Floral visitors in urban gardens and natural areas: Diversity and interaction networks in a neotropical urban landscape

Because cities concentrate 50% of the world’s population, and are experiencing a re-emergence of urban agriculture, we investigated the influences of urban agriculture and surrounding natural areas on floral visitors (bees, wasps, butterflies and flies) and plant species in San Cristóbal de Las Casas, Mexico.Throughout the frost, dry and rainy seasons of 2015, we sampled floral visitors in nine urban gardens and nine natural areas. We found 210 floral visitor species: 78% pollinators, 18% predators, and 4% florivores. Rarefaction curves showed that natural areas harbor significantly more floral visitor species (148) than home gardens (132). However, the differences in species composition between habitats and seasons highlight the need to view natural areas and home gardens as complementary habitats with which floral visitors interact in varying ways, during successive seasons, to meet different needs. Furthermore, mean species richness of floral visitors was influenced mainly by seasonality, and increased as seasons progressed from the dry, frost season to the rainy season. Nonetheless, some taxa were influenced by both season and habitat type. Floral visitor abundance was influenced by both habitat type and season, with home gardens showing higher abundance across seasons. Moreover, interaction networks for each season were more asymmetric in natural areas than in home gardens. Urban cover in the surrounding landscape influenced in a quadratic way the species number of floral visitors, but not their abundance. Thus, our results are evidence that natural areas surrounding cities and urban agriculture contribute to floral visitor communities and their networks.     

Pollination in Geonoma macrostachys and Three Congeners,G. acaulis,G. gracilis,and G. interrupta

A continuous 15 month study of the floral ecology of four syntopic understorey palm species of Genoma was conducted in Amazonian Peru lowland rainforest. The spicate inflorescences of G. macrostachys, G. acaulis and G. gracilis are strictly protandrous and the plants are functionally dioecious. Data suggest that in G. macrostachys and G. acaulis pollination is based on a mimicry system, the pistillate flowers mimicking the staminate ones in colour, shape and scent. Pollen-collecting meliponine bees (Hymenoptera, Apidae, Meliponinae) and pollen-feeding syrphid flies (Diptera, Syrphidae) which visit inflorescences during both sexual stages are the pollinators of G. macrostachys. Geonoma acaulis is pollinated by small pollen-feeding weevils (Coleoptera, Curculionidae, Derelomini) that visit male and female spikes. Additionally, in G. macrostachys another pollinator type, viz. euglossine bees (Hymenoptera, Apidae, Euglossinae), which are attracted and rewarded by both types of flowers may account for long-distance pollination. The palm G. gracilis shows a very distinct pollination system. Although opportunistic insect visitors are attracted to the inflorescences of this species it seems to be mainly anemophilous because pollen becomes powdery during an thesis. The branched inflorescences of G. interrupta are also protandrous, but unlike the other species of Geonoma observed, staminate and pistillate anthesis of individual flowers are, for the most, overlapping. A broad spectrum of visitors is attracted (bees, wasps, flies, and beetles), which all may act as pollinators. Outcrossing is especially encouraged during the purely female phase at the end of the flowering cycle when there are no more staminate flowers in the inflorescence. Effects on the reproductive biology and population structure of different pollination systems and breeding system are discussed.     

Diverse visitors with various pollinator importance and temporal change in the important pollinators of <Emphasis Type="Italic">Geranium thunbergii</Emphasis> (Geraniaceae)

Assessing pollinator importance of each floral visitor to a plant species is a key to understanding plant–pollinator interaction. The present study examined visitation frequency, pollination efficiency, and pollinator importance of the full range of floral visitors to Geranium thunbergii natural population, by measuring seed-set. During 2 years of observations, the flowers were visited by at least 45 insect species belonging to four orders. Among the main 22 visitor species, 11 species belonging to three orders (Hymenoptera, Diptera, and Lepidoptera) acted as the efficient pollinators. In both years, Hymenoptera, especially bees, was the most important pollinator to G.thunbergii. Thus, the flowers could be considered as bee-pollinated. However, the most important species were not constant between years. The study also documented that the efficient pollinators have larger body sizes. The dish-shaped floral morphology, taxonomically diverse pollinators, and temporal change in the most important pollinators indicate that G.thunbergii–pollinator interaction is a rather generalized system. The results suggest that casual observations of visitation, or even precise measurement of pollinator importance in a single season is insufficient to identify important pollinators.     

