Triterpenoid saponins from Echinopsis macrogona (Cactaceae)

Triterpene saponins, pachanosides C1, E1, F1 and G1 (1-4), and bridgesides A1, C1, C2, D1, D2, E1 and E2 (5-11) were isolated from Echinopsis macrogona. Compounds 1-4 were saponins with pachanane type triterpene saponins, while the others (5-11) were oleanane type triterpene saponins. While the aglycones of 2-4 and 8-11 were hitherto unknown, the structure of pachanol C was revised in this paper. Their structures were elucidated on the basis of chemical and physicochemical evidence.     

Physiology and genetics of self-incompatibility in Echinopsis chamaecereus (Cactaceae)

The self-incompatibility (SI) system of a geophytic cactus (Echinopsis chamaecereus Friedrich & G. Rowley) was examined in a series of experiments. Pollination tests indicated that E. chamaecereus is an obligate outbreeding species with a functional SI system. Incompatible matings were characterized by stylar inhibition of pollen tube growth and lack of fruit set. Two S₁ seedlings were recovered when plants of one clone were exposed to 42°C for 16 h and flowers were selfed immediately after incubation. The two S₁ seedlings and the parental (S₀) clone were crossed in a full diallel. Results were consistent with a one-locus, gametophytic SI system with two different S alleles. Disturbed segregation at isozyme locus Lap-1 was attributed to close linkage with the S locus (recombination frequency = 11±8%). This is the second report of close linkage between Lap-1 and S in the Cactaceae. Recevied: 1 February 2001 / Revision accepted: 13 March 2001     

Reproductive biology of the melon cactus, Melocactus curvispinus (Cactaceae)

We examined the reproductive biology of Melocactus curvispinus Pfeiffer (Cereeae, Cactoideae) in xeric areas of northern Venezuela. Floral traits correspond to a classic hummingbird-pollination syndrome; however, pollination is shared between hummingbirds, Leucippus fallax (Bourcier 1843), and anthophorid bees, Ceratina sp. Reproduction occurs during most of the year. Anthesis and nectar secretion occur between noon and sunset. Average daily nectar production per flower was 163.1 l, nectar sugar concentration between 29.1 and 30.2% (w/w). Hummingbirds promoted inter-plant pollen movement and were relatively more reliable floral visitors than anthophorid bees, but these insects had a higher frequency of floral visits (28.75/day) than hummingbirds (4.96/day). M. curvispinus is self-compatible and autogamous. By combining extended reproductive activity, frequent animal-mediated pollination, and selfing capabilities, this cactus possesses a mating strategy that guarantees reproduction under variable environmental conditions. We argue that based on its reproductive biology, M. curvispinus should be considered an example of evolutionary transition towards selfing within tribe Cereeae.     

Beyond the pollination syndrome: nectar ecology and the role of diurnal and nocturnal pollinators in the reproductive success of Inga sessilis (Fabaceae)

Inga species present brush‐type flower morphology allowing them to be visited by distinct groups of pollinators. Nectar features in relation to the main pollinators have seldom been studied in this genus. To test the hypothesis of floral adaptation to both diurnal and nocturnal pollinators, we studied the pollination ecology of Inga sessilis, with emphasis on the nectar secretion patterns, effects of sequential removals on nectar production, sugar composition and the role of diurnal and nocturnal pollinators in its reproductive success. Inga sessilis is self‐incompatible and pollinated by hummingbirds, hawkmoths and bats. Fruit set under natural conditions is very low despite the fact that most stigmas receive polyads with sufficient pollen to fertilise all ovules in a flower. Nectar secretion starts in the bud stage and flowers continually secreting nectar for a period of 8 h. Flowers actively reabsorbed the nectar a few hours before senescence. Sugar production increased after nectar removal, especially when flowers were drained during the night. Nectar sugar composition changed over flower life span, from sucrose‐dominant (just after flower opening, when hummingbirds were the main visitors) to hexose‐rich (throughout the night, when bats and hawkmoths were the main visitors). Diurnal pollinators contributed less than nocturnal ones to fruit production, but the former were more constant and reliable visitors through time. Our results indicate I. sessilis has floral adaptations, beyond the morphology, that encompass both diurnal and nocturnal pollinator requirements, suggesting a complementary and mixed pollination system.     

