Ant visitation to extrafloral nectaries decreases herbivory and increases fruit set in Chamaecrista debilis (Fabaceae) in a Neotropical savanna

Studies of ant-plant relationships elucidate how top-down effects of the third trophic level can affect the biomass, richness, and/or species composition of plants. Although widespread in the neotropics, few studies have so far examined the direct effects of ants on plant fitness. Here, through experimental manipulation (ant-exclusion) under natural conditions, we examined the effect of ant visitation to extrafloral nectaries on leaf herbivory and fruit set in Chamaecrista debilis in the Brazilian savanna. As opposed to other Chamaecrista species, our results showed that visiting ants (15 species) significantly reduce herbivory and increase fruit set by more than 50% compared to plants from which ants were excluded. This mutualistic system is facultative in nature, and corroborates the potential beneficial role of exudate-feeding ants as anti-herbivore agents of tropical plants. (C) 2010 Elsevier GmbH. All rights reserved.

Morphology, ontogeny and structure of the stipular nectaries in Caamembeca spectabilis (Polygalaceae)

ABSTRACT Nodal glands are found in one third of the Polygalaceae genera and have valuable taxonomic, ecological and evolutionary significance. In Brazil, they occur in five of the eleven genera already registered. However, there is still a controversy regarding the origin of these structures. The objective of this study was to characterize the morphology and the origin of nodal glands inCaamembeca spectabilis, in order to increase the structural and functional knowledge of these glands in the genera. Samples of nodal regions were collected, fixed and processed according to the methods of light microscopy and electron scanning. Ants were observed and identified along the stem axis. The glucose in exudate allows us to classify these glands as extrafloral nectaries. They are located in pairs on the nodal region. However, its origin is in the leaf trace. In the longitudinal section, the nectaries were present in the apex of cells with anticlinal walls impregnated with suberin, which represents the first record for the family. In this region there is also the formation of a hole by lysis. The secretory tissue is surrounded by phloem. Xylem vessels were observed only on the basis of the nectary, where there are also idioblasts with crystals in druse type. We have studied the ontogeny of the glands nodal in Caamembeca spectabilis and unveiled that these glands are linked to the leaves as stipular nectaries. In addition, the new findings presented here may add support for the understanding of morphology and anatomy of nodal glands in Caamembeca.

New record of predatory ladybird beetle (Coleoptera, Coccinellidae) feeding on extrafloral nectaries

New record of predatory ladybird beetle (Coleoptera, Coccinellidae) feeding on extrafloral nectaries. Feeding by Exoplectra miniata (Germar) on extrafloral nectaries of Inga edulis Mart. was observed in Nova Friburgo, Rio de Janeiro, Brazil. This is the first record of this behavior for Exoplectrini.

When are ant-attractant devices a worthwhile investment? Vicia faba extrafloral nectaries and Lasius niger ants

Most studies aiming to determine the beneficial effect of ants on plants simply consider the effects of the presence or exclusion of ants on plant yield. This approach is often inadequate, however, as ants interact with both non-tended herbivores and tended Homoptera. Moreover, the interaction with these groups of organisms is dependent on ant density, and these functional relationships are likely to be non-linear. A model is presented here that segregates plant herbivores into two categories depending on the sign of their numerical response to ants (myrmecophiles increase with ants, non-tended herbivores decline). The changes in these two components of herbivores with increasing ant density and the resulting implications for ant-plant mutualisms are considered. It emerges that a wide range of ant densities needs to be considered as the interaction sign (mutualism or parasitism) and strength is likely to change with ant density. The model is used to interpret the results of an experimental study that varied levels of Aphis fabae infestation and Lasius niger ant attendance on Vicia faba bean plants. Increasing ant density consistently reduced plant fitness and thus, in this location, the interaction between the ants and the plant can be considered parasitic. In the Vicia faba system, these costs of ants are unlikely to be offset by other beneficial agents (e.g., parasitoids), which also visit extrafloral nectaries.

