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The genus Utricularia includes around 250 species of carnivorous plants, commonly known as bladderworts. The generic name Utricularia was coined by Carolus Linnaeus in reference to the carnivorous organs (Utriculus in Latin) present in all species of the genus. Since the formal proposition by Linnaeus, many species of Utricularia were described, but only scarce information about the biology for most species is known. All Utricularia species are herbs with vegetative organs that do not follow traditional models of morphological classification. Since the formal description of Utricularia in the 18th century, the trap function has intrigued naturalists. Historically, the traps were regarded as floating organs, a common hypothesis that was maintained by different botanists. However, Charles Darwin was most likely the first naturalist to refute this idea, since even with the removal of all traps, the plants continued to float. More recently, due mainly to methodological advances, detailed studies on the trap function and mechanisms could be investigated. This review shows a historical perspective on Utricularia studies which focuses on the traps and body organization.

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BACKGROUND AND AIMS: Floral food bodies (including edible trichomes) are a form of floral reward for pollinators. This type of nutritive reward has been recorded in several angiosperm families: Annonaceae, Araceae, Calycanthaceae, Eupomatiaceae, Himantandraceae, Nymphaeaceae, Orchidaceae, Pandanaceae and Winteraceae. Although these bodies are very diverse in their structure, their cells contain food material: starch grains, protein bodies or lipid droplets. In Pinguicula flowers, there are numerous multicellular clavate trichomes. Previous authors have proposed that these trichomes in the Pinguicula flower play the role of 'futterhaare' ('feeding hairs') and are eaten by pollinators. The main aim of this study was to investigate whether the floral non-glandular trichomes of Pinguicula contain food reserves and thus are a reward for pollinators. The trichomes from the Pinguicula groups, which differ in their taxonomy (species from the subgenera: Temnoceras, Pinguicula and Isoloba) as well as the types of their pollinators (butterflies/flies and bees/hummingbirds), were examined. Thus, it was determined whether there are any connections between the occurrence of food trichomes and phylogeny position or pollination biology. Additionally, we determined the phylogenetic history of edible trichomes and pollinator evolution in the Pinguicula species. METHODS: The species that were sampled were: Pinguicula moctezumae, P. esseriana, P. moranensis, P. emarginata, P. rectifolia, P. mesophytica, P. hemiepiphytica, P. agnata, P. albida, P. ibarrae, P. martinezii, P. filifolia, P. gigantea, P. lusitanica, P. alpina and P. vulgaris. Light microscopy, histochemistry, and scanning and transmission electron microscopy were used to address our aims with a phylogenetic perspective based on matK/trnK DNA sequences. KEY RESULTS: No accumulation of protein bodies or lipid droplets was recorded in the floral non-glandular trichomes of any of the analysed species. Starch grains occurred in the cells of the trichomes of the bee-/fly-pollinated species: P. agnata, P. albida, P. ibarrae, P. martinezii, P. filifolia and P. gigantea, but not in P. alpina or P. vulgaris. Moreover, starch grains were not recorded in the cells of the trichomes of the Pinguicula species that have long spurs, which are pollinated by Lepidoptera (P. moctezumae, P. esseriana, P. moranensis, P. emarginata and P. rectifolia) or birds (P. mesophytica and P. hemiepihytica), or in species with a small and whitish corolla that self-pollinate (P. lusitanica). The results on the occurrence of edible trichomes and pollinator syndromes were mapped onto a phylogenetic reconstruction of the genus. CONCLUSION: Floral non-glandular trichomes play the role of edible trichomes in some Pinguicula species (P. agnata, P. albida, P. ibarrae, P. martinezii, P. filifolia and P. gigantea), which are mainly classified as bee-pollinated species that had originated from Central and South America. It seems that in the Pinguicula that are pollinated by other pollinator groups (Lepidoptera and hummingbirds), the non-glandular trichomes in the flowers play a role other than that of a floral reward for their pollinators. Edible trichomes are symplesiomorphic for the Pinguicula species, and thus do not support a monophyletic group such as a synapomorphy. Nevertheless, edible trichomes are derived and are possibly a specialization for fly and bee pollinators by acting as a food reward for these visitors.

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The insectivorous genus Pinguicula occurs along the whole Andean mountain chain from Colombia-Venezuela in the north to Tierra del Fuego in the south with a short interruption in the Peruvian-Chilean desert range. This paper describes a new and striking species of Pinguicula that occurs in the south-eastern part of the Amotape-Huancabamba Zone in north Peru. It grows either as a lithophyte on moist rocks or as an epiphyte on Polylepis multijuga Pilg. in the wet highlands of the Cordillera Central. Pinguicula rosmarieae Casper, Bussmann & T.Henning, sp. nov. is clearly distinguished by a basal rosette of ovate-obovate leaves spread out flat on the ground and especially by a two-partite corolla with a straight uniform tube-spur complex, two features unknown from other Andean Pinguicula species. The morphological similarity to P. calyptrata Kunth is discussed and the habitat and distribution of P. rosmarieae are characterised.

