Early Cretaceous floral structures and in situ tricolpate‐striate pollen: New early eudicots from Portugal

Five new taxa with affinities to extant lineages that diverged at an early stage from the main line of eudicot evolution are established from the Early Cretaceous (late Aptian or early Albian) Vale de Agua locality, Portugal. Staminate flowers of Lusistemon striatus and pistillate flowers of Lusicarpus planatus are unisexual without rudiments of the opposite gender. They are linked by the association of an unusual pollen type found in situ in the stamens and adhering to the stigmatic surface. The staminate flower, Lusistemon striatus, is composed of six stamens subtended by small perianth parts. The arrangement of the stamens is difficult to ascertain, but their variable sizes suggests a spiral arrangement. Pollen found in situ is tricolpate and striate with densely‐spaced, sparsely diverging and anastomosing muri that are aligned more or less parallel to the polar axis. The muri have a conspicuous supratectal ornamentation of fine transverse ridges. The granular infratectal layer forms an indistinct internal reticulum. The foot layer is thin. Pollen is closely similar to dispersed grains from the Aptian of Egypt described as STRIOTRI‐SEGMUR. It also resembles pollen of the dispersed pollen genus Rutihesperipites, as well as some dispersed pollen assigned to Striatopollis. Pistillate flowers of Lusicarpus planatus consist of a bicarpellate, syncarpous gynoecium borne on a short stalk. The styles are bent outwards and expose the double‐crested stigmatic regions on their ventral sides. The only organ preserved besides the gynoecium is a lateral scale‐like organ at the base of the stalk. Pollen of the same type found in Lusistemon striatus occurs on the stigmatic surface of the carpels. Comparisons with extant taxa demonstrate that Lusistemon and Lusicarpus share many characters with early diverging groups of eudicots, in particular Buxaceae. In addition to the Lusistemon‐Lusicarpus flowers, the Vale de Agua samples also contain three other pistillate reproductive structures that may be related to early diverging lineages of eudicots. Silucarpus camptostylus has a bicarpellate and syncarpous gynoecium with two styles; Valecarpus petiolatus and Aguacarpus hirsutus have tricarpellate gynoecia that are distinguished from each other in the shape and extension of the stigma as well as other details.     

Flower structure and potential bisexuality in Freycinetia reineckei (Pandanaceae), a species of the Samoa Islands

In Freycinetia reineckei the staminate flower (on the staminate spikes) comprises 3 or 4 (sometimes 2) stamens and a pistillode with 2 (sometimes 4) carpellodes, and the pistillate flower (on the pistillate spikes) is formed of a pistil with 2 (sometimes 4) carpels and of 3 or 4 (sometimes 2) staminodes. This perfect floral homology, also observed in all the other species that were studied with both pistillate and staminate material, strongly suggests that the flower of Freycinetia is basically and potentially bisexual, and may explain the occasional sexual lability and bisexuality of that flower (occurrence of both pistillate and staminate inflorescences, and/or of bisexual inflorescences with bisexual flowers and/or unisexual flowers, on the same individuals) in some species, and also the frequent occurrence of bisexual spikes in this species. These may be partitioned into pistillate, staminate, mixed and sterile zones. In the pistillate zones the flowers have the same aspect and structure as the pistillate flowers. In the staminate zones the flowers generally comprise 3 or 4 (sometimes 2) stamens and a ‘semi-pistil’ some have both stamens and staminodes. The semi-pistils are intermediate between pistils and pistillodes in length, aspect and structure, but always have placentas and ovules. In the mixed zones the flowers are generally formed of a pistil and 3 or 4 (sometimes 2) stamens, and are therefore true hermaphrodite flowers; some have both stamens and staminodes. In the sterile zones the flowers comprise a semi-pistil and 3 or 4 (sometimes 2) staminodes. The staminodes are anatomically very similar to the stamens, especially in the staminate, mixed, and sterile zones, in which they exhibit a wide range of variation in length, aspect and structure. The perfect floral homology as generic character on one hand, and the occasional bisexuality both with and without bisexual flowers and other aspects of sex expression (e.g. occurrence of both pistillate and staminate shoots on the same individuals) in some species on the other hand, seem to indicate that Freycinetia is a basically monoecious, sex changing genus.     