Genetic analysis reveals a novel locus in Vicia faba decoupling pigmentation in the flower from that in the extra-floral nectaries

The genetics of the stipule spot pigmentation (SSP) in faba bean (Vicia faba L.) was studied using four inbred lines, of which Disco/2 was zero-tannin (zt2) with colourless stipule spots, ILB938/2 was normal-tannin (ZT2) with colourless stipule spots, and both Aurora/2 and Mélodie/2 were ZT2 with coloured stipule spots. Crosses Mélodie/2 × ILB 938/2, Mélodie/2 × Disco/2, ILB 938/2 × Aurora/2 and ILB 938/2 × Disco/2 (A, B, C and D, respectively) were prepared, along with reciprocals and backcrosses, and advanced through single-seed descent. All F1 hybrid plants had pigmented stipule spots, and in the F2 generation, the segregation ratio fit 3 coloured:1 colourless in crosses A, B and C and 9:7 in cross D. In the F3 generation, the ratio fit 5:3 in crosses A and C and 25:39 in cross D, and in the F4 generation, 9:7 in cross A. SSP was linked to the zero-tannin characteristics (white flower) only in cross B. The results show that coloured stipule spot is dominant to colourless and that colouration is determined by two unlinked complementary recessive genes. We propose the symbols ssp2 for the gene associated with zt2 in Disco/2 and ssp1 for the gene not associated with tannin content in ILB938/2. The novel ssp1 locus was mapped at F5 in cross ‘A’ using Medicago truncatula-derived single-nucleotide polymorphism and was on chromosome 1 of faba bean, in a well-conserved region of M. truncatula chromosome 5 containing some candidate Myb and basic helix–loop–helix transcription factor genes.

Do extrafloral nectaries present a defensive role against herbivores in two species of the family Bignoniaceae in a Neotropical savannas?

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Leaf glands act as nectaries in Diplopterys pubipetala (Malpighiaceae)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Nectaries and reproductive biology of Croton sarcopetalus (Euphorbiaceae)

Flower morphology, nectary structure, nectar chemical composition, breeding system, floral visitors and pollination were analysed in Croton sarcopetalus, a diclinous-monoecious shrub from Argentina. Male flowers have five receptacular nectaries, with no special vascular bundles, that consist of a uniserial epidermis with stomata subtended by a secretory parenchyma. Female flowers bear two different types of nectaries: inner (IN) and outer (ON) floral nectaries. IN, five in all, are structurally similar to the nectaries of male flowers. The five ON are vascularized, stalked, and composed of secretory, column-shaped epidermal cells without stomata subtended by secretory and ground parenchyma. In addition, ON act as post-floral nectaries secreting nectar during fruit ripening. Extrafloral nectaries (EFN) are located on petioles, stipules and leaf margins. Petiolar EFN are patelliform, stalked and anatomically similar to the ON of the female flower. Nectar sampled from all nectary types is hexose dominant, except for the ON of the female flower at the post-floral stage that is sucrose dominant. The species is self-compatible, but geitonogamous fertilization is rarely possible because male and female flowers are not usually open at the same time in the same individual, i.e. there is temporal dioecism. Flowers are visited by 22 insect species, wasps being the most important group of pollinators. No significant differences were found in fruit and seed set between natural and hand pollinated flowers. This pattern indicates that fruit production in this species is not pollen/pollinator limited and is mediated by a wide array of pollinators. (C) 2001 the Linnean Society of London.

Floral anatomy of Bromeliaceae, with particular reference to the evolution of epigyny and septal nectaries in commelinid monocots

Floral anatomy is described in ten genera of Bromeliaceae, including three members of subfamily Bromelioideae, three Tillandsioideae, and four genera of the polyphyletic subfamily Pitcairnioideae (including Brocchinia, the putatively basal genus of Bromeliaceae). Bromeliaceae are probably unique in the order Poales in possessing septal nectaries and epigynous or semi-epigynous flowers. Evidence presented here from floral ontogeny, vasculature, and the relative positions of nectary and ovules indicates that there could have been one or more reversals to apparent hypogyny in Bromeliaceae, although this hypothesis requires a better-resolved phylogeny. Such evolutionary reversals probably evolved in response to specialist pollinators, and in conjunction with other aspects of floral morphology of Bromeliaceae, such as the petal appendages of some species. The ovary is initiated in an inferior position even in semi-epigynous or hypogynous species. The ovary of all so-called hypogynous Bromeliaceae is actually semi-inferior, because the septal nectary is infralocular; in these species the nectaries have a labyrinthine surface and many vascular bundles. Brocchinia differs from most other fully epigynous species in that each carpel is secretory at the apex and reproductive, rather than secretory, at the base.