Deciduous sapling responses to season and large herbivores in a semi‐arid African savanna

Impacts of large herbivores (>5 kg) on woody plants in African savannas are potentially most severe among plants shorter than 1.6 m. It is well established that severe browsing leads to longer shoots, yet prevents saplings from recruiting into adult size‐classes in African savannas. Increased shoot length, indicating faster shoot growth, is often associated with reduced concentrations of tannins and increased nutrient concentrations, suggesting carbon limitation. We hypothesized that, on average, large herbivores suppress stem height or circumference, but increase shoot length. We also hypothesized that if there were concomitant positive effects on nutrients, or negative effects on tannin concentrations, they would be greatest early in the wet season. We sampled saplings of four deciduous woody species (Acacia grandicornuta, Dichrostachys cinerea, Combretum apiculatum and Grewia flavescens) at different stages of the wet season in a large‐scale, long‐term herbivore exclusion experiment in Kruger National Park, South Africa. Plant height, shoot length and stem circumference were generally not adversely affected by large herbivores, suggesting C limitation is rarely present among deciduous saplings in semi‐arid African savannas, allowing them to tolerate browsing. Time since first rainfall emerged as a predominant factor consistently affecting nutrient and tannin concentrations, rather than large herbivores. Nitrogen and phosphorus generally decreased (by 20–50%), while condensed tannin concentration increased (150–350%) during the wet season, except for one species. We postulate that A. grandicornuta is less prone than other species to accumulating tannins during the wet season because of high investment of C in spines. Although nutrient and tannin concentrations were generally not affected by large herbivores, species‐specific responses were evident very early in the wet season, which is when herbivore populations are most likely to be affected by differential forage quality among plants.     

Pollen partitioning of three species of Convolvulaceae among oligolectic bees in the Caatinga of Brazil

During the rainy season many species of Convolvulaceae bloom simultaneously in the Caatinga of northeast Brazil. In a Caatinga nature reserve we studied pollination and breeding systems of three sympatric species of Convolvulaceae, Ipomoea bahiensis, I.?nil, and Merremia aegyptia, focusing on pollen partitioning among flower visitors and pollen flow. The study showed that only oligolectic bees collected pollen and that these species had different preferences among the three species of Convolvulaceae: pollen of Ipomoea bahiensis, the only self-incompatible species, was collected mainly by Melitoma segmentaria, M.?osmioides, and Melitomella murihirta; pollen of I.?nil by Lithurgus huberi; and that of Merremia aegyptia by Ancyloscelis apiformis and an undescribed species of this genus. Introduced honey bees visited only flowers of Merremia aegyptia, where they were extraordinarily frequent flower visitors. However, they discarded the pollen grains, which led to almost 50% pollen loss. No polylectic bee species compete for pollen with the oligolectic species. Partitioning of pollen diminishes competition for floral resources in this specialized plant?Cpollinator association.     

Spatial predictability and resource specialization of bees (Hymenoptera: Apoidea) at a superabundant, widespread resource

