Stigmatic exudate in the Annonaceae: Pollinator reward,pollen germination medium or extragynoecial compitum?

Although “dry‐type” stigmas are widely regarded as ancestral in angiosperms, the early‐divergent family Annonaceae has copious stigmatic exudate. We evaluate three putative functions for this exudate: as a nutritive reward for pollinators; as a pollen germination medium; and as an extragynoecial compitum that enables pollen tube growth between carpels. Stigmatic exudate is fructose dominated (72.2%), but with high levels of glucose and sucrose; the dominance of hexose sugars and the diversity of amino acids observed, including many that are essential for insects, support a nutritive role for pollinators. Sugar concentration in pre‐receptive flowers is high (28.2%), falling during the peak period of stigmatic receptivity (17.4%), and then rising again toward the end of the pistillate phase (32.9%). Pollen germination was highest in sugar concentrations <20%. Sugar concentrations during the peak pistillate phase therefore provide optimal osmolarity for pollen hydration and germination; subsequent changes in sugar concentration during anthesis reinforce protogyny (in which carpels mature before stamens), enabling the retention of concentrated exudate into the staminate phase as a pollinator food reward without the possibility of pollen germination. Intercarpellary growth of pollen tubes was confirmed: the exudate therefore also functions as a suprastylar extragynoecial compitum, overcoming the limitations of apocarpy.     

The mechanism of delayed selfing in Collinsia verna (Scrophulariaceae)

Collinsia verna, blue-eyed Mary, has floral attributes of an outcrossing species, yet most flowers readily self-pollinate under greenhouse conditions. Here we describe the mechanism of self-pollination in C. verna via changes in relative positions of the stigma and anthers and late timing of receptivity, resulting in delayed selfing. Each flower contains four anthers that dehisce sequentially over ∼1 wk. Pollen that is not collected by pollinators accumulates in the keel petal and retains high viability (>80% pollen germination) up to the time of corolla abscission. The stigmatic surface does not become receptive until after the third anther dehisces. This overlap in the sexual phases is concurrent with a change in herkogamy during floral development. In most flowers (70%), the stigma has moved to the front of the keel and is positioned near the anthers when the third anther dehisces. Under field conditions, fruiting success of plants within pollinator exclosures was ∼75% of the fruiting success in open-pollinated plants (33% fruiting success via autogamy vs. 44% fruiting success, respectively). Collinsia verna plants in pollinator exclosures exhibit variation in autogamy rates within natural populations (range 0–80%). In addition, only half of naturally pollinated, receptive flowers examined had pollen tubes growing in their styles. In contrast, shortly after corolla abscission, nearly all flowers examined (96%) had pollen tubes in their styles. Thus we find that in C. verna, autogamy occurs late in floral development, which has the potential to provide substantial reproductive assurance, and that individuals vary in their ability to set fruit through this mechanism. We suggest that delayed selfing mechanisms may be overlooked in other species and that variable pollinator availability may play a significant role in the maintenance of mixed mating in species with delayed selfing, such as C. verna.     

Specialised protagonists in a plant–pollinator interaction: the pollination of Blumenbachia insignis (Loasaceae)