Morphological patterns of extrafloral nectaries in woody plant species of the Brazilian cerrado

Extrafloral nectaries are nectar-secreting structures that are especially common among the woody flora of the Brazilian cerrado, a savanna-like vegetation. In this study, we provide morphological and anatomical descriptions of extrafloral nectaries (EFNs) occurring on vegetative and reproductive organs of several plant species from the cerrado, and discuss their function and ecological relevance. We describe the morphology and anatomy of EFNs of 40 species belonging to 15 woody families using scanning electron microscopy and light microscopy. We categorise EFNs following a structural-topographical classification, and characterise the vascularised and complex nectaries, amorphous nectaries and secretory trichomes. Fabaceae, Bignoniaceae, Malpighiaceae and Vochysiaceae were the plant families with the majority of species having EFNs. Ten species possess more than one morphotype of gland structure. Observations and experimental field studies in the cerrado support the anti-herbivore role of EFN-gathering ants in this habitat. Additional morphological studies of EFNs-bearing plants, including other growth forms (e.g. herbs and lianas), are being undertaken and will hopefully cast further light on the ecological relevance of these glands in the cerrado, especially with respect to their attractiveness to multiple visitors.

A macro- and micromorphological survey of floral and extrafloral nectaries in the epiphytic cactus Rhipsalis teres (Cactoideae: Rhipsalideae)

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Morphology, development and homologies of the perianth and floral nectaries in Croton and Astraea (Euphorbiaceae-Malpighiales)

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

EXTRAFLORAL NECTARIES IN THE TROPICAL TREE GUAREA-MACROPHYLLA (MELIACEAE)

This paper describes the anatomy and morphology of the complex nectary systems of the tropical tree Guarea macrophylla Vahl (Meliaceae) and presents the first record of extrafloral nectaries occurring on reproductive organs (fruits) of a member of the order Sapindales. The extrafloral nectaries of G. macrophylla occur on petioles, petiolules, the abaxial surface of all leaflets, leaf buds, and over the surface of fruits. All extrafloral nectaries are distinctly raised above the surface. Foraging ants collect extrafloral nectar on Guarea trees both day and night. We suggest that the presence of extrafloral nectaries might be a useful taxonomic character for the identification of Guarea species.

Distribution of extrafloral nectaries in different vegetation types of Amazonian Brazil

Surveys were carried out in terra firme' forest, successional forest, buritirana' (palm vegetation) and shrub canga' (savanna). Extrafloral nectaries (EFNs) were present in 30 plant species belonging to 22 genera and 14 families. Nectary species represented 17.6-53.3% of the species samples in different areas, with local abundances varying from 19.1-50.0%. The percentage of species with EFNs was greater in the flora of the shrub canga than in the terra firme and successional forests. Nectary plants were more abundant in the shrub canga and successional forest. The high abundance of EFNs may be the result of intense foraging activity by ants on plants, leading to the formation of facultative mutualism. -from Authors

Anatomy of the floral nectaries of some neotropical Salacioideae (Celastraceae)

Floral nectaries have contributed to the systematics of different taxonomic groups. Since those of the neotropical genera included in subfamily Salacioideae-Cheiloclinium Miers, Peritassa Miers, Salacia L. and Tontelea Aubl.-have different forms and positions, we explored their anatomy to delimit more precisely the genera of subfamily Salacioideae. Buds and open flowers of six species were treated following the usual techniques in plant anatomy. The obtained data were helpful in characterizing the floral nectary anatomy of the studied species. Furthermore, some features such as form, position and surface of nectaries; form of their epidermal cells; presence and distribution of stomata; occurrence of idioblasts containing druses in the nectariferous parenchyma; and absence of nectary vascularization can contribute to the taxonomy and phylogeny of the Salacioideae studied. In most of the studied species the nectar is probably released by both the stomata and the nectary epidermal surface. In Cheiloclinium cognatum, the structure acknowledged as nectary is actually a vestigial tissue and the functions of attracting and rewarding pollinators has phylogenetically migrated to the stigmatic region. The druses and phenolic substances observed in the nectariferous parenchyma probably help defend flowers against herbivore attacks. The minute size of the nectaries of Salacioideae may explain the absence of vascularization. The floral nectaries of Salacia elliptica are epithelial while those of the other species are mesenchymal. © 2012 Springer-Verlag Wien.