For reciprocal specialization (coevolution) to occur among floral visitors and their host plants the interactions must be temporally and spatially persistent. However, studies repeatedly have shown that species composition and relative abundance of floral visitors vary dramatically at all spatial and temporal scales. We test the hypothesis that, on average, pollen specialist bee species occur more predictably at their floral hosts than pollen generalist bee species. Taxonomic floral specialization reaches its extreme among species of solitary, pollen-collecting bees, yet few studies have considered how pollen specialization by floral visitors influences their spatial constancy. We test this hypothesis using an unusually diverse bee guild that visits creosote bush (Larrea tridentatd), the most widespread, dominant plant of the warm deserts of North America. Twenty-two strict pollen specialist and 80 + generalist bee species visit Larrea for its floral resources. The sites we sampled were separated by 0.5 to > 1450 km, and spanned three distinct deserts and four vegetation zones. We found that species of Larrea pollen specialist bees occurred at more sites and tended to be more abundant than generalists. Surprisingly, spatial turnover was high for both pollen specialist and generalist bee species at all distances, and species composition of samples from sites 1–5 km apart varied as much as repeat samples made at single sites. Nevertheless, the pattern of bee species turnover was not haphazard. As distance among sites increased faunal similarity of sites decreased. Faunal similarities among sites within 250 km of each other were generally greater than if randomly distributed over all sites (the null model). No single ecological category of species (widespread, localized, Larrea pollen specialist, floral generalist) accounted for this spatial predictability. Evidently, concordant local distribution patterns of many ecologically diverse species contribute to the non-random spatial pattern. The ecological dominance of creosote bush does not confer obvious ecological advantages to its specialist floral visitors. Spatial turnover is comparable to that found for bee guilds from other biogeographic regions of the world and is not therefore limited to those bee species that inhabit highly seasonal climates, such as deserts. Philopatry and differences in bloom predictability among sites are probably more important causes for spatial turnover of bee species than are interspecific competition for nest sites or floral resources.     

Insects associated with flowers of weeds of the Botanical Garden of Santiago de Cuba,with emphasis on Hymenoptera

The insect visitors of flowers in nine weeds species were studied in the Botanical Garden of Santiago de Cuba, Eastern Cuba, during 1993 (March-June, spring season) and 1994 (January-March, end of winter and beginning of spring season). About 50 hours of collecting efforts were made at three times (0900-0930 hr in 1993; 0900-0930 hr, 1200-1230 hr and 1500-1530 hr in 1994). More than 140 species of at least 37 families were found; Hymenoptera dominated (with more than a half of specimens), followed by Diptera, Coleoptera and Lepidoptera. Among Hymenoptera, bees (Apoidea) were the largest group, especially Apis mellifera L; followed by wasps (Vespidae, Pompilidae, Sphecidae) and ichneumon flies (Ichneumonidae); Microhymenopterans were not sampled. Hymenopterans of each weed were compared for diversity, similarity, dominant and subdominant species, visitation time, sampling efficiency, etc. Each plant species had a particular Hymenoptera complex, almost one third of which were natural enemies of agricultural pests, and most are believed to be potential pollinators. Closely related species showed similar patterns of daily activity, with a peak at 0900-0930 hr for all plant species. Second grade polynomial equations were the best fitted models to describe the relationships between number of species and number of specimens, and between total number of species and number of samples (R2 = 0.9734 and R2 = 0.9573, p < 0.01). The role of weeds in the biodiversity of the agroecosystems is analyzed; as well as the effectiveness of this collection method to study Hymenoptera.     

Pollination, breeding system and seed abortion in some African acacias

TYBIRK, K., 1993. Pollination, breeding system and seed abortion in some African acacias . Studies of the flower visitors and pollination ecology of Acacia albida Del., A. nilotica (L.) Willd. ex Del. A. tortilis (Forssk.) Hayne and A. Senegal (L.) Willd. in Senegal and Kenya showed a high diversity of floral foragers. One hundred and eighteen taxa of insects mainly from Hymenoptera, Lepidoptera, Coleoptera and Diptera were collected. The most important pollen vectors were bees from the families Megachilidae and Halictidae, and wasps from the families Scoliidae and Eumenidae. Beetles, flies and butterflies were secondary pollen vectors. Diversity and frequency of flower visitors of species with floral nectar ( A. Senegal and A. albida ) were not clearly different from species without floral nectar ( A. tortilis, A. nilotica ). Acacia tortilis was almost exclusively outcrossed (Index of Self Incompatibility = 0.2), with 5.5% of the inflorescences (0.13% of the flowers) developing fruits. The number of seeds per pod was positively correlated with dispersal type. Wind dispersed species had fewer seeds per pod than animal dispersed species. About 5% of the seeds were aborted in A. tortilis and A. nilotica , while 22–48% of the seeds were aborted in A. Senegal, A. ataxacantha DC. and A. polyacantha Willd. Most abortions in the latter three species occur in the proximal end of the pods, indicating selective seed abortion.     