• Analyses of resource presentation, floral morphology and pollinator behaviour are essential for understanding specialised plant‐pollinator systems. We investigated whether foraging by individual bee pollinators fits the floral morphology and functioning of Blumenbachia insignis, whose flowers are characterised by a nectar scale‐staminode complex and pollen release by thigmonastic stamen movements.
• We described pollen and nectar presentation, analysed the breeding system and the foraging strategy of bee pollinators. We determined the nectar production pattern and documented variations in the longevity of floral phases and stigmatic pollen loads of pollinator‐visited and unvisited flowers.
• Bicolletes indigoticus (Colletidae) was the sole pollinator with females revisiting flowers in staminate and pistillate phases at short intervals, guaranteeing cross‐pollen flow. Nectar stored in the nectar scale‐staminode complex had a high sugar concentration and was produced continuously in minute amounts (~0.09 μl·h^?1). Pushing the scales outward, bees took up nectar, triggering stamen movements and accelerating pollen presentation. Experimental simulation of this nectar uptake increased the number of moved stamens per hour by a factor of four. Flowers visited by pollinators received six‐fold more pollen on the stigma than unvisited flowers, had shortened staminate and pistillate phases and increased fruit and seed set.
• Flower handling and foraging by Bicolletes indigoticus were consonant with the complex flower morphology and functioning of Blumenbachia insignis. Continuous nectar production in minute quantities but at high sugar concentration influences the pollen foraging of the bees. Partitioning of resources lead to absolute flower fidelity and stereotyped foraging behaviour by the sole effective oligolectic bee pollinator.

     

Anther structure and pollen development in Melicoccus lepidopetalus (Sapindaceae): An evolutionary approach to dioecy in the family

Anther and pollen development in staminate and pistillate flowers of dioecious Melicoccus lepidopetalus (Sapindaceae) were examined by light and electron microscopy. Young anthers are similar in both types of flowers; they consist of epidermis, endothecium, two to four middle layers and a secretory tapetum. The microspore tetrads are tetrahedral. The mature anther in staminate flowers presents compressed epidermal cells and endothecium cells with fibrillar thickenings. A single locule is formed in the theca by dissolution of the septum and pollen grains are shed at two-celled stage. The mature anthers of pistillate flowers differ anatomically from those of staminate flowers. The epidermis is not compressed, the endothecium does not develop fibrillar thickenings, middle layers and tapetum are generally persisting, and the stomium is nonfunctional. Microspore degeneration begins after meiosis of microspore mother cells. At anthesis, uninucleate microspores and pollen grains with vegetative and generative nuclei with no cytokinesis are observed. Some pollen walls display an abnormal exine deposition, whereas others show a well formed exine, although both are devoid of intine. These results suggest that in the evolution towards unisexuality, the developmental differences of anther wall tissues and pollen grains between pistillate and staminate flowers might become more pronounced in a derived condition, such as dioecy.     

FUNCTIONAL DIOECY AND ANDROMONOECY IN SOLANUM

Field and laboratory studies of 19 diclinous species endemic to Australia help to clarify the nature and evolution of andromonoecy, androdioecy, and dioecy in the genus Solanum. Ten species are andromonoecious; typically these species bear inflorescences with a single, large basal hermaphroditic flower and 12–60 distal, smaller staminate flowers. We suggest that the andromonoecious condition was derived from hermaphroditic-flowered ancestors in part by hemisterilization of flowers but largely by addition of staminate flowers. The resultant larger inflorescences are hypothesized to serve both to attract and to entrain pollinators, yielding more or higher-quality seed set in hermaphroditic flowers and/or greater dispersion of pollen from staminate flowers. We suggest that andromonoecy may also serve to reduce selling. Nine other species are morphologically androdioecious but functionally dioecious. In these species, staminate flowers, like those of the andromonoecious species, bear anthers with copious tricolporate pollen and a highly reduced gynoecium. The morphologically hermaphroditic flowers are functionally pistillate and borne singly in inflorescences, and they bear anthers with inaperturate pollen. The inaperturate pollen, although viable, never germinates and is hypothesized to be retained in pistillate flowers as a reward to pollinators in the nectarless Solanum flowers. All other species of Solanum studied with pollen dimorphism in which one pollen morph is inaperturate are also best treated as functionally dioecious. We conclude that there is no evidence for androdioecy in Solanum. A review of other families suggests that there is little support for this unusual breeding system in any other angiosperm group either. Preliminary analyses suggest that andromonoecy and dioecy are polyphyletic in Solanum. Furthermore, dioecy is as likely to have arisen from hermaphroditic as from andromonoecious ancestors.     

