Natural Pollination Research Articles

Suitable pollen source for the improvement of fruit and seed traits in Xanthoceras sorbifolium

Xanthoceras sorbifolium with high oil content in seeds has been used for various purposes, including as edible oil and feedstock for biodiesel production; however, the large-scale commercial cultivation of this species is severely hindered by the poor seed yield. As a self-incompatible species, whether a suitable pollen source can effectively improve yield and fruit traits of super clones has not been confirmed in X. sorbifolium. In the present study, cross pollination, mixed pollination, and natural pollination were conducted in five super clones. For all clones, the highest pollen germination was observed one day before flowering 84.26–90.25% (**p < 0.01). The final fruit set ratio (FFSR) was significantly influenced by the pollen source (*p < 0.05). Hand pollination had a positive effect on FFSR and seed yield (*p < 0.05). Fruit width and 1000-seed weight were significantly influenced by pollen source (*p < 0.05). Fruit weight, length, seed number per fruit, seed-producing ratio, seed yield, seed size, kernel ratio, and chemical components were significantly affected by the recipient × pollen source (**p < 0.01). In all combinations, the maximum seed yield occurred in the combination No. 2 × No. 3. The xenia effect occurred in the 1000-seed weight and kernel starch concentration. Based on these traits, the most suitable pollen source for each super clone was identified. This study suggests the significance of a suitable pollen source and collocation of pollinizer trees, especially in X. sorbifolium plantations, which can be applied in its cultivation and yield improvement.

Cacao flower visitation: Low pollen deposition, low fruit set and dominance of herbivores

Abstract Pollination services of cacao are crucial for global chocolate production, yet remain critically understudied, particularly in regions of origin of the species. Notably, uncertainties remain concerning the identity of cacao pollinators, the influence of landscape (forest distance) and management (shade cover) on flower visitation and the role of pollen deposition in limiting fruit set. Here, we aimed to improve understanding of cacao pollination by studying limiting factors of fruit set in Peru, part of the centre of origin of cacao. Flower visitors were sampled with sticky insect glue in 20 cacao agroforests in two biogeographically distinct regions of Peru, across gradients of shade cover and forest distance. Further, we assessed pollen quantities and compared fruit set between naturally and manually pollinated flowers. The most abundant flower visitors were aphids, ants and thrips in the north and thrips, midges and parasitoid wasps in the south of Peru. We present some evidence of increasing visitation rates from medium to high shade (40%–95% canopy closure) in the dry north, and opposite patterns in the semi‐humid south, during the wet season. Natural pollination resulted in remarkably low fruit set rates (2%), and very low pollen deposition. After hand pollination, fruit set more than tripled (7%), but was still low. The diversity and high relative abundances of herbivore flower visitors limit our ability to draw conclusions on the functional role of different flower visitors. The remarkably low fruit set of naturally and even hand pollinated flowers indicates that other unaddressed factors limit cacao fruit production. Such factors could be, amongst others, a lack of effective pollinators, genetic incompatibility or resource limitation. Revealing efficient pollinator species and other causes of low fruit set rates is therefore key to establish location‐specific management strategies and develop high yielding native cacao agroforestry systems in regions of origin of cacao.

Native Flowering Border Crops Attract High Pollinator Abundance and Diversity, Providing Growers the Opportunity to Enhance Pollination Services.

Over the past century, habitat loss from agricultural intensification has contributed to pollinator decline. One way to mitigate the harmful effects of agricultural intensification is through the re-introduction of native flowering plants as border strips that provide supplemental floral and nesting resources to pollinators. However, border crop species vary in bloom period and flower densities, and are thus likely to attract different suites of pollinator species. Resulting differences in pollinator community composition are likely to affect their ability to provide pollination services to adjacent crop habitat. To address these issues, we implemented a two-year study on the impact of different flowering border crops on pollinator abundance, richness, and community composition. We also examined which crop features (bloom duration, number of flowers, floral area) were most likely to affect pollinator densities. We found that native flowering plant border crops of diverse prairie mix and monocultures of silflower (Silphium integrifolium Michx.) and cup plant (Silphium perfoliatum L.) attracted the highest abundance and species richness of bees and pollinator groups combined, while alfalfa (Medicago sativa L.) attracted the highest lepidopteran abundance and species richness. We also found a significant, positive relationship between pollinator abundance and floral resource amount and bloom duration. These findings offer valuable insight into the impacts of different land management strategies on different pollinator groups, and thus provide landowners with management options for attracting specific pollinator groups and species.