Pollination of Campomanesia phaea (Myrtaceae) by night‐active bees: a new nocturnal pollination system mediated by floral scent

• Bees are the most important diurnal pollinators of angiosperms. In several groups of bees a nocturnal/crepuscular habit developed, yet little is known about their role in pollination and whether some plants are adapted specifically to these bees. We used a multidisciplinary approach to investigate the reproductive biology and to understand the role of nocturnal/crepuscular bees in pollination of Campomanesia phaea (Myrtaceae), popularly named cambuci.
• We studied the floral biology and breeding system of C. phaea. We collected the floral visitors and tested the pollinators' effectiveness. We also determined the floral scents released at night and during daytime, and studied behavioural responses of crepuscular/nocturnal bees towards these scents.
• The flowers of cambuci were self‐incompatible and had pollen as the only resource for flower visitors. Anthesis lasted around 14 h, beginning at 04:30 h at night. The flowers released 14 volatile compounds, mainly aliphatic and aromatic compounds. We collected 52 species of floral visitors, mainly bees. Nocturnal and crepuscular bees (four species) were among the most frequent species and the only effective pollinators. In field bioassays performed at night, nocturnal/crepuscular bees were attracted by a synthetic scent blend consisting of the six most abundant compounds.
• This study describes the first scent‐mediated pollination system between a plant and its nocturnal bee pollinators. Further, C. phaea has several floral traits that do not allow classification into other nocturnal pollination syndromes (e.g. pollinator attraction already before sunrise, with pollen as the only reward), instead it is a plant specifically adapted to nocturnal bees.

     

Growth responses of a woody species to clipping and goat saliva

Studies on the role of mammalian herbivore saliva in plant–animal interactions have mostly focused on graminoid species and bovine saliva. A trial was performed in Botswana with clipping treatments to simulate browsing of shoots and the application of goat saliva on the woody species Combretum apiculatum Sonder (Combretaceae). Treatments were performed during early growing season while shoot growth was rapid, and responses of trees were recorded later in the same season. Clipped shoots with saliva had significantly enhanced shoot growth (tripled in length) and leaf production (2.7 times more leaves) compared to clipped shoots without saliva. However, unclipped shoots still grew more than clipped shoots, with or without saliva treatment.     

Intrafloral patterns of color and scent in Capparis spinosa L. and the ghosts of its selection past

Premise

Capparis spinosa is a widespread charismatic plant, in which the nocturnal floral habit contrasts with the high visitation by diurnal bees and the pronounced scarcity of hawkmoths. To resolve this discrepancy and elucidate floral evolution of C. spinosa, we analyzed the intrafloral patterns of visual and olfactory cues in relation to the known sensory biases of the different visitor guilds (bees, butterflies, and hawkmoths).

Methods

We measured the intrafloral variation of scent, reflectance spectra, and colorimetric properties according to three guilds of known visitors of C. spinosa. Additionally, we sampled visitation rates using a motion-activated camera.

Results

Carpenter bees visited the flowers eight times more frequently than nocturnal hawkmoths, at dusk and in the following morning. Yet, the floral headspace of C. spinosa contained a typical sphingophilous scent with high emission rates of certain monoterpenes and amino-acid derived compounds. Visual cues included a special case of multisensory nectar guide and color patterns conspicuous to the visual systems of both hawkmoths and bees.

Conclusions

The intrafloral patterns of sensory stimuli suggest that hawkmoths have exerted strong historical selection on C. spinosa. Our study revealed two interesting paradoxes: (a) the flowers phenotypically biased towards the more inconsistent pollinator; and (b) floral display demands an abundance of resources that seems maladaptive in the habitats of C. spinosa. The transition to a binary pollination system accommodating large bees has not required phenotypic changes, owing to specific eco-physiological adaptations, unrelated to pollination, which make this plant an unusual case in pollination ecology.     