Floral biology and pollinators of Trichocereus pasacana (Cactaceae) in Parque Nacional Los Cardones, Argentina]

Many columnar cacti are bat pollinated. It has been suggested that this kind of pollination would be more important in tropical than in temperate regions where flowers are open only one night. Thrichocereus pasacana produces big and resistant white flowers. We analyzed flower characteristics, floral cycle, stigmatic receptivity, nectar production, pollen presence and floral visitors in a T. pasacana population at National Park Los Cardones (Salta, Argentina) in November 1997. Flower features were constant between individuals of the population. Flowers start opening at evening and anthesis time is from 18 to 40 hs. The estigma was receptive throughout the floral cycle. Anther dehiscence occurs with flower opening. Nectar production was highest between 18 to 24 hs. Although T. pasacana are open during the night, floral visitors are diurnal. The most frequent was Xylocopa sp. In the study area, nectarivorous bats were not detected. The morphological features of T. pasacana flowers were similar but bigger compared to other columnar cacti. Anthesis time was also longer while nectar production was lower. T. pasacana pollination at National Park Los Cardones is done by bees.     

Stamen movements in flowers ofOpuntia (Cactaceae) favour oligolectic pollinators

Opuntia brunneogemmia andO. viridirubra occur sympatrically in the Serra do Sudeste, Rio Grande do Sul, Brazil. Their flowers have 450–600 thigmonastic stamens and provide large amounts of pollen and nectar for bees. Bees of 41 species were registered at the flowers ofO. brunneogemmia and 30 at the flowers ofO. viridirubra. Females of three oligolectic species are the only effective pollinators:Ptilothrix fructifera (Anthophoridae),Lithurgus rufiventris (Megachilidae), andCephalocolletes rugata (Colletidae). During their visits inOpuntia-flowers, bees touch the filaments and stimulate the movement of the stamens to the centre of the flower. At the end of this movement, the anthers are densely packed around the style. As a consequence the pollen is presented in an easily accessible upper layer of anthers and various, nearly inaccessible lower layers. The lower layers contain about 80% of the pollen reward. Only females of the three oligolectic pollinators exploit the pollen from the lower layers and reach the nectar furrow. Therefore, through their stamen movements,Opuntia flowers hide most of their pollen from flower visitors but favour effectively pollinating, oligolectic bees.     

Reproductive biology of Echinodorus longipetalus (Alismataceae): Sexual morphs,breeding system and pollinators

The floral biology, pollinators and breeding system of Echinodorus longipetalus Micheli were studied in a marshy area of the district of Taquaritinga (State of São Paulo), southeastern Brazil. E. longipetalus is gynodioecious and as far as is known, this is the first record of unisexual flowers, besides perfect flowers, in Echinodorus. Proportion of female individuals in the studied population is 50% and produces 31% more flowers than hermaphrodites. Perfect and pistillate flowers of E. longipetalus are similar in appearance and are pollinated by several species of Hymenoptera (mainly by Xylocopa (Neoxylocopa) suspecta Moure & Camargo). Perfect flowers offer pollen as a reward. Pistillate flowers attract floral visitors by deceit with their staminodes that resemble the stamens of the perfect flowers. Visits to pistillate flowers are quick (1–2 s), while visits to perfect flowers last up to 120 s. The perfect flowers are self-compatible and produce fruits through spontaneous self-pollination (control flowers), whereas the pistillate ones only set fruits through cross-pollinations. Perfect and pistillate flowers set more fruits under natural conditions than in manual treatments, respectively. Although the pistillate and perfect flowers bear a strong similarity, the selective pollinator behavior seems to be responsible for the increase of fruit set in perfect flowers.     

Pollination biology of Lilium japonicum var. abeanum and var. japonicum: evidence of adaptation to the different availability of diurnal and nocturnal pollinators