Supplemental Stingless Bee Pollination in Fennel (Foeniculum vulgare Mill.): An Assessment of Impacts on Native Pollinators and Crop Yield

The yield of many crops benefits from pollinating insects; thus, recently reported declines in pollinator abundance and diversity are concerning for global food production. The pollinator dependency and amount of yield enhancement may vary according to crop species and geographical location. Fennel (Foeniculum vulgare) is an important spice crop cultivated in Indian states. However, comprehensive knowledge about pollination demand and yield enhancement potential of managed bees is still unknown. Here, we conducted a replicated study in two successive years (2020 and 2021) in West Bengal by combining pollinator surveys, pollinator-exclusion experiments, and field manipulation on fennel, which quantifies the impacts of supplemental stingless bees (Tetragonula iridipennis) pollination on native pollinators and crop yield. The crop species attracted many insect species belonging to diverse groups. Among those, important native pollinators (based on “approximate pollination value”) were Apis cerana, Apis dorsata, Apis florea, and Oxybelus furculatus in open condition (i.e., without field manipulation and in the absence of managed bees). We derived the coefficient of pollination deficit (D) from the fruit set percentages in open and manual cross-pollination treatments. The obtained value (D = 0.18) implies that the crop species have pollen transfer limitations, resulting in retardation of crop yield. From field manipulation with managed stingless bee colonies, the abundance of visitors (especially stingless bee foragers) on fennel increased (without altering the foraging activity of other native pollinators), thereby fruit set and crop yield increased by about 14.89 and 19.31%, respectively. Native managed stingless bees had no negative impacts on other native unmanaged species and can be promoted as complementary and short-term means of boosting yields and improving agricultural sustainability.

Is the High Proportion of Males in a Population of the Self-Incompatible Fraxinus platypoda (Oleaceae) Indicative of True Androdioecy or Cryptic-Dioecy?

Androdioecy is a rare reproductive system. Fraxinus platypoda, a woody canopy species in Japan’s mountainous riparian zones, is described as a morphologically androdioecious species. In this study, we tried to detect whether F. platypoda is also functionally androdioecious. We analyzed its sexual expression, seed development, pollen morphology and germination ability, pollination systems, and mast flowering behavior. We found that the hermaphrodite trees are andromonoecious, with inflorescences bearing male and hermaphroditic flowers, whereas male individuals had only male flowers. Pollen morphology was identical in male flowers, in hermaphrodite flowers of an andromonoecious individual, and in male flowers of male individuals. Pollen from both types of individuals was capable of germination both ex vivo (on nutrient medium) and in vivo in pollination experiments. However, compared with pollen from andromonoecious trees, pollen from male trees showed a higher germination rate. The self-pollination rate of bagged hermaphroditic flowers was almost zero. The fruit set rate following cross-pollination with male pollen from a male tree was higher than that following natural pollination, whereas the rate with hermaphroditic pollen was the same. The flowering and fruiting of F. platypoda have fluctuated over 17 years; the flowering of the two types of sexual individuals exhibited clear synchronization during this period. The frequency of male individuals within the populations is 50%. The maintenance of such a proportion of males in populations of the self-incompatible F. platypoda is either indicative of a true androdioecious species with a diallelic self-incompatibility system or a cryptic-dioecious species. This alternative is discussed here.

Effective and feasible mechanisms to support native invertebrate pollinators in agricultural landscapes: A meta‐analysis

AbstractPollinator declines have emerged as a major conservation concern across a wide diversity of systems and taxa worldwide. In response to these concerns, active efforts to conserve pollinators are increasingly considered in agricultural landscape design and management, natural area management, and public land decision‐making. An important conservation challenge is translating these efforts to feasible recommendations that agriculturalists across a diversity of systems and contexts can employ to support pollinator communities. In order to develop such a toolbox for agriculturalists, we conducted a meta‐analysis to examine feasible management practices, tools, or decisions, employed in agricultural landscapes, that directly impact pollinator forage and nest site resources. Our search terms netted a total of 743 unique references; from this, we developed a dataset of 209 unique records, derived from 89 unique references, for inclusion in our meta‐analysis. We found that across all treatments aimed at boosting pollinators, there were significant increases in bee abundance, bee species richness, and overall pollinator abundance and visitation rate. We also found specifically that across all study types, the following practices resulted in positive changes in pollinator community responses: reduced distance to natural areas, reduced regional intensity of human modification, increased quantity and quality of pollinator resources within a focal system, plantings adjacent to agricultural areas, and the use of nonconventional agricultural practices. The most consistent pollinator response to treatments was bee species richness, which was significantly increased by plantings adjacent to agricultural areas, distance from natural areas, and quantity and diversity of pollinator resources. Bee species richness also decreased with intensity of human modification. These findings are presented in a toolbox for agriculturalists, containing evidence‐based actions and outcomes to assist practitioners in decision‐making specific to pollinator habitat improvement that could be actionable in the near term.

