Gender expression and inflorescence structure of Pappea capensis Eckl. and Zeyh. (Sapindaceae)

Gender and the structure of the inflorescence and flowers of Pappea capensis (Sapindaceae) are investigated in a locality around Pretoria (22-27°S and 25-32°E). The trees flower over a long period (December to April) and are basically monoecious, starting with male flowers followed by female flowers towards the end of the flowering period, although some trees may be predominantly male and some predominantly female. The inflorescence is a reduced thyrse with small flowers. Male flowers have five ephemeral petals, eight stamens and a rudimental pistil. Female flowers comprise a 3-lobed ovary, a single style and stigma and eight staminodes.     

Spatial and temporal dispersion patterns of pollinators and their relationship to the flowering strategy of Yucca whipplei (Agavaceae)

Summary A field investigation of the mutualistic interaction between a monocarpic perennial plant, Yucca whipplei, and its host-specific pollinator and seed predator, Tegeticula maculata (Lepidoptera: Prodoxidae), was conducted to determine how the resource utilization pattern and population dynamics of the pollinator have influenced the evolution of the flowering and fruiting pattern of the plant. Although the temporal pattern of emergence of pollinators results in a relatively close tracking of flower abundance within a season, the ratio of pollinators to open flowers does vary significantly within a season, as well as between seasons. At any point in time during the flowering season, the population of adult yucca moths is distributed evenly among the available flowers, so that the number of pollinators on an inflorescence is directly proportional to the number of open flowers available. The relative isolation of individual flowering plants appears to have little effect on the distribution of pollinators among inflorescences. The number of fruits initiated on a plant is directly proportional to the number of flowers produced, and is also partially determined by the time of flowering. Yucca whipplei always produces many more flowers than fruits. Most flowers are not fertilized, and the plants also generally abort and abscise immature fruits after flowering. Fruit production of at least some plants, however, appeared limited by pollination. It is also expected that in some years the relative abundance of pollinators will be low enough that most plants will be pollinator-limited. It is suggested that the pattern of flowering and fruiting of this species has evolved in response to the unpredictability of pollinator availability, both within and between seasons. Resource uncertainty and selection acting on the male component of fitness may also be involved.     

REPRODUCTIVE SUCCESS AND INFLORESCENCE SIZE OF CALOPOGON TUBEROSUS (ORCHIDACEAE)

Reproductive success of Calopogon tuberosus, which produces no nectar, was investigated in relation to inflorescence size and dispersion pattern. Mean inflorescence size was 2.56 (range 1–10). A bagging experiment showed that insects are required for pollen transfer and that fruits are produced from self-, geitonogamous, and cross-pollinations; fruit set was not 100%. Fruit set of nonmanipulated plants was limited by the number of pollinator visits. Reproductive success increased with increasing inflorescence size, although not above theoretical predictions. However, the probability of producing no fruit or contributing no pollinia decreased with increasing inflorescence size since sequential flowering increased the probability of a pollinator visit to the inflorescence over the blooming period. Large inflorescences did not provide a greater pollinator attraction than small ones, because inflorescences only presented a few open flowers at a time. In addition, flowers on plants growing in clumps of 2–8 plants had a higher probability of setting fruit, apparently because of increased pollinator attraction. Although there are obvious selective advantages for large inflorescences, the sequential flowering habit, and low resource availability may reduce the advantages of large inflorescence size at our study site.     

Fine-scale temporal dynamics of flower visitors sheds light on insect-assemblage overlap between sexes in a dioecious Ecuadorian palm

Dioecious plants generally display sexual dimorphism in male and female floral traits, potentially attracting slightly different pollinator communities. The sharing of common floral visitors between male and female flowers and their timing of visits to both sexes is of critical importance to ensure plant's reproductive success. Palm inflorescences are visited by abundant and diverse insect communities, yet the temporal patterns of insect visits on both sexes remain poorly known. We characterized the composition of a community of flower-visiting arthropods associated with the dioecious ivory palm (Phytelephas aequatorialis, Spruce) in a pre-montane forest of Ecuador. We monitored the temporal variations in insect visits along the flowering of 12 inflorescences (eight female and four male) using interception traps recovered every 4 h. We report 59 morphospecies in the arthropod community, dominated by three beetle families: Staphylinidae, Nitidulidae, and Curculionidae. Male inflorescences were more abundantly visited than female, but visitors of the later were taxonomically more diverse. Among the 16 pollinator candidates identified, nine visited both inflorescence sexes synchronously at dusk /night whereas the others did so asynchronously during the day. Our study provides new insights into the pollination mechanism of P. aequatorialis. We found evidence of differential pollinator attraction between floral sexes, which may be explained by the sexual dimorphism of both flowers. Synchronicity in dusk/night visits of both inflorescence sexes suggests a sexual synchronization of the signal used to attract pollinators.     