MORPHOLOGY,VASCULAR ANATOMY AND EMBRYOLOGY OF PISTILLATE AND STAMINATE FLOWERS OF ASPARAGUS OFFICINALIS

Flowers of three pistillate (female), two heterogametic staminate (male) and two homogametic male genotypes of Asparagus officinalis L. were compared for morphology and vascular anatomy of the flower and for embryological development to the stage of mature ovules and pollen. Flowers are liliaceous, the staminate with rudimentary pistils and the pistillate with collapsed anthers. The uncomplicated vascular pattern differs between staminate and pistillate flowers only in the size and degree of maturation of bundles to stamens and carpels. Longer styles appear to be correlated with a greater extent of ovule development in ovaries of staminate flowers. Microsporogenesis in males is normal with wall development corresponding to the Monocotyledonous type. The tapetum is glandular and binucleate, cytokinesis successive, the tetrads isobilateral or occasionally decussate, and the mature pollen grain two-celled. A pair of heteromorphic, possibly sex, chromosomes was observed in heterogametic male plants. Anther development is initially the same in pistillate flowers, but the tapetum degenerates precociously followed by collapse of microspore mother cells. In pistillate flowers the ovules are hemitropous, bitegmic, and slightly crassinucellate. Megasporogenesis-megagametogenesis conforms to the Polygonum type. In staminate flowers ovule development is like that in pistillate flowers until degeneration starts in nucellar and integumentary cells at the chalazal end. Ovules in both homogametic male genotypes rarely complete meiosis, while in the heterogametic males it is normally completed with about one ovule in 20 flowers forming a mature megagametophyte. Since manipulation of sex expression in Asparagus could be important in developing inbred male and female lines for breeding purposes, those aspects of the morphological and embryological observations presented which might be useful in planning experiments to induce sex changes are discussed briefly.     

Flower morphology and biology of Myrsine laetevirens, structural and evolutionary implications of anemophily in Myrsinaceae

This study deals with the phenology, pollination biology and floral morphology of Myrsine laetevirens , a neotropical dioecious tree. In Punta Lara (Argentina), its flowering period occurs during January-February. Both pistillate and staminate flowers are small, with a yellowish-green perianth and produce neither nectar nor odour. Staminate flowers have five stamens and a vestigial gynoecium while pistillate flowers, have non-functional anthers and a conspicuous stigma. The floral characteristics of staminate and pistillate plants are related to the syndrome of anemophily. Fruit set in inflorescences covered with mesh bags and observations prove that animals are not involved in the pollination process. A provisional cladistic analysis of Myrsinaceae shows that dioecy evolved as one of the most recent apomorphies of Myrsine and is part of the anemophilous syndrome.     

SEX DIFFERENTIAL FLORAL LONGEVITY,NECTAR SECRETION,AND POLLINATOR FORAGING IN A PROTANDROUS SPECIES

Sex differential nectar production, floral longevity and pollinator foraging were examined in Lobelia cardinalis, a self-compatible, protandrous species that is hummingbird pollinated. The staminate phase of the flowers lasts significantly longer and produces significantly more nectar (total sugar) per day than the pistillate phase of the flowers. Additional pollen is presented throughout the staminate phase. Because inflorescences of L. cardinalis mature acropetally, the nectar reward on any given day is greatest at the top of the inflorescence (where staminate phase flowers are located). Hummingbirds appear to be sensitive to this pattern of nectar presentation as they most commonly began foraging in the middle of an inflorescence and proceeded upward. This foraging pattern tends to promote outcrossing and suggests that staminate phase flowers are visited more often than pistillate phase flowers. We conclude that L. cardinalis emphasizes the male function at anthesis. Others have hypothesized that the features of this species are a logical consequence of intrasexual selection, but further research is needed before we place great confidence in a sexual selection interpretation of our data.     