ResumenEl género insectívoro Pinguicula se encuentra a lo largo Andes desde Colombia y Venezuela en el norte hasta Tierra Fuego en el sur, con una breve interrupción en el los desiertos peruano-chilenos. Este artículo describe una nueva y distintiva especie de Pinguicula que se encuentra en la parte sur de la zona Amotape-Huancabamba en el norte del Perú. Puede crecer tanto como litófita sobre rocas húmedas o como epífita sobre Polylepis multijuga Pilg. en las tierras altas y húmedas de la Cordillera Central. Pinguicula rosmarieae Casper, Bussmann & T.Henning, sp. nov. se distingue claramente por tener una roseta basal de hojas ovadas-obovadas, postradas sobre el suelo y, especialmente, por la corola bipartida con un espolón uniforme recto, una combinación de características desconocidas de otras especies andinas de Pinguicula. Se discute la similitud morfológica con P. calyptrata Kunth y se caracterizan el hábitat y la distribución de P. rosmarieae.

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Utricularia amethystina Salzm. ex A.St.-Hil. & Girard (Lentibulariaceae) is a highly polymorphic carnivorous plant taxonomically rearranged many times throughout history. Herein, the complete chloroplast genomes (cpDNA) of three U. amethystina morphotypes: purple-, white-, and yellow-flowered, were sequenced, compared, and putative markers for systematic, populations, and evolutionary studies were uncovered. In addition, RNA-Seq and RNA-editing analysis were employed for functional cpDNA evaluation. The cpDNA of three U. amethystina morphotypes exhibits typical quadripartite structure. Fine-grained sequence comparison revealed a high degree of intraspecific genetic variability in all morphotypes, including an exclusive inversion in the psbM and petN genes in U. amethystina yellow. Phylogenetic analyses indicate that U. amethystina morphotypes are monophyletic. Furthermore, in contrast to the terrestrial Utricularia reniformis cpDNA, the U. amethystina morphotypes retain all the plastid NAD(P)H-dehydrogenase (ndh) complex genes. This observation supports the hypothesis that the ndhs in terrestrial Utricularia were independently lost and regained, also suggesting that different habitats (aquatic and terrestrial) are not related to the absence of Utricularia ndhs gene repertoire as previously assumed. Moreover, RNA-Seq analyses recovered similar patterns, including nonsynonymous RNA-editing sites (e.g., rps14 and petB). Collectively, our results bring new insights into the chloroplast genome architecture and evolution of the photosynthesis machinery in the Lentibulariaceae.

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Genlisea violacea is a Brazilian endemic carnivorous plant species distributed in the cerrado biome, mainly in humid environments, on sandy and oligotrophic soil or wet rocks. Studies on reproductive biology or pollination in the Lentibulariaceae are notably scarce; regarding the genus Genlisea, the current study is the first to show systematic and standardised research on reproductive biology from field studies to describe the foraging of visiting insects and determine the effective pollinators of Genlisea. We studied two populations of G. violacea through the observation of flower visitors for 4 months of the rainy and dry seasons. Stigmatic receptivity, pollen viability, and breeding system were evaluated together with histochemistry and morphological analyses of flowers. The flowers showed stigmatic receptivity of 100% in open buds and mature flowers, reducing to 80% for senescent flowers. Nearly 80% of pollen grains are viable, decreasing to 40-45% after 48 h. Nectar is produced by glandular trichomes inside the spur. Two bee species are effective pollinators: one of the genus Lasioglossum (subgenus Dialictus: Halictidae) and the other of the genus Ceratina (subgenus Ceratinula: family Apidae). Moreover, bee-like flies of the Syrphidae family may also be additional pollinators. Genlisea violacea is an allogamous and self-compatible species. The differences in flower-visiting fauna for both populations can be attributed to factors such as climate, anthropogenic effect, seasonal factors related to insects and plants, as well as the morphological variation of flowers in both populations.

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Utricularia cornigera and Utricularia nelumbifolia are giant, aquatic-epiphytic species of carnivorous bladderwort from southeastern Brazil that grow in the central 'urns' of bromeliads. Both species have large, colourful flowers. The main aim of our study is to ascertain whether the prominent floral palate of U. cornigera and U. nelumbifolia functions as an unguentarius-i.e. an organ that bears osmophores. Floral tissues of both species were investigated using light microscopy, scanning electron microscopy, transmission electron microscopy and histochemistry. Floral palates of U. cornigera and U. nelumbifolia provide clear visual signals for pollinating insects. In both species, the palate possesses diverse micro-morphology, comprising unicellular, conical to villiform papillae and multicellular, uniseriate, glandular trichomes that frequently display terminal branching. The most characteristic ultrastructural feature of these papillae was the presence of relatively large, polymorphic plastids (chromoplasts) containing many plastoglobuli. Similar plastids are known to occur in the fragrance-producing (osmophores) and oil-producing (elaiophores) tissues of several orchid species. Thus, these palate papillae may play a key role in providing the olfactory stimulus for the attraction of insect pollinators. Nectariferous trichomes were observed in the floral spurs of both species, and in U. nelumbifolia, free nectar was also recorded. The location, micro-morphology, anatomy and ultrastructure of the floral palate of the two species investigated may thus indicate that the palate functions as an unguentarius. Furthermore, the flowers of these taxa, like those of U. reniformis, have features consistent with bee pollination.