Are pollination syndromes useful predictors of floral visitors in Tasmania?

Abstract Diurnal visitors to the flowers of many native plant species were identified in a wide range of Tasmanian sclerophyllous vegetation between September 1996 and April 1997. These foraging profiles were analysed to determine whether they were characteristic of various floral morphologies in predictable ways. It was found that although visitor profiles were sometimes consistent with classic pollination syndromes, these syndromes were unreliable predictors of floral visitors. Very few flowers were exclusively bird‐pollinated, and none were strictly fly‐, beetle‐, wasp‐, or butterfly‐pollinated. The majority of flowering plants were unspecialized in their morphology, and consequently hosted a diverse array of visitors. In addition, visitor profiles to congeners with similar floral morphologies, and even to conspecifics, differed between habitats. Altitude was a major factor in determining visitors, with flies being the most abundant visitors above 700 m. However, congeners in several genera of Epacridaceae, as well as the genus Correa, which differed in floral morphology also differed in visitor profiles. Tubular flowers were associated with birds, while flowers with more accessible nectar were visited by insects. The only taxa exhibiting a bee‐pollination syndrome that were largely visited by bees were the Fabaceae and Goodenia ovata Sm. Several species with purple or pink flowers were also predominantly visited by bees, but did not strictly conform to the melittophilous syndrome. In contrast, other flowers exhibiting an ostensibly mellitophilous syndrome hosted very few bees. Of these, species that occurred at high altitude were mainly visited by flies, while others received very few potential pollen vectors.     

Interaction between oil-collecting bees and seven species of Plantaginaceae

Oil-bee/oil-flower mutualism evolved through multiple gains and losses of the ability to produce floral oil in plants and to collect it in bees. Around 2000 plant species are known to produce floral oils that are collected by roughly 450 bee species, which use them for the construction of nests and for the larval food. The Plantaginaceae contain several Neotropical species that produce floral oils, the main reward offered by these plants. In the genera Angelonia, Basistemon, Monopera and Monttea, mainly associated with Centris bees, the floral oil is produced in trichomes that are located in the inner corolla. The pollinators of a few species in this neotropical clade of Plantaginaceae are known, and the role of flower morphology as well as the requirements from pollinators and the role of other groups of bees in the pollination of these flowers remains unclear. In this paper we provide a list of the flower visitors of seven Plantaginaceae species (six Angelonia species and Basistemon silvaticus) analyzing their behavior to highlight the legitimate pollinators and illustrating little known aspects of flower morphology and oil-collecting apparatuses of the bees. Two general morphological patterns were observed in the Angelonia flowers: deep corolla tube with short lobes, and short corolla tube with long lobes. Corolla tubes of different length result in pollen adherence to different parts of the insect body. The six Angelonia species and B. silvaticus flowers were visited by 25 oil-collecting bee species (10 Centris, 11 Tapinotaspidini and 4 Tetrapedia species), the majority acting as legitimate visitors. The flowers were also visited by illegitimate bee pollinators, which collected pollen but do not transfer it to the female organ. Specialized collectors of Plantaginaceae floral oils present modifications on the first pair of legs, mainly in the basitarsi but also extended to the tarsomeres. The new records of Tapinotaspidini and Centridini species acting as specialized pollinators of Plantaginaceae suggest that there is a geographic variation in the pollinators of the same plant species, and that the evolutionary scenario of the historical relationships between oil-collecting bees and floral oil producing plants is more complex than previously considered.     

