content of chlorophyll, sugar in extrafloral nectar and trichome density. Families with higher phenotypic integration among leaf traits grew faster and produced more flowers. We suggest that the coordinated expression of leaf traits has an adaptive value, probably related to optimisation in the expression of traits related to water conservation and nitrogen acquisition.

• Distyly is an intriguing floral adaptation that increases pollen transfer precision and restricts inbreeding. It has been a model system in evolutionary biology since Darwin. Although the S-locus determines the long- and short-styled morphs, the genes were unknown in Turnera. We have now identified these genes. • We used deletion mapping to identify, and then sequence, BAC clones and genome scaffolds to construct S/s haplotypes. We investigated candidate gene expression, hemizygosity, and used mutants, to explore gene function. • The s-haplotype possessed 21 genes collinear with a region of chromosome 7 of grape. The S-haplotype possessed three additional genes and two inversions. TsSPH1 was expressed in filaments and anthers, TsYUC6 in anthers and TsBAHD in pistils. Long-homostyle mutants did not possess TsBAHD and a short-homostyle mutant did not express TsSPH1. • Three hemizygous genes appear to determine S-morph characteristics in T. subulata. Hemizygosity is common to all distylous species investigated, yet the genes differ. The pistil candidate gene, TsBAHD, differs from that of Primula, but both may inactivate brassinosteroids causing short styles. TsYUC6 is involved in auxin synthesis and likely determines pollen characteristics. TsSPH1 is likely involved in filament elongation. We propose an incompatibility mechanism involving TsYUC6 and TsBAHD.

Plant biomass allocation is mainly affected by the environment where each individual grows. In this sense, through the rapid global expansion of impermeable areas, urbanization has strong, albeit poorly understood, consequences on the biomass allocation of plants found in this environment. Nevertheless, the comprehension of biomass allocation processes in urban shrubs remains unclear, because most studies of urban ecology focus on tree species. This is an important gap of knowledge because a great part of urban vegetation is composed of shrubs and their association with trees have positive impacts in urban ecosystem services. In this study, we explored the ecological and potential selective pressure effects of an urbanization gradient on the biomass allocation patterns of aboveground organs of Turnera subulata, a widely distributed tropical shrub. We have demonstrated that, for certain reproductive organs, biomass allocation decreases in locations with higher urbanization. Unlike expected, the biomass of vegetative organs was not affected by urbanization, and we did not observe any effect of urbanization intensity on the variance in biomass allocation to vegetative and reproductive organs. We did not record urbanization-mediated trade-offs in biomass allocation for reproductive and vegetative organs. Instead, the biomass of these structures showed a positive relationship. Our data suggest that urbanization does not result in radical changes in biomass allocation of T. subulata, and neither in the variation of these traits. They indicate that the ability of T. subulata to thrive in urban environments may be associated with life history and morphological mechanisms. Our findings contribute to the understanding of shrub plant responses to urbanization and highlight urbanization as a potential factor in resource allocation differences for different structures and functions in plants living in these environments.

Changes in resource availability, functional demands, hormonal regulation and developmental constraints can promote differences in the expression of leaf traits during plant development and foster changes in the targets of natural selection. As a consequence, the pattern and magnitude of covariation among traits, and therefore their phenotypic integration and modularity are equally expected to change throughout ontogeny. However, these changes have not been described yet. We measured leaf economic, defensive and morphological traits in plants of Turnera velutina and estimated the magnitude and pattern of foliar integration and modularity for juvenile and reproductive individuals. In addition, we assessed the relationship between plant biomass and foliar integration within and among ontogenetic stages. Both the pattern and magnitude of foliar integration changed across plant ontogeny. Foliar integration was lower in juvenile than in reproductive plants, and the pattern of phenotypic integration and modularity was different between ontogenetic stages, whereas leaves from juvenile plants showed two functional modules related to plant defence and leaf economy, traits from reproductive plants had greater interconnectivity and hence lower modularity. The relationship between plant biomass and foliar integration was negative within each ontogenetic stage but positive between ontogenetic stages, suggesting that processes intrinsic to plant development influenced the magnitude of foliar integration to a greater extent than plant size. Our findings indicate that plants can change the patterns of covariation among leaf traits during their development. However, a lower foliar integration in juvenile plants could allow for greater lability to explore a multi‐trait phenotypic space, canalisation of leaf attributes along ontogeny should promote greater phenotypic integration, constraining the number of multi‐trait combinations that plants can express. Hence, we suggest that ontogenetic changes in foliar integration allow plants to deal with changing selective dynamics and physiological priorities along their development. A plain language summary is available for this article. Plain Language Summary

