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The evolution of key innovations, novel traits that promote diversification, is often seen as major driver for the unequal distribution of species richness within the tree of life. In this study, we aim to determine the factors underlying the extraordinary radiation of the subfamily Bromelioideae, one of the most diverse clades among the neotropical plant family Bromeliaceae. Based on an extended molecular phylogenetic data set, we examine the effect of two putative key innovations, that is, the Crassulacean acid metabolism (CAM) and the water-impounding tank, on speciation and extinction rates. To this aim, we develop a novel Bayesian implementation of the phylogenetic comparative method, binary state speciation and extinction, which enables hypotheses testing by Bayes factors and accommodates the uncertainty on model selection by Bayesian model averaging. Both CAM and tank habit were found to correlate with increased net diversification, thus fulfilling the criteria for key innovations. Our analyses further revealed that CAM photosynthesis is correlated with a twofold increase in speciation rate, whereas the evolution of the tank had primarily an effect on extinction rates that were found five times lower in tank-forming lineages compared to tank-less clades. These differences are discussed in the light of biogeography, ecology, and past climate change.

Objectives In vitro seed propagation can enhance plant species growth and enable the rapid production of seedlings while preserving genetic variability. This study aimed to develop in vitro seed propagation and acclimatization protocols for Dyckia rariflora to support conservation efforts of this bromeliad endemic to ferruginous campos rupestres. Seed germination and plant growth were tested using MS (Murashige & Skoog) culture medium with varying salt concentrations, sucrose levels, and the presence or absence of polyvinylpyrrolidone (PVP). Following these treatments, seedlings were acclimatized after removal from the controlled environment. Results Germination rates varied between 65 and 90%, unaffected by treatment. The highest germination speed index was in half MS salts without PVP, while full MS salts, sucrose, and PVP slowed germination. Half MS salts resulted in seedlings with greater height, more leaves, and longer roots. Complete MS salts were less effective. No seed oxidation was observed. After 120 days of acclimatization, survival rates exceeded 70%, with plants in half MS salts and 15 g sucrose showing the best growth. In vitro propagation of D. rariflora is viable for large-scale plant production, with half MS salt and sucrose concentrations, without PVP, recommended for better plant growth and cost reduction.

Epiphytic bromeliads frequently present a structure formed by the overlapping of leaf bases where water and nutrients can be accumulated and absorbed, called tank. However, this structure is not present during the juvenile ontogenetic phase, leading to differences in nutrient acquisition strategies. Recent studies have shown a high capacity of the bromeliad Vriesea gigantea, an epiphytic tank-forming bromeliad, to absorb urea by their leaves. Since plant aquaporins can facilitate the diffusion of urea through the membranes, we cloned three foliar aquaporin genes, VgPIP1; 1, VgPIP1;2 and VgTIP2;1 from V. gigantea plants. Through functional studies, we observed that besides water, VgTIP2;1 was capable of transporting urea while VgPIP1;2 may facilitate ammonium/ammonia diffusion. Moreover, aiming at identifying urea and ammoniuminduced changes in aquaporin expression in leaves of juvenile and adult-tank plants, we showed that VgPIP1;1 and VgPIP1;2 transcripts were up-regulated in response to either urea or ammonium only in juvenile plants, while VgTIP2;1 was up-regulated in response to urea only in adult-tank plants. Thereby, an ontogenetic shift from juvenile to adult-tank-forming-plant appears to occur with metabolic changes regarding nitrogen metabolism regulation. Investigating urea metabolism in wild species that naturally cope with organic N sources, such as V. gigantea, may provide the knowledge to modify nitrogen use efficiency of crop plants.