Floral traits such as color and size are highly diversified in lilies, but their adaptive significance remains uncertain. In the present study, we compared pollination processes between Lilium japonicum var. abeanum and var. japonicum to clarify how the two varieties are adapted to different pollinators. Lilium japonicum var. japonicum is known to be pollinated by moths, and we hypothesized from its flower traits that var. abeanum is pollinated by diurnal insects. Using waterproof digital cameras set to a recording interval of 10–30 s, we recorded flower visitors for 7–9 flowers of var. japonicum and 4–6 flowers of var. abeanum over 24‐h periods. We also recorded the number of versatile (T‐shaped) and rigid (I‐shaped) anthers per flower. For var. abeanum, we observed flowers at 05.00 and 17.00 hours to determine flower opening time and measured the intensity of floral scent using a metal‐oxide semi‐conductor odor sensor. Both diurnal and nocturnal insects visited flowers of the two varieties, but visitation of diurnal insects was more frequent in var. abeanum. Anthers of var. abeanum are usually rigid, as in many bee‐pollinated flowers, whereas those of var. japonicum are mostly versatile. Although flowers of var. japonicum are known to open in the evening, 32% of the flowers of var. abeanum started to flower during the day. Lilium japonicum var. abeanum emitted scent at night, with a maximum intensity at 20.00 hours, just as in var. japonicum. These findings suggest that the floral traits of the two varieties are adapted to the different relative availabilities of nocturnal versus diurnal pollinators. The flower of var. abeanum is more adapted to diurnal pollinators, but retains adaptations to nocturnal pollinators, particularly with regard to its scent.     

Reproductive biology and mating system estimates of two Andean melocacti, Melocactus schatzlii and M. andinus (Cactaceae)

BACKGROUND AND AIMS: The genus Melocactus comprises 36 species of globose cacti with the most derived traits in the Cereeae tribe. It is the proper study system to examine what are the most derived reproductive strategies within that tribe. This study aims to characterize the reproductive biology and to estimate the mating system parameters of two Andean melocacti, Melocactus schatzlii and M. andinus. METHODS: The reproductive attributes of the two species were described, including floral morphology, anthesis patterns, floral rewards, floral visitors and visitation patterns. Levels of self-compatibility and autonomous self-pollination were estimated by hand-pollination experiments. Mating system estimates were obtained by conducting progeny array analyses using isozymes. KEY RESULTS: The flowers of the two species present the typical hummingbird-pollination syndrome. Despite their morphological resemblance, the two species differ in flower size, pollen and ovule production and anthesis pattern. Their main pollinator agents are hummingbirds, four species in M. schatzlii and one species in M. andinus. Both cacti are self-compatible and capable of self-pollination without the aid of pollen vectors. Population-level outcrossing rate was higher for M. schatzlii (t(m)=0.9) than for M. andinus (t(m)=0.4). At the family level, outcrossing rates for most mothers of M. schatzlii were higher (t(m)>0.8) than for M. andinus (t(m)<0.5). CONCLUSIONS: Although the two cacti are capable of selfing, M. schatzlii is a predominantly outcrossing species, while M. andinus behaves as a mixed-mating cactus. Hummingbirds are the only pollinators responsible for outcrossing and gene flow events in these species. In their absence, both melocacti set seeds by selfing. Based on its low population size, restricted distribution in Venezuela, low rates of floral visits, and high levels of inbreeding, M. andinus is considered to be an endangered species deserving further study to define its conservation status.     

Diurnal and nocturnal pollination of Marginatocereus marginatus (Pachycereeae: Cactaceae) in Central Mexico

BACKGROUND AND AIMS: Chiropterophillous and ornithophillous characteristics can form part of a single reproductive strategy in plants that have flowers with diurnal and nocturnal anthesis. This broader pollination strategy can ensure seed set when pollinators are scarce or unpredictable. This appears to be true of hummingbirds, which presumably pollinate Marginatocereus marginatus, a columnar cactus with red nocturnal and diurnal flowers growing as part of dense bat-pollinated columnar cacti forests in arid regions of central Mexico. The aim of this study was to study the floral biology of M. marginatus, and evaluate the effectiveness of nocturnal vs. diurnal pollinators and the contribution of each pollinator group to overall plant fitness. METHODS: Individual flower buds were marked and followed to evaluate flower phenology and anthesis time. Flowers and nectar production were measured. An exclusion experiment was conducted to measure the relative contribution of nocturnal and diurnal pollinators to seed set. KEY RESULTS: Marginatocereus marginatus has red hermaphroditic flowers with nocturnal and diurnal anthesis. The plant cannot produce seeds by selfing and was pollinated during the day by hummingbirds and during the night by bats, demonstrating that both pollinator groups were important for plant reproduction. Strong pollen limitation was found in the absence of one of the pollinator guilds. CONCLUSIONS: Marginatocereus marginatus has an open pollination system in which both diurnal and nocturnal pollinators are needed to set seeds. This represents a fail-safe pollination system that can ensure both pollination, in a situation of low abundance of one of the pollinator groups (hummingbirds), and high competition for nocturnal pollinators with other columnar cacti that bloom synchronously with M. marginatus in the Tehuacan Valley, Mexico.     