Pollination biology and sexual differentiation ofOsyris alba (Santalaceae) in the Mediterranean region

Osyris alba L. is a widespread dioecious hemiparasitic shrub of S Europe, N Africa, and SW Asia. Male inflorescences are multiflowered whereas each female inflorescence is reduced to a single flower with persistent enlarged bracts. Pollination is a prerequisite for fruit and seed development and wind is unlikely to be an effective means of pollen spread. In southern Italy pollen is transported by small unspecialized flies and beetles. Both male and female flowers produce an indistinguishable sweet odour. Male flowers are produced in large numbers and over a larger period than the females and provide pollen, nectar, and staminal hairs as rewards for pollinators. The presence and function of staminal hairs with tip cells inOsyris alba has been reported for the first time. Female flowers are rewardless, producing neither mature pollen, nectar nor staminal hairs, but possess three modified yellow indehiscent anthers containing no viable pollen which may provide a strong visual feeding stimulus for pollinators. It is suggested that pollinators are attracted by deceit to female flowers by mimicry of the males and the floral mimicry is, therefore, intraspecific and intersexual. The floral characteristics and flowering phenology of male and female plants are consistent with this kind of mimicry. The female flower possesses a tricarpellary ovary with three ovules of which only one develops. The single seed, containing a small embryo and a large, rich endosperm, is borne in a red fleshy bird-dispersed fruit. The reduction in seed number per flower to one highly nutrient-invested seed, together with a reduction of the multiflowered inflorescence to a solitary flower and the sequential production of ripe fruits over an extended fruiting season, suggest that the female function is markedly resource-limited. It is suggested that, although all the reproductive characteristics present inOsyris alba, as well as hemiparasitism, had probably evolved before the end of the tropical Tertiary, they are of adaptive advantage in the nutrient and water-limited environment of the Mediterranean maquis.     

Flowering phenology,floral display and reproductive success in dioecious,Aralia nudicaulis L. (Araliaceae)

Summary Aralia nudicaulis L. is a dioecious, perennial, herbaceous plant that is commonly found in the understory vegetation throughout the boreal forest of North America. Female remets have fewer flowers per inflorescence, initiate flowering earlier, and reach peak flowering before male ramets. The consequences of the asynchrony in flowering between the sexes on pollination and seed set were examined during a two-year study. In both years there was significant variation in seed set associated with the flowering times of individual female ramets. In 1983, seed production was highest in the middle of the flowering season. In 1984, seed production was greatest in the later stages of flowering. Variation in seed set was not attributed to lack of pollination in 1983. In 1984, pollination limited seed set per flower during peak flowering. However, seed production never reached the potential five seeds per flower, suggesting that resource limitation was the most important factor affecting fecundity in both years. The asynchronous pattern of flowering is suggested to be the result of the different inflorescence sizes between the sexes.     

Patterns and implications of floral nectar secretion, chemical composition, removal effects and standing crop in Mandevilla pentlandiana (Apocynaceae)

Flower morphology, nectar features (chemical composition, production pattern, removal effects, and standing crop) and floral visitors are analysed in an Argentine population of Mandevilla pentlandiana. Nectar variability was examined during the lifetime of a single flower, over the course of the flowering season, and at different times of the day. Nectar is sucrose dominant. There were some variations in the proportions of sugar throughout both the flower lifespan and the flowering season. Flowers produced most nectar during bud-stage. Nectar secretion ceased near the end of the first day after flower opening. Nectar quantity varied as a function of flower age due to a combination of nectar secretion, cessation, and resorption periods. Overall sugar production was increased by nectar removal. Standing crop data showed that each open flower and inflorescence offers c. 2 and 11 mg of sugar respectively at any time of the flowering season. There was higher nectar availability at the beginning of the flowering season compared with the rest of the period. Flowers were visited by bumblebees, honeybees and hummingbirds. The greater the number of open flowers and the nectar variance, the more the mean reward quantity per flower available in the inflorescence. The sources of nectar variability in M. pentlandiana seem to be linked with both the female function (nectar resorption, nectar cessation) and the male one (early and comparatively large nectar availability, variation in nectar production as the flower ages, nectar secretion stimulation by nectar removals).     