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.     

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.

     

Reproductive Biology of the Monoecious Understory Palm Prestoea schultzeana in Amazonian Ecuador1

Prestoea schultzeana is a monoecious, protandrous palm in the forest understory of Amazonian Ecuador. We studied its leaf production, population density, sexual expression, phenology, pollination, and the specificity of the floral visitors. On average, 1.4 leaves and 0.9 inflorescences are produced per individual per year. The number of staminate flowers per inflorescence is relatively constant compared with the number of pistillate flowers which varies greatly. Flowering occurs in staminate and pistillate phases of approximately 19 and 0–7 days duration, respectively. Flowers open in the morning, and staminate flowers abscise in the afternoon of the same day whereas pistillate flowers last for two days. Flowers are whitish-yellow with a sweet odor and produce nectar. They were visited by Coleoptera (Chrysomelidae, Curculionidae, Nitidulidae, Ptiliidae, Staphylinidae), Hemiptera, Diptera (Drosophilidae, Syrphidae, Ceratopogonidae), Lepidoptera (Nymphalidae), and Hymenoptera (Formicidae, Halictidae). All examined individuals of the syrphid fly Copestylum sp. visiting pistillate flowers carried 100–500 grains of P. schultzeana pollen. Pollen occurred on all body parts, but especially on the legs, and this makes Copestylum sp. the most important pollinator. Most floral visitors were also frequent on the flowers of co-occurring plant species; notably the palm Hyospathe elegans shared most visitor species with P. schultzeana.     

The role of growth regulators in the differentiation of flowers and inflorescences

Growth regulators participate in the differentiation of floral parts, determining the developmental path of the respective type of inflorescence. The effect depends on the expression of the peculiarities of floral part differentiation, the recognition of the character of endogenous substances in certain stages and the choice of the suitable regulator for application. In the primitive flower ofPapaver petals and stamens are formed from the peripheral meristem with a lower content of auxins and a higher level of gibberellic substances. The pistil arises later from central tissues with a higher level of auxins and inhibitory substances. The stamens are more sensitive to the higher level of auxin substances, and by a suitable application of GA₃ and BAP they can be transformed into petals; in this way double flower forms arise. In the differentiation of floral parts ofCampanula, Rosa andMelandrium similar regularities assert themselves in time successions, but in another spatial arrangement. Sex differentiation of diclinous flowers ofMelandrium is based on differences in heterochromosomes XY and XX. The rise of the zygomorphic flower ofVeronica is accompanied by a different distribution of endogenous substances which affect the development of petals, stamens and the pistil. The differentiation of flowers in the racemose inflorescence occurs in the acropetal succession, and lateral primordia inCampanula develop into actinomorphic regular flowers, whereas inDigitalis they are zygomorphic and only the terminal flower is peloric. In the initial phases the staminate tassel and the pistillate ear in maize are identical. Earlier differentiation of the terminal pistillate tassel is connected with a higher level of gibberellins and the later development of the lateral pistillate ear is accompanied by the increase in auxin-like substances and inhibitions. Similar correlations were found in the development of staminate catkins and the differentiation of pistillate flowers in terminal buds ofJuglans regia. By the application of auxin-like substances it is possible to achieve the transformation of primordia of the staminate tassel into the pistillate ear in maize or to regulate the number of staminate catkins and pistillate flowers on twigs of the walnut tree. In the capitulum of the sunflower differences arise between peripheral pistillate ray flowers and hermaphrodite tubular ones. By applying GA₃ and BAP the number of ray flowers is increased. If the normal course of inflorescence differentiation is affected with a suitable type of regulator, a range of floral abnormalities appears which permit to assess the intervention in different developmental stages and the reaction of the primordium to the applied type of regulator. Abnormalities also suggest some phylogenetic correlations.     