Protandry,synchronized flowering and sequential phenotypic unisexuality in neotropical Pentagonia macrophylla (Rubiaceae)

Summary The neotropical shrub Pentagonia macrophylla Benth. (Rubiaceae) has protandrous two-day flowers. Synchronous development among flowers on a single individual results in sequential phenotypic unisexuality: the entire plant alternates gender from day to day. Pistillate flowers produce more nectar than staminate flowers at comparable hours, but this difference does not result in different paterns of visitation to male and female flowers by hummingbird pollinators. Rare periods of bisexuality occur due to asynchronous floral development within or between inflorescences and are always followed by return to a synchronized pattern of alternation of gender. Reestablishment of synchrony usually involves a timeframe shift in the pattern of flowering (i.e., a plant produces staminate flowers on days when it previously would have been pistillate). It is suggested that timeframe shifts occur in response to inadequate pollination and serve to either desynchronize the plant from neighboring conspecifics or to temporarily allow self-pollination.     

Resource partitioning to male and female flowers of Spinacia oleracea L. in relation to whole-plant monocarpic senescence

Male plants of spinach (Spinacea oleracea L.) senesce following flowering. It has been suggested that nutrient drain by male flowers is insufficient to trigger senescence. The partitioning of radiolabelled photosynthate between vegetative and reproductive tissue was compared in male (staminate) versus female (pistillate) plants. After the start of flowering staminate plants senesce 3 weeks earlier than pistillate plants. Soon after the start of flowering, staminate plants allocated several times as much photosynthate to flowering structures as did pistillate plants. The buds of staminate flowers with developing pollen had the greatest draw of photosynthate. When the staminate plants begin to show senescence 68% of fixed C was allocated to the staminate reproductive structures. In the pistillate plants, export to the developing fruits and young flowers remained near 10% until mid-reproductive development, when it increased to 40%, declining to 27% as the plants started to senesce. These differences were also present on a sink-mass corrected basis. Flowers on staminate spinach plants develop faster than pistillate flowers and have a greater draw of photosynthate than do pistillate flowers and fruits, although for a shorter period. Pistillate plants also produce more leaf area within the inflorescence to sustain the developing fruits. The (14)C in the staminate flowers declined due to respiration, especially during pollen maturation; no such loss occurred in pistillate reproductive structures. The partitioning to the reproductive structures correlates with the greater production of floral versus vegetative tissue in staminate plants and their more rapid senescence. As at senescence the leaves still had adequate carbohydrate, the resources are clearly phloem-transported compounds other than carbohydrates. The extent of the resource redistribution to reproductive structures and away from the development of new vegetative sinks, starting very early in the reproductive phase, is sufficient to account for the triggering of senescence in the rest of the plant.     

The role of autonomous self-pollination in floral longevity in varieties of Impatiens hypophylla (Balsaminaceae)

The floral longevity of unpollinated, hand self-, and hand cross-pollinated flowers was compared in two varieties of Impatiens hypophylla, which contrasts with their mating systems. When flowers were emasculated and hand-pollinated every day after anthesis, their longevity was reduced. In the absence of emasculation and hand pollination, the staminate phase of the flowers of both varieties was 1 d longer. After the staminate phase, flowers of the outcrossing variety dropped their androecium, exposing the stigma and initiating the pistillate phase, which lasted for ～2 d. In contrast, flowers of the mixed-mating variety self-pollinated autonomously and then terminated their flowering. Under great seasonal variation in the pollinator visitation rate, which was observed in their natural populations, the outcrossing variety should maximize expected outcross success through the phenology of floral sex phases, whereas the mixed-mating variety self-pollinated ovules that were not outcrossed within the staminate phase. Based on these results, I suggest that the autonomous self-pollination in I. hypophylla induced differences both in the selfing coefficient and in floral longevity between the varieties.     