Influence of inflorescence size on sexual expression and female reproductive success in a monoecious species

Sex allocation theory forecasts that larger plant size may modify the balance in fitness gain in both genders, leading to uneven optimal male and female allocation. This reasoning can be applied to flowers and inflorescences, because the increase in flower or inflorescence size can differentially benefit different gender functions, and thus favour preferential allocation to specific floral structures. We investigated how inflorescence size influenced sexual expression and female reproductive success in the monoecious Tussilago farfara, by measuring patterns of biomass, and N and P allocation. Inflorescences of T.?farfara showed broad variation in sex expression and, according to expectations, allocation to different sexual structures showed an allometric pattern. Unexpectedly, two studied populations had a contrasting pattern of sex allocation with an increase in inflorescence size. In a shaded site, larger inflorescences were female-biased and had disproportionately more allocation to attraction structures; while in an open site, larger inflorescences were male-biased. Female reproductive success was higher in larger, showier inflorescences. Surprisingly, male flowers positively influenced female reproductive success. These allometric patterns were not easily interpretable as a result of pollen limitation when na?vely assuming an unequivocal relationship between structure and function for the inflorescence structures. In this and other Asteraceae, where inflorescences are the pollination unit, both male and female flowers can play a role in pollinator attraction.     

Flower herbivory and seed predation in Silene vulgaris (Caryophyllaceae): effects of pollination and phenology

Pollination and seed predation were studied in Silene vulgaris populations during two seasons, one with much lower pollinator abundance than the other. Among the pollinators, noctuid moths of the genus Hadena also acted as seed predators. Nectar-foraging female moths oviposited in flowers, and their larvae consumed flowers and seed capsules.
Despite a lower percentage of pollinated flowers in the year of low pollinator abundance, similar numbers of flowers set fruit in both years, because fewer flower buds and flowers were eaten by Hadena larvae during the year of low pollinator visitation. The number of seed capsules preyed upon was also lower in the year with low pollinator abundance, resulting in a higher seed set. The positive correlation between the percentage of pollinated flowers and the percentage of seed capsules destroyed was also observed when comparing flowers opening in different parts of the season.
Early flowering plant individuals had the same pollination success but suffered higher seed predation than late flowering ones. Selection for maximized pollination success through synchronous flowering, is probably the main reason for the compressed flowering period in 5. vulgaris , but the high level of predation early in the season may further increase the reproductive success of synchronous flowering individuals.     

FLOWER DEVELOPMENT IN DIOECIOUS SPINACIA OLERACEA (CHENOPODIACEAE)

Spinacia oleracea (Chenopodiaceae) is a potential model system for studies of mechanisms of sex expression and environmental influences on gender in dioecious species. Development of the male and female flowers and inflorescences of spinach were studied to determine when the two sex types can be distinguished. We found that female inflorescence apices are significantly larger than those of the male. Flower primordia are similar in size prior to perianth initiation, but the male primordia develop at a faster rate. Another distinguishing feature at this early stage is the larger bract subtending the female primordium. The two flower types become readily distinguishable when the perianth initiates. Male flowers produce four sepals and four stamens in a spiral pattern in close succession. Female flowers produce two alternate perianth parts that enlarge somewhat before the gynoecium becomes visible. There are no traces of gynoecia in male flowers or of stamens in female flowers. We propose that plant sex type is determined before inflorescence development, prior to or at evocation.     