• The evolution of plant defenses has traditionally been studied at single plant ontogenetic stages, overlooking the fact that natural selection acts continuously on organisms along their development, and that the adaptive value of phenotypes can change along ontogeny. • We exposed 20 replicated genotypes of Turnera velutina to field conditions to evaluate whether the targets of natural selection on different defenses and their adaptative value change across plant development. • We found that low chemical defense was favored in seedlings, which seems to be explained by the assimilation efficiency and the ability of the specialist herbivore to sequester cyanogenic glycosides. Whereas trichome density was unfavored in juvenile plants, it increased relative plant fitness in reproductive plants. At this stage we also found a positive correlative gradient between cyanogenic potential and sugar content in extrafloral nectar. • We visualize this complex multi-trait combination as an ontogenetic defensive strategy. The inclusion of whole-plant ontogeny as a key source of variation in plant defense revealed that the targets and intensity of selection change along the development of plants, indicating that the influence of natural selection cannot be inferred without the assessment of ontogenetic strategies in the expression of multiple defenses.

Many plants attract insect pollinators with floral nectar (FN) and ant “bodyguards” with extrafloral nectar (EFN). If nectar production is costly or physiologically linked across glands, investment in one mutualism may trade off with investment in the other. We confirmed that changes in FN and EFN availability alter pollination and ant defense mutualisms in a field population of Turnera ulmifolia. Plants with additional FN tended to produce more seeds, while plants with reduced EFN production experienced less florivory. We then mimicked the consumptive effects of mutualists by removing FN or EFN daily for 50 days in a full factorial design using three Turnera species (T. joelii, T. subulata, and T. ulmifolia) in a glasshouse experiment. For T. ulmifolia and T. subulata, but not T. joelii, removing either nectar reduced production of the other, showing for the first time that EFN and FN production can trade off. In T. subulata, increased investment in FN decreased seed set, suggesting that nectar production can have direct fitness costs. Through the linked expression of EFN and FN, floral visitors may negatively affect biotic defense, and extrafloral nectary visitors may negatively affect pollination.

Ant guards protect plants from herbivores, but can also hinder pollination by damaging reproductive structures and/or repelling pollinators. Natural selection should favour the evolution of plant traits that deter ants from visiting flowers during anthesis, without waiving their defensive services. The Distraction Hypothesis posits that rewarding ants with extrafloral nectar could reduce their visitation of flowers, reducing ant‐pollinator conflict while retaining protection of other structures. We characterised the proportion of flowers occupied by ants and the number of ants per flower in a Mexican ant‐plant, Turnera velutina. We clogged extrafloral nectaries on field plants and observed the effects on patrolling ants, pollinators and ants inside flowers, and quantified the effects on plant fitness. Based on the Distraction Hypothesis, we predicted that preventing extrafloral nectar secretion should result in fewer ants active at extrafloral nectaries, more ants inside flowers and a higher proportion of flowers occupied by ants, leading to ant‐pollinator conflict, with reduced pollinator visitation and reduced plant fitness. Overall ant activity inside flowers was low. Preventing extrafloral nectar secretion through clogging reduced the number of ants patrolling extrafloral nectaries, significantly increased the proportion of flowers occupied by ants from 6.1% to 9.7%, and reduced plant reproductive output through a 12% increase in the probability of fruit abortion. No change in the numbers of ants or pollinators inside flowers was observed. This is the first support for the Distraction Hypothesis obtained under field conditions, showing ecological and plant fitness benefits of the distracting function of extrafloral nectar during anthesis. Synthesis. Our study provides the first field experimental support for the Distraction Hypothesis, suggesting that extrafloral nectaries located close to flowers may bribe ants away from reproductive structures during the crucial pollination period, reducing the probability of ant occupation of flowers, reducing ant‐pollinator conflict and increasing plant reproductive success. Our study provides the first field experimental support for the Distraction Hypothesis, suggesting that extrafloral nectaries located close to flowers may bribe ants away from reproductive structures during the crucial pollination period, reducing the probability of ant occupation of flowers, reducing ant‐pollinator conflict and increasing plant reproductive success.

Two new species of Turnera (Passifloraceae s. l., Turneroideae), T. acangatinga and T. ibateguara, from the Atlantic Forest of northeastern Brazil, are described. Some specimens of T. acangatinga were originally treated as T. annularis. Turnera ibateguara was recently considered part of T. pernambucensis, a species which also occurs in the Atlantic Forest in northeastern Brazil, and which was, until then, only known from the type material. In this study, recent collections of T. pernambucensis in the state of Alagoas revealed that both are distinct species and that have been mistakenly identified so far. We provide data on morphology, distribution, habitat, and phenophases, a preliminary conservation status assessment for the three species, and a key to the species with capitate inflorescences from Alagoas and Pernambuco.