Short Communication: Impact of introduction of managed honey bee colony on wild bee diversity and abundance in an agroecosystem in Indonesia

Abstract. Widhiono I, Sudiana E, Suryaningsih S. 2022. Short Communication: Impact of introduction of managed honey bee colony on wild bee diversity and abundance in an agroecosystem in Indonesia. Biodiversitas 23: 1099-1104. Wild bees are natural pollinators of crops in agroecosystems, but managed honeybee colonies may diminish their diversity and abundance. We aimed to determine the impact of introducing managed honeybee (Apis cerana and A. mellifera) colonies on the diversity and abundance of wild bees in an agroecosystem in Indonesia. We introduced managed honeybee colonies into the plots containing four crops (tomato, green bean, chili, and green mustard). Sampling was conducted from March to July 2020 on the eastern slopes of Mount Slamet, Indonesia, at an elevation of 1241 m asl. Wild bee diversity was not affected by the introduction of managed honeybee colonies, but their abundance decreased at the end of the flowering period due to resource limitations. The most abundant wild bee species were Amegilla cyrtandrae and A. burneensis, and some wild bee species tended to be more abundant on certain crop species. We recommend that managed colonies be introduced during the early and mid-stages of crop flowering to reduce potential adverse effects on wild bee populations.

Efficacy of hybrid seed production through natural pollinators employing male sterility: Case study of ridge gourd [Luffa acutangula (Roxb.) L.] CGMS system

Potential of natural pollinators and manually assisted pollination for producing F1 hybrid seeds of ridge gourd hybrid ‘KRH 1’ was evaluated under open field condition for two seasons by growing male sterile line (KAU-LA-MS-101) and monoecious male fertile line (LA-102) in alternate rows. Investigation was carried out at the Department of Vegetable Science, College of Agriculture, Vellanikkara, Thrissur, India. Species diversity of the pollinators, peak foraging time of the pollinators, number of floral visits by pollinating bees and number of bees visiting a flower during stigma receptivity were recorded. Nectary glands of male and female flowers of KAU-LA-MS-101 and LA 102 were measured. Seed yield and seed quality related traits were also recorded following standard protocol. Male sterile plants produce small male flowers with tiny nectary glands when compared to that of LA- 102, but female flowers of male sterile line have larger nectary glands. Stingless bee (Tetragonula sp.), was the pollinator observed in the field during anthesis time with peak activity after 5.30 pm. Number of fruits, number of seeds per fruit and average seed weight were significantly different between male sterile line and monoecious male fertile line LA-102. Pollination efficiency of bees found to be high when compared to manual hands in terms of quality of seeds produced from male sterile plants. However manual hands proved to be better for inducing more number of seeds per fruit. Total seed yield per plant depends on the total number of fruits developed and pollination by bees induces the development of more flowers to maturity, thereby ensuring competitive seed yield. Seed yield per plant exhibited significant positive correlation with seed length, shoot length, root length, fresh weight, vigour index I and germination velocity index. Present investigation revealed the potential of male sterile line developed in ridge gourd for hybrid seed production. Alternate planting of fertility restorer line with the MS line and allowing open pollination with the assistance of bees will reduce the labour charge involved in hybrid seed production and quality seed can be made available at a competitive rate.