POLLEN REMOVAL AND POLLEN DEPOSITION AFFECT THE DURATION OF THE STAMINATE AND PISTILLATE PHASES IN CAMPANULA RAPUNCULOIDES

We examined factors affecting the duration of the staminate and pistillate phases in the protandrous flowers of Campanula rapunculoides L. (Campanulaceae). Under conditions of natural pollinator visitation, flowers experiencing low rates of pollen removal lasted significantly longer than flowers that had faster rates of pollen removal. Experimental manipulations showed that low levels of pollen removal resulted in extension of the staminate phase. Hand-pollinations in which we varied the amount and source of pollen showed that when the number of fertilized ovules within an ovary is low, senescence of the flower is delayed, resulting in extension of the pistillate phase. We also report on pollinator foraging patterns within the vertical inflorescences of C. rapunculoides and the limiting factor for seed set in this population. The results are relevent to recent suggestions that floral characters often serve to reduce interference between the sexual functions in cosexual plants.     

Pollination Biology in a TropicalAcaciaHybrid (A. mangiumWilld. xA. auriculiformisA. Cunn. ex Benth.)

Floral biology and pollination mechanism of theAcaciahybrid(A. mangiumWilld. xA. auriculiformisA. Cunn. ex Benth.) growingin Thailand are investigated using light and electron microscopy.The hybrid is andromonoecious. A floral spike consists of about150 loosely arranged flowers. Flowers are cream coloured, fragrantand have no floral nectaries. The pistil has a solid style witha smooth, wet stigma and amphitropous ovules with immature integuments.The anther consists of eight loculi, each bearing only one 16-grainpolyad. The flowers are weakly protogynous. Anthesis is completeat 0500–0600 h but peak female receptivity begins at 0200–0300h and is completed that day. The stigmatic exudate is of thelipophilic type and is secreted from the stigmatic cells bya holocrine mechanism. Pollen is the main floral reward forthe insect pollinators. There are several floral characteristicswhich facilitate pollen transfer from anthers and depositionon stigmas.Apis melliferaandCeratinasp. are the most effectivepollinators because they are the most common visitors and carrya heavy load of hybrid polyads. However, their behaviour inforaging for pollen in the same tree and weak protogynous dichogamymay promote self-pollination in the hybrid. The hybrid has lowpollination success due to low pollinator number. An increasein exposure time of flowers to pollinators or pollinator numbermay increase pollination success but may not affect the rateof pollen deposition on stigmas due to the relatively smallsize of the stigma in relation to the polyad.Copyright 1998Annals of Botany Company Acaciahybrid,Acacia mangium, Acacia auriculiformis, pollination, pistil receptivity, anthesis, pollinators.     

Morphological variation in the flowers of Jacaratia mexicana A. DC. (Caricaceae), a subdioecious tree

The Caricaceae is a small family of tropical trees and herbs in which most species are dioecious. In the present study, we extend our previous work on dioecy in the Caricaceae, characterising the morphological variation in sexual expression in flowers of the dioecious tree Jacaratia mexicana . We found that, in J. mexicana , female plants produce only pistillate flowers, while male plants are sexually variable and can bear three different types of flowers: staminate, pistillate and perfect. To characterise the distinct types of flowers, we measured 26 morphological variables. Our results indicate that: (i) pistillate flowers from male trees carry healthy-looking ovules and are morphologically similar, although smaller than, pistillate flowers on female plants; (ii) staminate flowers have a rudimentary, non-functional pistil and are the only flowers capable of producing nectar; and (iii) perfect flowers produce healthy-looking ovules and pollen, but have smaller ovaries than pistillate flowers and fewer anthers than staminate flowers, and do not produce nectar. The restriction of sexual variation to male trees is consistent with the evolutionary path of dioecy from hermaphrodite ancestors through the initial invasion of male-sterile plants and a subsequent gradual reduction in female fertility in cosexual individuals (gynodioecy pathway), but further work is needed to confirm this hypothesis.     