Relative effects of nocturnal vs diurnal pollinators and distance on gene flow in small Silene alba populations

Silene alba exists in natural metapopulations throughout its range and is visited by a suite of both diurnal and nocturnal pollinators. Pollen-mediated gene flow may help reduce genetic isolation of subpopulations. Here, we compared the relative effects of nocturnal vs diurnal pollinators on pollen-mediated gene flow in subpopulations separated by two distance treatments. We established populations consisting of genetically marked individuals in an old field in Tennessee (USA). Electrophoretic examination of seedlings produced by plants exposed to nocturnal, diurnal and control pollinator treatments and separated by either 20 or 80 m allowed us to directly measure pollen-mediated gene flow. Gene flow was more common between populations separated by only 20 m. Nocturnal pollinators were responsible for most gene flow between populations, regardless of distance. Diurnal pollinators played only a small role in pollen-mediated gene flow. The results suggest that nocturnal pollinators are better than diurnal pollinators at moving pollen between small S. alba subpopulations. However, their effectiveness declines as the distance between subpopulations increases, making them relatively ineffective at moving genes between isolated subpopulations.     

Reproductive biology of the genusDamasonium (Alismataceae)

The reproductive biology of the genusDamasonium was studied in cultivation and in nature. A correlation was shown between flowering time and latitude. All species are self-compatible but the N. American member of the genus (D. californicum) is strongly protandrous and probably experiences high levels of outcrossing in nature. In the European taxa the stamens and the stigmas mature simultaneously but the levels of inbreeding appear to be greater in the southern species (D. bourgeai, D. polyspermum) than in the northern (tetraploid) speciesD. alisma. The Australian memberD. minus resembles the southern European species in being strongly autogamous and bud-pollination was shown to occur. Floral morphology, stigma exertion, pollen/ovule (P/O) ratio and seed weight each show a strong correlation with the degree of autogamy. Vegetative reproduction occurs only inD. californicum andD. alisma through the production of corm-buds. This study suggests that autogamy inDamasonium is accompanied by a shift in resource allocation from male (pollen) toward female (seed) function. However, further studies are required to assess the quality of seeds and their survivorship in nature.     

Floral biology and pollinators of three co-occurring Cistus species (Gistaceae)

BOSCH, J., 1992. Floral biology and pollinators of three co-occurring Cistus species (Cistaceae). The pollination of three species of Cistus, Cistus albidus L., Cistus salvifolius L. and Cistus monspeliensis L. is studied. Insect visitors were censused and some were captured to analyse pollen distribution on their bodies. Floral phenology, breeding systems, some aspects of floral biology (pollen and nectar production, pollen-ovule ratios) and floral morphology were also studied for each species. Flowers of C. albidus produce more pollen and nectar than the other two species studied, and attract more insects, especially those with higher nutritional requirements. All three species are highly self-incompatible, but C. salvifolius and C. monspeliensis show some features that hinder xenogamy, and they have lower pollen-ovule ratios than C. albidus. As a result, this latter species apparently receives higher levels of cross-pollination, but at the cost of a higher investment to attract and reward reliable pollinators.     

The reproductive biology and effective pollinators of the endangered beardtonguePenstemon penlandii (Scrophulariaceae)

Penland's beardtongue, a rare endemic plant of the Colorado Plateau, displays a mixed breeding system. Plants are partially self-compatible but set more fruits when cross-pollinated than when self-pollinated. Fruit production is significantly increased by pollinators. However, in two years of study there was no indication that fruit set was being limited by inadequate pollinator visitation. Pollinator effectiveness was judged by correlating bee behavior at the flowers with analysis of the pollen carried on bee bodies. The most important pollinators were native megachilid bees, particularly in the genusOsmia. The bees that pollinate Penland's beardtongue are essential to its reproduction and must be preserved along with this rare plant.     