Reproductive biology of the invasive Gleditsia triacanthos L. (Fabaceae)

Biotic invasions constitute a threat for natural communities, affecting ecological and evolutionary integrity of natural ecosystems. Gleditsia triacanthos is an invasive species in many regions of the world. In Argentina, its populations are expanding over native vegetation of different ecoregions. The aim of this work is to study the reproductive biology of the invasive honey locust, characterizing the reproductive individuals, the assemblage of pollinators, the pollination process, the compatibility system, and the reproductive success. Considering that G. triacanthos could be characterized as a novel invader in Argentina and plant-pollinator interactions could be important to facilitate plant invasions, we expect to find (a) an assembly of generalist floral visitors that promote pollination; (b) that fruits and seeds could be produced through self- (autogamy) or cross-pollination (mediated by pollinators); (c) that pollinators can increase reproductive success of exposed flowers compared to those experimentally isolated from pollinators. The studied populations (Córdoba Province, central Argentina) can be characterized as polygamous andro-dioecious, with a higher proportion of male individuals. Although honeybees were the most frequent pollinator, all floral visitors could be potential pollinators and the honey locust can be characterized as pollinator-generalist. G. triacanthos produces fruits and seeds by spontaneous self-pollination, but a higher reproductive success was observed after hand cross- or natural pollinations. Producing both self- and cross-pollinated seeds could be advantageous for this species during different stages of the invasion process. The generalist pollination system presented by G. triacanthos, the high production of pollen by male individuals compared to the individuals with perfect flowers, and the higher fruit and seed set by cross- or natural pollinated flowers are all traits favouring the invasion process. These reproductive traits could be considered when designing management strategies to control the expansion of this invasive species.

Analysis of The Diversity Of Butterfly (Rhopalocera) Based On Environmental Conditions In Muria Kudus Tourism Area Central Java

Mount Muria has abundant natural resources, both flora and fauna, one of which is butterflies. Butterflies as natural pollinators have an important role in the ecosystem. The deterioration of air quality has a negative impact on butterflies because they are quite sensitive to poor air quality and poor lighting. This study aims to analyze the relationship between environmental conditions and butterflies in the Muria Kudus area, carried out in March and June 2020. Butterfly observations were carried out using the quadratic sample transect method with 3 observation stations and measured environmental factors, among others: temperature, humidity, light intensity, altitude and Pb concentration analysis with AAS. The results of the study found 40 species of butterflies consisting of 6 families, with a moderate diversity index, almost merit index and low dominance. Muria area still has good environmental conditions for butterfly breeding. Relative abundance is negatively correlated with light intensity (rcount = -0,855, rtable = 0,811). Averageness is negatively correlated with heigh (rcount=-0,836, rtable=-0,881). The butterfly species Ypthima nigricans, Orsotriaena medus, and Melanitis leda is a specialist butterfly species and the butterfly species Papilio memnon, Leptosia nina, Chersonesia rahria, Mycalesis horsifieldii, and Yphtima horsifieldii is a generalist species butterfly

Features of cost accounting and calculation of the cost of production of honey-pollinating direction of beekeeping

Beekeeping has a high value due to the commercial products obtained and the creation of opportunities for natural pollination of agricultural crops in order to increase their yield. Based on this, intensive farming territories have a great potential in the development of the industry. The article describes the features of calculating the products of honey-pollinating direction of beekeeping; the composition and structure of production costs, the procedure for calculating the actual cost of production of honey-pollinating direction of beekeeping are considered. The subject of the study is the system of cost accounting and calculation of the cost of production of honey-pollinating direction of beekeeping in the Stavropol Territory. The information base was the forms of specialized reporting of agricultural organizations of the Stavropol Territory. The scientific novelty of the research consists in the development of methods for calculating the cost of beekeeping products of an economy that has a honey-pollinating direction.

Assessment of in Vitro Pollen Germination and Pollen Tube Growth of Annona Cherimola Mill

ABSTRACT Yield in cherimoya (Annona cherimola Mill.) is affected by various factors reducing fertilization and fruit production such as: the protogynous dichogamy of the flowers, deficient natural pollination, and low pollen quality. The objective of this study was to determine in vitro the differences in pollen quality of nine ecotypes and one cultivar of A. cherimola from pollen samples collected at the female and male phenological stages of flowering. The percentage of pollen germination and the length of pollen tubes was evaluated in vitro at 2 and 24 hours after incubation at 21°C, in two phenological stages of the flowers during bloom in 2018 and 2019 seasons. The germination medium contained: 10% sucrose, 1% agar, 0.005% boric acid, and 0.025% calcium nitrate. Fabulosa and Austro ecotypes and the Fino de Jete cultivar had the highest pollen germination in the male phenological stage, after 2 and 24 hours in both seasons. Fabulosa had the highest percentage of germination in the female stage in both seasons. The ideal phenological stage for pollen collection was at anther dehiscence. Austro and Fabulosa ecotypes had the longest pollen tubes after 2 and 24 hours, in both phenological stages for both seasons. Fabulosa and Austro ecotypes, and Fino de Jete cultivar, showed promising pollen characteristics for hand pollination of cherimoya.