Two alien invasive acacias in Italy: Differences and similarities in their flowering and insect visitors

In Italy, alien acacias have been introduced for ornamental and reforestation purposes, and some species became invasive occupying patches of the Mediterranean landscape. On the Island of Elba (Central Italy), Acacia dealbata and A. pycnantha form dense stands at short distance, showing an impressive massive flowering at the end of the winter/early spring. Our aim was to investigate the behaviour of the two species in relation to the flowering features, from phenology to floral characteristics, and their replay to the observed flower visitors. Differences between the two species emerged on all the parameters considered. A. pycnantha peak of flowering occurred later than A. dealbata and showed larger flower heads (FHs), more flowers/head, stamens/flower and polyads. On A. dealbata, we recorded longer racemes and more FHs/raceme, determining a more flower-dense crown. Even if contacts with flower visitors were generally low on both species, A. dealbata showed a more heterogeneous visitor assemblage. Both acacias species interacted with local generalist pollinators, as bumblebees and honey bees. Flower handling and resource collection strategy by the honey bee indicate a long-term relationship between the bee and the acacias, with bees investing longer time on the larger A. pycnantha flower heads.     

Solitary and social bees as pollinators of Wahlenbergia (Campanulaceae): single-visit effectiveness, overnight sheltering and responses to flower colour

Solitary bees often form specialised mutualisms with particular plant species, while honeybees are considered to be relatively opportunistic foragers. Thus, it may be expected that solitary bees are more effective pollinators than honeybees when foraging on the same floral resource. To test this, we studied two Wahlenbergia species (Campanulaceae) in South Africa that are visited by both social honeybees and solitary bees, and which are shown here to be genetically self-incompatible and thus reliant on pollinator visits for seed production. Contrary to expectation, the solitary bee Lipotriches sp. (Halictidae) and social bee Apis mellifera (Apidae), which were the two most frequent visitors to flowers of the study species, were equally effective pollinators in terms of the consequences of single visits for fruit and seed set. Both bee species preferentially visited female phase flowers, which contain more nectar than male phase flowers. Male solitary bees of several genera frequently shelter overnight in flowers of both Wahlenbergia species, but temporal exclusion experiments showed that this behaviour makes little contribution to either seed production or pollen dispersal (estimated using a dye particle analogue). Manipulation of flower colour using a sunscreen that removed UV reflectance strongly reduced visits by both bee groups, while neither group responded to Wahlenbergia floral odour cues in choice tests. This study indicates that while flowers of Wahlenbergia cuspidata and W. krebsii are pollinated exclusively by bees, they are not under strong selection to specialise for pollination by any particular group of bees.     

Exploitation of floral resources on Sparattosperma leucanthum (Bignoniaceae): foraging activity of the pollinators and the nectar and pollen thieves

The interaction between floral visitors and plants in natural ecosystems has become a major subject in studies on biodiversity conservation. However, there is a very complex interaction between the community of floral visitors and the plant species foraged by them in the natural environment and little is known about the foraging activity of these visitors. Thus, the objective of this study was to determine the composition of the community of floral visitors to Sparattosperma leucanthum in an area under natural regeneration, focusing on the assessment of the exploitation of floral resources by the predominant visitors and effective pollinators. To accomplish this objective, we recorded the activity and behavior of the floral visitors throughout the day in a 2- to 3-m² area of flowering branches of S. leucanthum. The model established in the current study comprises the occurrence of a high number of species with few individuals, whereas few species are represented by many individuals. The five predominant species were Apis mellifera, Trigona spinipes, Bombus sp1, Hylocharis chrysura, and Halictidae type 1. Of these species, only Bombus sp1 seemed to effectively pollinate the flowers.     

REPRODUCTIVE BIOLOGY OF APIACEAE. I. FLORAL VISITORS TO THASPIUM AND ZIZIA AND THEIR IMPORTANCE IN POLLINATION

Based on the ready availability of nectar and pollen, and on the large numbers and great diversity of insect visitors, species in the Apiaceae have been labeled promiscuous. The distinction between floral visitors and effective pollinators, however, is extremely important and is rarely discerned. Floral visitation was documented for plants of nine populations in a comparative study of three species of the closely related apioid genera, Thaspium Nutt. and Zizia Koch. A pollinator importance index was calculated for each floral visitor using visitor abundance, pollen load composition and foraging behavior as its basic components. Results showed that, despite a high diversity of insect visitors, generally 1–4 species accounted for a minimum of about 74% of the pollinations in all populations. This specialization in pollination appears in part to be the result of an oligolectic relationship between Andrena ziziae (Hymenoptera; Andrenidae) and plants of the taxa studied, but solitary bees of Andrenidae, Colletidae and Halictidae in general were efficient and important pollinators. This study emphasizes that visitation records, when considered alone, effectively disguise specialization in the pollination system.     