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ResumenLas plantas carnívoras del género Utricularia, capturan un amplio ámbito de organismos acuáticos. La mayoría se desarrollan en ambientes con carencias de nutrientes y tienen la capacidad de cambiar las condiciones de su microambiente. El objetivo de la presente investigación fue estudiar la selectividad, en la captura de zooplancton por Utricularia foliosa en la Ciénaga de Paredes; entre febrero y noviembre 2014. Se determinó si existe selección en los recursos alimentarios de la planta con el índice de Czekanowski y el grado de selección, a través de los índices de Savage e Ivlev. Se estableció la relación existente entre los patrones de captura y selección del zooplancton, con variables físicoquímicas y la disponibilidad de nutrientes en la Ciénaga. En nuestros resultados, el índice de Czekanowski evidenció selección, en los recursos alimentarios de la planta, durante todo el pulso de inundación, con valores entre 0.28 y 0.41. Se encontró selección positiva significativa, con el índice de Savage, para los géneros Lecane sp., Alona sp., Ceriodaphnia sp., y Bosmina sp. (p < 0.05). Se obtuvieron resultados similares, con el índice de Ivlev. La intensidad en la selección de los géneros varió entre periodos hidrológicos y por periodos de aguas altas y bajas. Se identificó que los cambios en la concentración de amoniaco y nitrato, además de la variación en la conductividad eléctrica de la ciénaga influencian la captura que realiza U. foliosa. Este es el primer trabajo que permite dar un acercamiento al entendimiento de la selección de recursos alimentarios dentro de la especie a través de índices de selectividad y uno de los pocos para el género.

Abstract:Utricularia is a genus of carnivorous plants that capture a wide range of aquatic organisms. Most of these plants grow in environments with nutrients deficiency and have the ability to change the conditions of their microenvironment. The aim of this research was to study the selectivity in the zooplankton capture by Utricularia foliosa in the Ciénaga de Paredes. Our study was undertaken between February and November, 2014. We tried to determine if there is selection in the plant's food resources by the Czekanowski's index, and the selection degree by the Savage and Ivlev's indexes. Additionally, we studied the possible relation between the patterns of zooplankton capture and selection, with physicochemical variables in the swamp. The Czekanowski's index showed a food selection in plant resources throughout the flood pulse, with values between 0.28 and 0.41. We also found a significant positive selection with Savage's index for Lecane sp., Alona sp., Ceriodaphnia sp., and Bosmina sp. (p < 0.05); similar results were obtained with Ivlev's index. The intensity in the selection of each captured genus varied significantly between hydrological periods and between high and low water levels in the swamp. It was possible to identify some changes in the ammonia and nitrate concentration and some variability in the electric conductivity of the swamp, which influenced the captures made by U. foliosa. This is the first paper that allows an approach to understand the selection of food resources for the species, using a selectivity index, and one of the few for the genus. Rev. Biol. Trop. 64 (3): 1297-1310. Epub 2016 September 01.

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Las plantas carnívoras acuáticas del género Utricularia (Lentibulariaceae), capturan pequeños organismos acuáticos, como rotíferos, copépodos, y cladóceros, por medio de estructuras anatómicas semejantes a vesículas. Este estudio tuvo como objetivo determinar el tamaño y tipo de presas de U. gibba y U. inflata, recolectados en un pequeño lago y en un pastizal inundado, respectivamente, en la localidad de la Península de Paria (Estado Sucre, Venezuela). La acidez, conductividad, cantidad de oxígeno disuelto y la salinidad del agua se midieron in situ para cada sitio de muestreo; también se tomaron muestras de agua para determinar niveles de N-Kjeldahl, P-total, Na+, K+, Ca++, Mg++ y Cl-. Se midieron cincuenta vesículas de cada especie y se analizaron sus contenidos. Los valores de N-Kjeldahl y P-total fueron similares en ambos sitios, y semejantes a los conocidos para ecosistemas eutróficos; sin embargo, las concentraciones de Na+, K+, Ca++, Mg++ y los valores in situ de los parámetros del agua fueron más altos en el pastizal inundado. Las vesículas contenían los siguientes grupos de zooplancton: rotíferos, cladóceros, copépodos, anélidos, rizópodos, e insectos; y las siguientes divisiones de fitoplancton: Bacillariophyta, Chlorophyta, Cyanophyta y Euglenophyta. Las vesículas presentadas por U. inflata fueron menores en tamaño y en cantidad, pero mayores en abundancia y riqueza total de morfoespecies algales y animales en comparación con U. gibba. La similitud en cuanto a composición de presas a nivel taxonómico, entre las dos especies de plantas carnívoras, fue baja.

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