Pistillate and staminate flower development in dioecious Pistacia vera (Anacardiaceae)

The development of staminate and pistillate flowers in the dioecious tree species Pistacia vera L. (Anacardiaceae) was studied by scanning electron microscopy with the objective of determining organogenetic patterns and phenology of floral differentiation. Flower primordia are initiated similarly in trees of both sexes. Stamen and carpel primordia are initiated in both male and female flowers, and the phenology of organ initiation is essentially identical for flowers of both sexes. Vestigial stamen primordia arise at the flanks of pistillate flower apices at the same time functional stamens are initiated in the staminate flowers. Similarly, a vestigial carpel is initiated in staminate flowers at the same time the primary, functional carpel is initiated in pistillate flower primordia. Differences between the two sexes become apparent early in development as, in both cases, development of organs of the opposite sex becomes arrested at the primordial stage. Male flowers produce between four and six mature functional stamens and female flowers produce a gynoecium with one functional and two sterile carpels.     

Thrips pollination of the dioecious ant plant Macaranga hullettii (Euphorbiaceae) in Southeast Asia

Discussion about thrips (Thysanoptera) as main pollinators has been controversial in the past because thrips do not fit the preconception of an effective pollinator. In this study, we present evidence for thrips pollination in the dioecious pioneer tree genus Macaranga (Euphorbiaceae). Macaranga hullettii is pollinated predominantly by one thrips species, Neoheegeria sp. (Phlaeothripidae, Thysanoptera). As a reward for pollinators, the protective floral bracteoles function as breeding sites for thrips and trichomal nectaries on the adaxial surface of the floral bracteoles provide alimentation. Flowering phenology of both staminate and pistillate trees was highly synchronized within 3-4 wk periods. In contrast to pistillate trees, staminate trees start to breed the thrips inside the developing inflorescences ～2 wk before anthesis. Breeding of Neoheegeria sp. in the laboratory indicates that the thrips development is completed within ～17 d. Thus, staminate trees offer breeding sites for one thrips generation until the onset of pollen presentation. Intraspecific pollen transfer by thrips was proved by pollen loads of thrips taken from receptive pistillate inflorescences of M. hullettii. Bagging experiments of different mesh sizes showed that seed set reached almost the level of open-pollinated flowers when exclusively tiny insects like thrips were able to enter the net bags, but no apomictic seed set occurred when no insect access was given to the flowers.     

Pollination Biology of Distylous Rubiaceae in the Atlantic Rain Forest, SE Brazil

Abstract: Data on pollination biology constitute important clues for the comprehension of pollen flow and genetic differentiation in plant populations. Pollinator type, availability and behaviour may modify morphological and mating patterns in populations of typically distylous species. This study investigates the pollination biology of four distylous species of Rubiaceae in the Atlantic rain forest, SE Brazil. Data on flowering phenology, floral lifespan, stigmatic receptivity, pollen availability, nectar volume and concentration, and pollinator activity were collected. The species studied flower sequentially throughout the wet season, and produce terminal inflorescences which bear small, tubular, diurnal, nectariferous flowers. Despite these similarities, some of the species studied are pollinated by different groups of pollinators, probably due to their distribution, availability of flowers and corolla length. On the other hand, pollinator specificity does not seem to be so important for distylous species. Long mouthparts, like those of most of the recorded pollinators, may reach lower sexual organs and, together with the self- and intramorph-incompatibilities observed, be sufficient to perform legitimate pollination and maintain levels of intermorph mating.     