Differential pollination success in the course of individual flower development and flowering time in Gentiana pneumonanthe L. (Gentianaceae)

The effect of flowering time and anthesis stage on fecundity of Gentiana pneumonanthe L. (Gentianaceae) in a greenhouse experiment was studied. The species is self-compatible, not apomictic, and rarely self-pollinates spontaneously. On average, flowers lived for 11.4 days, starting with a male stage of 3.6 and continuing with a female stage of 7.9 days. Pollen viability was highest during the first two days of the male stage, but never exceeded 50%. Seed-set from cross-pollination was higher earlier in the flowering period (77%) than later (63Vo). For self-pollinations, no significant difference was observed. The success of hand-pollinations declined dramatically after the fourth day of the female stage. Self-pollination yielded lower seed-set on the first day of the female stage than on the following three days. Also, there was a significant difference in seed-set between self- and cross-pollinations (25% vs 55%, respectively) only on the first day. This suggests that selection against self pollen was stronger during the first stage of the female phase and weakened as the stylar tissue aged. In conclusion, two different ways to enhance outcrossing were observed: (1) within the population's flowering period, and (2) within an individual flower's lifespan. Both may have important effects on the genetic variability of G. pneumonanthe offspring under unfavourable conditions. Wese include small population size (associated with reduced pollinator service), general pollinator scarcity, or rainy summers.     

Evidence for stabilising selection acting on flowering time in Arum maculatum (Araceae): the influence of phylogeny on adaptation

The relationship between flowering time and reproductive success was investigated in the fly-pollinated, monoecious perennial herb Arum maculatum L. (Araceae). This species temporarily traps its principle pollinator, a psychodid midge. Probability of fruit set was analysed in relation to early, peak and late periods of the flowering phenology of four British populations between 1992 and 1997. In three out of five cases, plants which flowered during early and late periods were significantly less likely to set fruit. In addition, one population showed a similar relationship for percentage fruit set of individual inflorescences, and seeds from peak-flowering plants were significantly heavier. There was no variation in number of female flowers per inflorescence over the flowering season. Probability of fruit set appears to be mediated by the likelihood of trapping psychodid midges that have previously been trapped and picked up pollen, an unlikely event during early and late flowering periods when few inflorescences are open. The majority of plants in all populations produce only one inflorescence which means that timing of flowering may be crucial to reproductive success. We interpret our findings as evidence that stabilising selection may be acting on some populations and/or during some years. The ultimate cause, however, can be related to the very short (12–18 h) female phase of each inflorescence, a phylogenetically conservative trait within the Araceae. Received: 19 August 1998 / 15 February 1999     

Flowering phenology, display size, and fruit set in an understory dioecious shrub, Aucuba japonica (Cornaceae)

I investigated the effects of display size and flowering phenology on fruit set in Aucuba japonica, an understory dioecious shrub pollinated by opportunistic insects. Natural variations in display size, flowering phenology, and fruit set were monitored in 1997. A hand-pollination experiment was also conducted to check whether pollen limitation was a factor in fruit set in the field. Increases in floral display size did not affect fruit set; the proportion of flowers that set fruit was almost constant irrespective of the total number of flowers per inflorescence, the total number of inflorescences per plant, and the total number of flowers per plant. The hand-pollination experiment showed that fruit set was not pollen limited despite the low mating probability that resulted from the combination of dioecism and the species' dependence on opportunistic pollinators. This was due, in part, to the fact that female flowers did not have a predetermined period of receptivity, but instead remained receptive until they received pollen. In contrast, flowering phenology did affect fruit set. Fruit set was most abundant when male and female flowering was most abundant. This suggests there was some degree of pollen limitation during the part of the flowering season when male flowers were scarce.     

Phenology of Ficus variegata in a seasonal wet tropical forest at Cape Tribulation, Australia

Abstract. We studied the phenology of 198 mature trees of the dioecious fig Ficus variegata Blume (Moraceae) in a seasonally wet tropical rain forest at Cape Tribulation, Australia, from March 1988 to February 1993. Leaf production was highly seasonal and correlated with rainfall. Trees were annually deciduous, with a pronounced leaf drop and a pulse of new growth during the August-September drought. At the population level, figs were produced continually throughout the study but there were pronounced annual cycles in fig abundance. Figs were least abundant during the early dry period (June-September) and most abundant from the late dry season (October-November) through the wet season (December-April). The annual peak in reproduction actually reflected two staggered peaks arising from gender differences in fig phenology. In this dioecious species, female and male trees initiated their maximal fig crops at different times and flowering was to some extent synchronized within sexes. Fig production in the female (seed-producing) trees was typically confined to the wet season. Male (wasp-producing) trees were less synchronized than female trees but reached a peak level of fig production in the months prior to the onset of female fig production. Male trees were also more likely to produce figs continually. Asynchrony among male fig crops during the dry season could maintain the pollinator population under adverse conditions through within- and among-tree wasp transfers.     