The large Neotropical family Bromeliaceae presents an outstanding example of adaptive radiation in plants, containing a wide range of terrestrial and epiphytic life-forms occupying many distinct habitats. Diversification in bromeliads has been linked to several key innovations, including water- and nutrient-impounding phytotelmata, absorptive epidermal trichomes, and the water-conserving mode of photosynthesis known as crassulacean acid metabolism (CAM). To clarify the origins of CAM and the epiphytic habit, we conducted a phylogenetic analysis of nucleotide sequences for 51 bromeliad taxa by using the plastid loci matK and the rps16 intron, combined with a survey of photosynthetic pathway determined by carbon-isotope ratios for 1,873 species representing 65% of the family. Optimization of character-states onto the strict consensus tree indicated that the last common ancestor of Bromeliaceae was a terrestrial C3 mesophyte, probably adapted to moist, exposed, nutrient-poor habitats. Both CAM photosynthesis and the epiphytic habit evolved a minimum of three times in the family, most likely in response to geological and climatic changes in the late Tertiary. The great majority of epiphytic forms are now found in two lineages: in subfamily Tillandsioideae, in which C3 photosynthesis was the ancestral state and CAM developed later in the most extreme epiphytes, and in subfamily Bromelioideae, in which CAM photosynthesis predated the appearance of epiphytism. Subsequent radiation of the bromelioid line into less xeric habitats has led to reversion to C3 photosynthesis in some taxa, showing that both gain and loss of CAM have occurred in the complex evolutionary history of this family.

Although there is unambiguous evidence for vascular epiphytic plants to be limited by insufficient water and nutrient supply under natural conditions, it is an open debate whether they are primarily phosphorus (P) or nitrogen (N) limited. Plant (15) N fractionation and foliar N : P stoichiometry of a tank epiphyte (Vriesea sanguinolenta), and its response to combined N-P fertilization, were studied under semi-natural conditions over 334 d to clarify the type of nutrient limitation. Plants collected in the field and experimental plants with limited nutrient supply showed significant plant (15) N fractionation (mean 5‰) and plant N : P ratios of c. 13.5. Higher relative growth rates and declines in plant (15) N fractionation (0.5‰) and in foliar N : P ratios to 8.5 in the high N-P treatment indicated that these epiphytes were P limited in situ. The critical foliar N : P ratio was 10.4, as derived from the breakpoint in the relationship between plant (15) N fractionation and foliar N : P. We interpret the widespread (15) N depletion of vascular epiphytes relative to their host trees as deriving from (15) N fractionation of epiphytes as a result of P limitation. High foliar N : P ratios (> 12) corroborate widespread P limitation (or co-limitation by N and P) of epiphytic bromeliads and, possibly, other epiphyte species.

Bromeliads are able to occupy some of the most nutrient-poor environments especially because they possess absorptive leaf trichomes, leaves organized in rosettes, distinct photosynthetic pathways [C , Crassulacean acid metabolism (CAM) or facultative C -CAM], and may present an epiphytic habit. The more derived features related to these traits are described for the Tillandsioideae subfamily. In this context, the aims of this study were to evaluate how terrestrial predators contribute to the nutrition and performance of bromeliad species, subfamilies and ecophysiological types, whether these species differ in their ecophysiological traits and whether the physiological outcomes are consistent among subfamilies and types (e.g. presence/absence of tank, soil/tank/atmosphere source of nutrients, trichomes/roots access to nutrients). Isotopic ( N-enriched predator faeces) and physiological methods (analyses of plant protein, amino acids, growth, leaf mass per area and total N incorporated) in greenhouse experiments were used to investigate the ecophysiological contrasts between Tillandsioideae and Bromelioideae, and among ecophysiological types when a predatory anuran contributes to their nutrition. It was observed that Bromelioideae had higher concentrations of soluble protein and only one species grew more (Ananas bracteatus), while Tillandsioideae showed higher concentrations of total amino acids, asparagine and did not grow. The ecophysiological types that showed similar protein contents also had similar growth. Additionally, an ordination analysis showed that the subfamilies and ecophysiological types were discrepant considering the results of the total nitrogen incorporated from predators, soluble protein and asparagine concentrations, relative growth rate and leaf mass per area. Bromeliad subfamilies showed a trade-off between two strategies: Tillandsioideae stored nitrogen into amino acids possibly for transamination reactions during nutritional stress and did not grow, whereas Bromelioideae used nitrogen for soluble protein production for immediate utilization, possibly for fast growth. These results highlight that Bromeliaceae evolution may be directly associated with the ability to stock nutrients.