Reproductive biology of the Andean-disjunct genusRetanilla (Rhamnaceae)

Various aspects of the reproductive biology of the Andean-disjunct genusRetanilla (DC.) Brongn. were studied in populations ofR. trinervia andR. ephedra located in central Chile and ofR. patagonica in southern Argentina. Flowering extends from -austral- winter (early spring inR. patagonica) to early summer. The small, white, entomophilous flowers are incompletely protandrous and last four to five days. A weak pleasant odor along with nectar and pollen attract thirty-seven diurnal insect species, ten of which are probable pollinators: these include the honey bee, eight species of solitary bees pertaining toAnthophoridae (1 sp.),Colletidae (3 spp.) andHalictidae (4 spp.), and a nemestrinid fly. Although flower morphology and individual flower phenology do not fully prevent self-pollination (and geitonogamy can easily take place), the level of autogamy is low. Therefore, some self-incompatibility mechanism seems operative inRetanilla. Fruit set of open-pollinated flowers is extremely low, with a maximal value of c. 3% inR. patagonica. In this species, ripe fruits contain on average 1.37 viable seeds. Predispersal (maternal) reproductive success (percent ovules becoming viable seeds) is 2.3%. For medium to full-sized individuals this corresponds to c. 4600 potential offspring per year. As presently known,Retanilla is a reproductively uniform group, in which the Andean disjunction seems to have exerted no particular impact, which is consistent with the view that pollination generalization exerts some stabilizing influence on floral morphology and other reproductive traits.Retanilla expresses a basic rhamnacean set of traits (including protandry and self-incompatibility) showing also high pollen production and secondary pollen presentation. The latter two traits seem characteristic of theRetanilla-Trevoa clade, suggesting that a trend to increased male effort and, perhaps, dependence upon polleneating has evolved within the tribe Colletieae.     

Reproductive biology ofAsphodelus aestivus (Asphodelaceae)

InAsphodelus aestivus the large number of showy flowers opened per inflorescence, high nectar and pollen production, and absence of automatic self-pollination, so that insect visitation is necessary for pollination, indicate that cross-fertilization is favoured. LargeHymenoptera are the main pollinators. Hand self-pollinations result in some degree of fruit-set, but cross-pollination also yield low fruit- and seed-set. The perennial habit and polyploid status are in agreement with the breeding system of this species, with sexual reproduction being rather inefficient and active vegetative propagation.     

Reproductive biology of Pseudococcus maritimus (Hemiptera: Pseudococcidae)

This study characterized several parameters associated with reproductive behaviors of a Californian population of the grape mealybug, Pseudococcus maritimus (Ehrhorn). Experiments with caged adults confirmed that females from this population could not reproduce parthenogenetically. Both sexes were capable of mating multiple times on the same day and on sequential days. Median times between copulations were short (< 10 min) on the first day that males were presented with females, but tended to increase with sequential copulation events. Unmated females lived for up to 19 wk, whereas mating and oviposition resulted in reduced longevity. We hypothesized that constant exposure to pheromone might stimulate males to leave their cocoons sooner and increase their activity levels compared with males that were not exposed to pheromone, resulting in shortened life-spans. However, whereas exposure to pheromone did stimulate males to emerge slightly sooner than control males, pheromone exposure had no effect on the longevity of males. The implications of the results of these experiments for pheromone-based methods of detection and control of mealybugs are discussed.     

Geographic variation in floral scent of Echinopsis ancistrophora (Cactaceae); evidence for constraints on hawkmoth attraction

Variation in floral phenotype (color, depth, nectar) suggests incipient specialization for bee or hawkmoth pollination across the geographic distribution of Echinopsis ancistrophora , with flower depth ranging from 4.5 to 24 cm. We used chemical and behavioral analyses to test whether fragrance has evolved in concert with morphology in these Andean cacti. Floral scent (145 total compounds) was collected using dynamic headspace methods and analyzed with gas chromatography–mass spectrometry, revealing subspecies-specific odors dominated by sesquiterpenes in E. ssp. ancistrophora and arachnacantha and fatty acid derivatives or aromatics in E. ssp. cardenasiana and pojoensis . Compounds indicative of sphingophily were not consistently found in moth-pollinated plants, and total scent emissions were significantly lower in populations with nocturnal anthesis. In wind tunnel assays, Manduca sexta moths were attracted to scent of ssp. ancistrophora from both bee and hawkmoth-pollinated populations, but not to scent of ssp . cardenasiana . However, hawkmoths were most attracted to the methyl benzoate-dominated scent of a distant relative, Echinopsis mirabilis . Thus, hawkmoth-pollinated descendants of the E. ancistrophora lineage may be phylogenetically constrained to emit weak, sesquiterpene-dominated fragrances that are not optimally attractive to hawkmoths, or floral scent may be under stronger selection by destructive flower visitors.