Native birds and insects,and introduced honey bees visiting Echium wildpretii (Boraginaceae) in the Canary Islands

In this paper, we report observations of flower visitors of the endemic Echium wildpretii in Tenerife, Canary Islands. This plant inhabits the high altitudinal sub-alpine zone, which is characterized by a harsh climate, low species diversity and a short growing season. Echium wildpretii is a monocarpic perennial, producing a 2–3 m column-shaped, red-flowered, nectar-rich inflorescence. Although these floral traits have previously been suggested as being typical of ornithophilous flowers, this is the first study reporting observations of native birds (Phylloscopus collybita and Serinus canarius) in addition to insects visiting the flowers for nectar. The purposes of this study were firstly to investigate levels of visitation by native birds, native insects, and introduced honey bees. Secondly, we studied the influence of floral display (plant height and number of flowers), nearest neighbours (distance and size) and local vegetation structure on visitation rate. Finally, we discuss the evolution of ornithophily in an otherwise entomophilous plant lineage. We found that the level of bird visitation was relatively high early in the flowering season, but decreased in mid/late season, while the opposite pattern was found for introduced honey bees. For native insects, the frequency of visits was similar in early and late season. Bird visits were correlated with floral display. In the early season, visitation rates of honey bees and the two most common native bee species were correlated with size of the plant or its nearest neighbours, consistent with preference patterns for larger resource patches. Since only insects visit the flowers of other species in the Echium clade, E. wildpretii appears to have evolved from a truly insect-pollinated lineage.     

Floral visitors of Chamaecrista debilis (Vogel) Irwin & Barneby (Fabaceae - Caesalpinioideae) at cerrado of Estação Ecológica de Jataí, São Paulo State, Brazil

Although Chamaecrista Moench genus is a very important source of pollen to bees in the Cerrado, this relationship is almost unknown. Within flower visitors of Ch. debilis, we found hymenopterans (Apidae) as the most abundants, but Coleoptera (Buprestidae) and Lepidoptera (Noctuidae) were also collected. Bees of great size are pointed out as effective pollinators of Cassiinae. Only five out of seventeen species of floral visitors of Ch. debilis sampled in cerrado, are indicated as possible pollinators, from which four species were not previously indicated in other papers.     

Recognition of scent marks in solitary bees to avoid previously visited flowers

Knowing how floral visitors forage efficiently among flowers is important to understanding plant-pollinator interactions. When bees search for rewarding flowers, they use several visual cues to detect the available floral resources. In addition to these cues, bees can recognize scent marks, which are olfactory cues left on flowers foraged by previous visitors. This behavior is well known in social bees, such as honeybees and bumblebees. Although solitary bees do not need to give information about which flowers were foraged to conspecifics, several pieces of evidence have indicated the use of scent marks. However, it is unknown whether the behavior is widely used in many different bee species. We investigated whether four different solitary bees, Colletes patellatus (Colletidae), Andrena prostomias (Andrenidae), Osmia orientalis (Megachilidae), and Tetralonia mitsukurii (Apidae), can recognize flowers that have been foraged previously by visitors within 3 min. All four bees showed rejection responses to flowers foraged by conspecifics. However, our results showed that responses to foraged flowers varied among bee species. The tendency of A. prostomias and T. mitsukurii to reject the foraged flowers was pronounced, while in C. patellatus and O. orientalis it was weak. In both A. prostomias and T. mitsukurii, the rejection rate of flowers foraged by conspecifics decreased as the time lag after the last visit increased. Both bees visited the flowers from which pollen or nectar had been artificially removed. We suggest that A. prostomias and T. mitsukurii would recognize scent marks left by previous visitors, while the other two bees would not recognize them so strongly. It is likely that the decision to use scent marks is dependent either on the richness of resources or on the complexity of floral structure.