The Bromeliaceae family is one of the most morphologically diverse families with a pantropical distribution. To schedule an appropriate flowering time for bromeliads, ethylene is commonly used to initiate flower development in adult plants. However, the mechanism by which ethylene induces flowering in adult bromeliads remains unknown. Here, we identified an APETALA2 (AP2)-like gene, AfAP2-1, in Aechmea fasciata. AfAP2-1 contains two AP2 domains and is a nuclear-localized protein. It functions as a transcriptional activator, and the activation domain is located in the C-terminal region. The expression level of AfAP2-1 is higher in juvenile plants than in adult plants, and the AfAP2-1 transcript level was rapidly and transiently reduced in plants treated with exogenous ethylene. Overexpression of AfAP2-1 in Arabidopsis thaliana results in an extremely delayed flowering phenotype. These results suggested that AfAP2-1 responds to ethylene and is a putative age-dependent flowering regulator in A. fasciata.

Salt stress is one of the most severe abiotic stresses affecting plant growth and development. The application of silicon (Si) is an alternative that can increase the tolerance of plants to various types of biotic and abiotic stresses. The objective was to evaluate salt stress’s effect in vitro and Si’s mitigation potential on Aechmea blanchetiana plants. For this purpose, plants already established in vitro were transferred to a culture medium with 0 or 14 µM of Si (CaSiO 3 ). After growth for 30 days, a stationary liquid medium containing different concentrations of NaCl (0, 100, 200, or 300 µM) was added to the flasks. Anatomical and physiological analyses were performed after growth for 45 days. The plants cultivated with excess NaCl presented reduced root diameter and effective photochemical quantum yield of photosystem II (PSII) (ΦPSII) and increased non-photochemical dissipation of fluorescence (qN). Plants that grew with the presence of Si also had greater content of photosynthetic pigments and activity of the enzymes of the antioxidant system, as well as higher values of maximum quantum yield of PSII (F V /F M ), photochemical dissipation coefficient of fluorescence (qP) and fresh weight bioaccumulation of roots and shoots. The anatomical, physiological and biochemical responses, and growth induced by Si mitigated the effect of salt stress on the A. blanchetiana plants cultivated in vitro , which can be partly explained by the tolerance of this species to grow in sandbank ( Restinga ) areas.

Conventional ecological theory predicts that predators affect nutrient cycling by decreasing the abundance or activity of prey. By using a predator-detritivore-detritus food chain in bromeliads, we show that predators can increase nutrient cycling by a previously undescribed, but broadly applicable, mechanism: reducing nutrient export by prey emigration. Contrary to expectations, predation on detritivores increases detrital nitrogen uptake by bromeliads. Predation reduces detritivore emergence and hence export of nitrogen from the system. Detritivores therefore benefit their host plant, but only when predators are present. More generally, our results show that predator loss or extinction can dramatically and unexpectedly affect ecosystem functioning.

$\\bullet$In contrast to terrestrial plants, epiphytic tank bromeliads take up nutrients mainly over their tank leaf surface. The form in which nutrients are available in the tanks is determined by the source and the complex interplay between tank microbes, which transform them and the epiphytes that take them up.$\\bullet$To elucidate the importance of different nitrogenous compounds for the nitrogen (N) nutrition of Vriesea gigantea from the Atlantic Rainforest, Brazil, N transformation processes in tank water as well as foliar uptake rates were estimated by15N labelling techniques.$\\bullet$Microorganisms actively transformed N compounds in the tank. Specifically, organic N compounds were rapidly mineralized to NH4 +, while nitrification was negligible.$\\bullet$Plants took up both organic and inorganic N forms, with a clear preference for$NH_4^{-}$. NH4 +comprised the largest and, because of fast mineralization rates, the most constant dissolved N pool in the tank water. Excretion of ureases by the plants together with an unusual uptake kinetic for urea also suggests that urea may be potentially important as an N source.