Natural and artificial pollination of white-fleshed pitaya

To produce pitaya (Hylocereus spp.), it is essential that pollination occur, either artificially or by pollinating agents. This study evaluated the viability of self-pollination, cross-pollination, and natural pollination, as well as pollen viability, stigma receptivity and ideal pollination window in pitaya flowers (Hylocereus spp.). An experiment was conducted with a randomized block design consisting of four treatments: T1: manual self-pollination; T2: nocturnal open pollination; T3: diurnal open pollination; and T4: manual cross-pollination - red-fleshed pitaya [ Hylocereus polyrhizus] pollen placed on white-fleshed pitaya [ Hylocereus undatus] stigma. The experiment had four replicates, with two plants per replicate (two flowers per plant), totaling 16 flowers per treatment. The analyzed variables were fruit weight, length, diameter, titratable acidity, pH, soluble solids, and SS/TA ratio; the germination percentage of pollen grains; and the receptivity of stigmas. Artificial pollination (self- and cross-pollination) of white-fleshed pitaya (Hylocereus undatus) is viable, resulting in larger fruits than natural pollination, with cross-pollination being the most recommended. Throughout the floral opening period, the pitaya flower (Hylocereus spp.) has receptive stigma capable of receiving pollen. The flowers exhibit the highest pollen germination rate at 7 p.m, which is the ideal pollination window for pitaya.

Produtividade de maracujá-amarelo orgânico em função da irrigação, cultivo semiprotegido e polinização artificial

Abstract This study aimed to evaluate the effect of the combination between irrigation, semiprotected cultivation, and artificial pollination on the yield, number of fruits, and mass of yellow passion fruit grown under organic production. The experimental design was in randomized blocks set up in split plots (2x2x2) with eight treatments and four replications. The plot consisted of irrigation or rainfed conditions, the second factor corresponds to plant cover with plastic or direct sunlight, and the third factor refers to natural or artificial pollination. Irrigation was performed using a micro-sprinkler system, and pollination was either manual or natural (entomophilous). The evaluated parameters were the number of fruits per plant, the mean fruit mass, and the total and commercial yields in two crop years: from January to August 2019 and from September 2019 to August 2020. The yield per hectare and the number of fruits per plant were not affected by irrigation and semi-protected cultivation. Artificial pollination increased the passion fruit yield by up to 31% under organic cultivation. The number of fruits per plant did not significantly change with the treatments. However, there was a positive linear correction with the yield. The mean fruit mass is higher using the combination between semi-protected cultivation, rainfed conditions, and artificial pollination or direct sunlight with natural pollination regardless of irrigation.

Floral biology and reproductive system of physic nut (Jatropha curcas L., Euphorbiaceae) in ‘Recôncavo da Bahia’, Brazil

Understanding the floral and reproductive biology of botanical species is crucial for the development of strategies in plant breeding systems. Jatropha curcas L. is a promising species for the manufacture of biofuels, being previously studied mainly in genetic improvement to develop characteristics suitable to biofuels. In order to contribute with data for hybridization and breeding programs, this paper studies the floral biology and reproductive system in two experimental populations of different ages of Jatropha curcas in Cruz das Almas/BA. Both of them were examined about their anthesis, durability, number of flowers, stigma receptivity, P:O ratio, and reproduction tests. As observed, Jatropha curcas is a monoecious species and its flowering occurs between September and April. Inflorescences are composed of unisexual flowers with daytime anthesis (♂: 05-06h; ♀: 07-08h), where the staminates last 10h and pistillates 60h. The physic nut is self-compatible, forming fruits by self-fecundation and cross-pollination, although the greatest number of fruits/seeds is generated by natural pollination. Experiment 02, presented a larger number of flowers, probably due to the plant's age and physiology. Performing artificial pollination between 08:00h and 09:30h is recommended for larger production since the stigma is receptive and the flowers have a large amount of pollen available.