Adaptive significances of sexual system in andromonoecious Capparis spinosa (Capparaceae)

The sexual system of the perennial shrub Capparis spinosa L. (Capparaceae), which is distributed in arid deserts of northern Xinjiang, was investigated. The main results can be summarized as follows. (1) The species is andromonoecious, i.e. individuals possess both male and perfect flowers. Stamens of two floral morphs are normal and can be divided into long and short ones in each flower. The perfect flowers have well developed pistils, but male flowers have aborted pistils and only function as males. (2) There were very significant differ-ences among the populations in daily ratio of the two floral morphs, number of long and short stamens of perfect flowers and length of filaments and anthers of short stamens in male flowers (P<0.01), but no significant differ-ences occurred in biomass of floral organ (P>0.05). (3) Anthesis was nocturnal and lasted 15–16 h. Both male and perfect flowers opened about 18:00 at dusk. The number of two floral morphs produced on each individual was indeterminate every day during flowering, which would make the individual temporally androdioecious. However, the total number of male flowers was more than that of perfect flowers within the population every day. (4) The P/O ratios of perfect flowers in three populations were 1.57×104, 1.65×104 and 1.71×104. There was no significant difference in pollen numbers (P>0.05) within population between male and perfect flowers or between long and short stamens, and also no significant difference among the populations in pollen numbers of male and perfect flowers, and in ovule numbers and P/O ratios of perfect flowers (P>0.05). (5) Dynamic curves of pollen viability of long and short stamens in the two floral morphs were similar in three populations. Pollen longevity of both long and short stamens was about 18–20 h, and duration of stigmatic receptivity was about 16–18 h. (6) Floral visitors were hymenopterous and lepidopterous insects. There was a total of seven species of floral visitors in the three populations. The activities of them were greatly affected by the climate of the desert environment. (7) Perfect flowers were not apomictic and could produce fruits after self-pollination (autogamy and geitonogamy) and cross-pollination; thus they had a mixed mating system. These results suggest that the sexual system of andro-monoecy may reflect an environment-heredity interaction in C. spinosa. The male flowers increase the number of pollen grains, the P/O ratio and enhance male fitness of individuals. They also increase the floral display and consequently effect cross pollination by attracting more pollinators that bring cross-pollen to the stigmas of per-fect flowers, thus increase outcrossing rate and female fitness of individuals, which insure reproductive success of C. spinosa in the extreme desert environment.     

Absence of pollinators and apomictic fruit production in an Atlantic rainforest population of Cymbopetalum brasiliense (Annonaceae)

Breeding system and pollination biology of an isolated population of Cymbopetalum brasiliense (Annonaceae), a large-flowered understory tree, was studied during two consecutive flowering seasons in October/November 2007 and September/October 2008. Flowers were morphologically comparable to other Cymbopetalum species and generally to other Annonaceae taxa adapted to pollination by large beetles, such as Annona spp. Flowers were thermogenic and emitted a strong scent during the pistillate and staminate flowering phases. Heating of the floral chamber was most intense during the pistillate phase between 1800 and 1900?h (up to 6°C difference to air temperature). Floral scent consisted almost entirely of p-methylanisole. None of the 133 flowers examined during two seasons received any visitation by dynastid beetles or other potential pollinators. Lycaenid larvae (Oenomaus ortygnus) attacked buds and flowers in both flowering seasons and destroyed about 20% of all buds in the 2007 season. Fruit set was high (72% of nonpredated flowers in 2007), despite the absence of pollinators. Field experiments showed that agamospermic reproduction rather than self-pollination was most likely responsible for fruit production. The study reports a probable case of apomixis, which would be the first in the Annonaceae family.     

The developmental floral morphology of Montrichardia arborescens (Araceae) revisited

The spadix of Montrichardia arborescens contains unisexual flowers without a perianth. The pistillate flowers are located in the basal portion of the inflorescence, and the staminate flowers are located in the apical portion. There is a narrow :zone between male flowers and female flowers consisting of atypical flowers. The portion of the atypical flowers facing the staminate zone exhibits staminate characters (stamens), and the portion facing the pistillate zone has an aborted gynoecium. The floral development of Montrichurdia is compared with that of Philodendron and a new interpretation of the morphology of atypical flowers of Montrichardia is proposed. Ontogenetic evidence supports relationships with Philodendron rather than Cercestis. 2001 The Linnean Society of London