A ballistic pollen dispersal system influences pollination success and fruit‐set pattern in pollinator‐excluded environments for the endangered species Synaphea stenoloba (Proteaceae)

The critically endangered Synaphea stenoloba (Proteaceae) has numerous scentless flowers clustered in dense inflorescences and deploys a ballistic pollen ejection mechanism to release pollen. We examined the hypothesis that active pollen ejection and flowering patterns within an inflorescence influence the reproductive success (i.e. fruit formation) of individual flowers within or among inflorescences of S. stenoloba in a pollinator‐excluded environment. Our results showed that: (1) no pollen grains were observed deposited on the stigma of their own flower after the pollen ejection system was manually activated, indicating self‐pollination within an individual flower is improbable in S. stenoloba; (2) fruit set in the indoor open pollination treatment and the inflorescence‐closed pollination treatment indicated that S. stenoloba is self‐compatible and pollen ejection can potentially result in inter‐floral pollination success; (3) fruit set in the inflorescence‐closed pollination treatment was significantly lower than that of indoor open pollination, indicating within‐ and between‐flower pollination events in an inflorescence are most likely limited, with pollination between inflorescences providing the highest reproductive opportunity; and (4) analysis of the spatial distribution of cumulative fruit set on inflorescences showed that pollen could reach any flower within an inflorescence and there was no functional limitation on seed set among flowers located at various positions within the inflorescence. These data suggest that the pollen ejection mechanism in S. stenoloba can enhance inter‐plant pollination in pollinator‐excluded environments and may suggest adaptation to pollinator scarcity attributable to habitat disturbance or competition for pollinators in a diverse flora. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 59–68.     

Variation in inflorescence size in a dioecious fig tree and its consequences for the plant and its pollinator fig wasp

The host-specific relationship between fig trees (Ficus) and their pollinator wasps (Agaonidae) is a classic case of obligate mutualism. Pollinators reproduce within highly specialised inflorescences (figs) of fig trees that depend on the pollinator offspring for the dispersal of their pollen. About half of all fig trees are functionally dioecious, with separate male and female plants responsible for separate sexual functions. Pollen and the fig wasps that disperse it are produced within male figs, whereas female figs produce only seeds. Figs vary greatly in size between different species, with female flower numbers varying from tens to many thousands. Within species, the number of female flowers present in each fig is potentially a major determinant of the numbers of pollinator offspring and seeds produced. We recorded variation in female flower numbers within male and female figs of the dioecious Ficus montana growing under controlled conditions, and assessed the sources and consequences of inflorescence size variation for the reproductive success of the plants and their pollinator (Kradibia tentacularis). Female flower numbers varied greatly within and between plants, as did the reproductive success of the plants, and their pollinators. The numbers of pollinator offspring in male figs and seeds in female figs were positively correlated with female flower numbers, but the numbers of male flowers and a parasitoid of the pollinator were not. The significant variation in flower number among figs produced by different individuals growing under uniform conditions indicates that there is a genetic influence on inflorescence size and that this character may be subject to selection.     

Gender variation of sequential inflorescences in a monoecious plant Sagittaria trifolia (Alismataceae)

In protogynous plants, female flowers of early blooming plants are at a reproductive disadvantage because they cannot set fruit due to the lack of available pollen. To study this phenomenon, gender expression of the monoecious herb Sagittaria trifolia was investigated over the entire flowering season in two field and two cultivated populations in Hubei and Hunan Provinces, China. In racemes of S. trifolia, flowers open sequentially from bottom to top, with female flowers opening first followed by male flowers. This creates a temporal separation of sexes in the species. Under field conditions small plants are often male, with production of both male and female flowers increasing with plant size. Femaleness increased among sequential inflorescences since female flower production increased whereas male flower production did not. Seed production was greater in large inflorescences because they contain more female flowers, and the number of ovules increased in female flowers at basal positions within the raceme. A consistent pattern of high seed set was observed in flowers from both field and cultivated populations. About 1 % of unfertilized ovules resulted from no pollination and 2 % of the seeds produced were only partly developed due to resource limitation. In the first inflorescence of the six experimental populations, 6.7-40.0 % of individuals produced only male flowers, and female flowers of 1.9-6.5 % individuals were aborted. The occurrence of male flowers in early blooming inflorescences could be an adaptive strategy to conserve resources and enhance pollination of female flowers in protogynous S. trifolia.     