Inter-Annual Changes in Abundance of Native and Exotic Pollinators of V. Faba Crops and Their Relationship with Landscape Variables

Inter-Annual Changes in Abundance of Native and Exotic Pollinators of V. Faba Crops and Their Relationship with Landscape Variables

Diversity and Effectiveness of Native Insect Pollinators in Carrot Seed Production

Diversity and Effectiveness of Native Insect Pollinators in Carrot Seed Production

Risks and opportunities associated with pollinators’ conservation and management of pollination services in Latin America

The conservation status of pollinators and pollination in Latin America (LA) is reviewed. The knowledge regarding native and managed pollinators (e.g., honeybee and stingless bees) and pollination services was synthetized, and the guidelines to improve the opportunities for conservation are provided, considering the threats to pollinators and the perspectives from traditional and local knowledge. The analysis indicates that diverse threats (e.g., large-scale agriculture, deforestation, overuse of agrochemicals) are linked with pollination and pollinator decline, which affect the reproduction of most native plants and the yields of many crops. LA harbours the highest bee diversity worldwide, with 26% of the total recorded species, and it is a biodiversity hotspot of vertebrate pollinators, including hummingbirds, perching birds, nectarivorous bats and other mammal pollinators. Specific recommendations to conserve native pollinators and to improve pollination services are provided, which could be considered by stakeholders and governments aiming to elaborate biocultural conservation. For example, introducing policies and legal responses for incentives to help farmers maintain natural habitats and forests, to replace or reduce agrochemicals and to improve diversified crop production with agroecological practices; refining agrochemical regulations to minimize the exposure of pollinators to insecticides and herbicides; improving knowledge and education on pollinators and pollination gives societies worldwide the opportunity to change current hegemonic agricultural practices and consumption patterns; integrating different land ethical views of ethnic minorities on a sustainable relationship between production and biodiversity. A wider view combining social, ecological, cultural dimensions may support better decision making. This holistic socio-agroecological perspective is urgently needed to conserve and manage pollinators at different spatial and temporal scales, and to integrate pollination services, pollinator-friendly habitat management approaches and diversified farming systems.

Assessing the labor productivity of two methods of artificial pollination in oil palm crops from Colombia

Oil palm interspecific hybrids Elaeis oleífera × Elaeis guineensis (O × G) are grown across approximately 68,000 hectares in Colombia. To address the limited natural pollination capacity of O × G hybrids and the difficulties associated with assisted pollination regarding the timing of E. guineensis pollen application, the Colombian Oil Palm Research Center (Cenipalma) conducted studies on induction of parthenocarpic fruits. Cenipalma confirmed that application of 1,200 ppm of 1-naphthaleneacetic acid (NAA) at different phenological stages enabled the formation of parthenocarpic fruits. This technological advance was termed artificial pollination. This paper presents the results from a research study aimed at assessing the labor productivity for two methods of NAA application (NAA in solid mixture and NAA in liquid suspension). From a methodological standpoint, a time and motion study was conducted to assess labor productivity for each NAA application method, with time data collected using the software Cybertracker on a mobile platform (smartphone). The results indicated that a worker can cover 3.9 hectares in a working day and sprinkle 303 inflorescences in a working day when applying NAA in solid mixture. On the other hand, when applying NAA in liquid suspension, the worker can cover 3.2 hectares and sprinkle 315 inflorescences (i.e. in a working day).

Review on Reproductive Biology of Caster bean (Ricinus communis L.)

Castor belongs to the genus Ricinus, a member of the Euphorbiaceae, which contains a vast number of plants mostly native to the tropics. The genus Ricinus is considered to be monotypic and R. communis is the only species, which includes many polymorphic types. The cultivated types are dwarf annuals. Castor plants have well developed root system with thick horizontal roots. Tap root looks like extension of the stem below the soil. The stem is erect, circular in section, partially hollow, glabrous, and smooth with good branching. Stem is either red or green or with shades of both. The stem is marked by well-defined nodes from each of which a leaf is arises. The lower internodes are shorter and their length increases with height. Leaves are alternate, large, and palmate with 5-11 lobes acuminate margins notched, serrate or indented. They are carried on long stout petioles. The inflorescence is borne terminally on the main and lateral branches. Flowers are large, in terminal sub panicled racemes, monoecious, apetalous, the upper portion of the raceme being occupied by the female flowers and the lower by male flowers the whole inflorescence may reach a length of 90 cm. The flowers are wind and insect pollinated and from 5 to 46 % natural pollination occurred. The fruit is round glucose capsules with three projecting sides covered with tough spines or smooth, 3 loculed and three seeded. Seeds are albuminous, anatropous, broad, oval, compressed with a marked caruncle and longitudinal raphe. The testa is thin, brittle, varying in color and mottling. Below the testa are thin legmen, covering a whitish oily endosperm containing the embryo.