EFFECTS OF DISPLAY SIZE,FLOWERING PHENOLOGY,AND NECTAR AVAILABILITY ON EFFECTIVE VISITATION FREQUENCY IN COMPARETTIA FALCATA (ORCHIDACEAE)

Comparettia falcata is an epiphytic, neotropical orchid that produces nectar as a pollinator reward. In Puerto Rico, C. falcata is allogamous and pollinated by the endemic hummingbird Chlorostilbon maugaeus. Autogamous pollinations are possible, but may result in reduced fruit set. For the 1989 and 1990 flowering seasons, the probability of pollinarium removals and natural pollinations increased with individual inflorescence display size. However, the frequency of effective pollinator visits was independent of flowering phenology in both years. A positive correlation between inflorescence size and reproductive success occurred in 1990 but not in 1989. In 1990 plants produced longer spurs, a higher standing crop of nectar, and a more concentrated nectar than in 1989. There was no relation between nectar availability (= standing crop of nectar) and sugar concentration in either year. Nectar availability and sugar concentration did not vary among the first four flowers of an inflorescence in either season. Nectar availability was not a good predictor of effective visitation. Comparettia falcata has a higher natural fruit set than tropical deceptive orchids, suggesting that pollinator visitation may be enhanced by nectar reward. The small, dilute nectar volumes secreted by C. falcata may benefit the plant by increasing interplant pollinator movement and pollen dispersal.     

Pollen and nectar as a reward in the basal epidendroid Psilochilus modestus (Orchidaceae: Triphoreae): A study of floral morphology, reproductive biology and pollination strategy

Psilochilus modestus Barb. Rodr. is a basal epidendroid orchid occurring in both the semi-deciduous and Atlantic rain forests of the state of São Paulo, southeastern Brazil. This species presents a perfect flowering synchrony within populations, since all the mature buds of each plant open simultaneously in the morning hours of the same day. These flowers are available only for 1 day and are pollinated by several species of small solitary and social native bees. These bees exploit both pollen and nectar as a reward. The bees collecting pollen promote a higher fruit set and perform mainly self-pollination while those collecting nectar, which are less numerous, are responsible for an increase in cross-pollinations. P. modestus is self-compatible but pollinator dependent. Natural fruit set (open pollination) is low when compared with the numbers obtained under manual self- and cross-pollination. Low fruit set in natural conditions is related to deficient pollen transfer, and pollinator inefficiency seems to be the main factor. Some factors, such as the small amount of nectar produced, the low number of flowers per inflorescence and their availability for 1 day only added to the perfect flowering synchrony seem to be responsible for the increase of cross-pollinations. The offering of both pollen and nectar as a reward can represent a transitional condition in basal Epidendroideae. Based on floral morphology, reward production and pollinator behavior, the relationship of P. modestus with the basal and most derived groups within Orchidaceae is discussed.     

Pollinators and Reproductive Success of the Wild Cucurbit Cucurbita maxima ssp. andreana (Cucurbitaceae)

Abstract: We studied the reproductive success and pollinators of Cucurbita maxima ssp. andreana in different disturbed habitats where it grows naturally. Data were obtained from three populations. One grew within a soybean crop, the other within a corn crop, and the third in an abandoned crop field. Cucurbita maxima ssp. andreana is an annual vine with a flowering period from December to April. Male flowers appear first, thereafter female and male flowers appear together. Flower lifetime (9 h) was similar in male and female flowers. The pollinator guild was comparable for the three populations but some differences in the frequency of the insect species were observed. Native bees were the main pollinators in the population in the abandoned field, while beetles pollinated the populations in crop fields. These differences were not linked with the pre-emergent reproductive success, fruit and seed set, or fruit quality. This is a self-compatible plant. Fruit and seed set and fruit traits (total mass, width and length of fruits, number of seeds per fruit, and seed mass) did not show significant differences between hand-cross and hand-self pollinated flowers. This wild cucurbit is a generalist with respect to pollinator guild, and flower visitors seem to be highly efficient in pollen transference. Cucurbita maxima ssp. andreana is well adapted to disturbed habitats because plants ripened fruits successfully, regardless of the group of insects visiting flowers.