单性木兰（木兰科）花的形态发生

报道了单性木兰(Kmeria septentrionalis Dandy)花的形态发生过程。发现过去一直被认为是雌花条状披针形的“内轮花被片”,实际为退化雄蕊,它形态发生的时间与位置均与雄花的雄蕊相同,在成熟结构中仍可见药室残迹,说明单性木兰的雌性花是由两性花退化而来。通过与K. duperreana(Pierre) Dandy和Magnolia thailandica Noot. &; Chalermglin雌花的比较,发现它们雌花的形态相同,从而得知人们长期以来对此3种植物雌花的认识有误,原一直认为的“内轮花被片”实为退化雄蕊。     

Gender variation in Actinidia deliciosa,the kiwifruit

Summary Flower and fruit characters were measured in ten female, five male and five fruiting male selections of A. deliciosa var deliciosa (A. Chev) Liang and Ferguson. Flowers from female vines had functional pistils, which contained many ovules. Stamens appeared to be fully developed but produced only empty pollen grains. Flowers from male vines had functional stamens that produced high percentages of pollen grains with stainable cytoplasmic contents. Pistils did not contain ovules and were generally small with vestigial styles. Fruiting male vines had both staminate and bisexual flowers. Staminate flowers were similar to those found on strictly male vines. Bisexual flowers produced ovules and stainable pollen. Pistils were smaller than in pistillate flowers. Although the three flower sexes differed in style length, ovary dimensions and ovules per carpel, staminate and bisexual flowers were similar in number of flowers per inflorescence, stamen filament length, pollen stainability, inflorescence rachis length and carpel number, and differed from pistillate flowers in these characters. The three flower sexes had similar sepal and petal numbers. The fruit of fruiting males were considerably smaller than those of females. Low ovule number appears to be the major factor limiting fruit size in the fruiting males studied. Prospects for developing hermaphroditic kiwifruit cultivars through breeding are discussed.     

Paisia,an Early Cretaceous eudicot angiosperm flower with pantoporate pollen from Portugal

A new fossil angiosperm, Paisia pantoporata, is described from the Early Cretaceous Catefica mesofossil flora, Portugal, based on coalified floral buds, flowers and isolated floral structures. The flowers are actinomorphic and structurally bisexual with a single whorl of five fleshy tepals, a single whorl of five stamens and a single whorl of five carpels. Tepals, stamens and carpels are opposite, arranged on the same radii and tepals are involute at the base clasping the stamens. Stamens have a massive filament that grades without a joint into the anther. The anthers are dithecate and tetrasporangiate with extensive connective tissue between the tiny pollen sacs. Pollen grains are pantoporate and spiny. The carpels are free, apparently plicate, with many ovules borne in two rows along the ventral margins. Paisia pantoporata is the oldest known flower with pantoporate pollen. Similar pantoporate pollen was also recognised in the associated dispersed palynoflora. Paisia is interpreted as a possibly insect pollinated, herbaceous plant with low pollen production and low dispersal potential of the pollen. The systematic position of Paisia is uncertain and Paisia pantoporata most likely belongs to an extinct lineage. Pantoporate pollen occurs scattered among all major groups of angiosperms and a close match to the fossils has not been identified. The pentamerous floral organisation together with structure of stamen, pollen and carpel suggests a phylogenetic position close to the early diverging eudicot lineages, probably in the Ranunculales.     

THE INTRAFLORAL PHENOLOGY OF STREPTANTHUS TORTUOSUS (BRASSICACEAE)

The sequence of floral events during anthesis was examined in Streptanthus tortuosus to determine the relationship between the male and female floral phases. The flowers are strongly protandrous. In the staminate phase, the anthers mature sequentially over a 3–4-day period. Because pollinators quickly remove pollen from the anthers, sequential anther maturation prolongs the male phase relative to what it would be if anthers did not mature sequentially. Pollen applied to the stigma during the staminate phase does not adhere readily and does not germinate. The length of the pistillate phase depends on pollinator activity, as pollination accelerates the abscission of floral parts. Unpollinated flowers remain pistillate for 3–4 days, during which time stigmatic receptivity declines gradually. In the field, 72% to 80% of flowers are staminate at any time, indicating that the staminate phase is three times longer than the pistillate phase when pollinators have access to the flowers. The consequences of the relative length of the floral phases and the schedule of stigmatic receptivity are discussed in terms of outcrossing mechanism, floral longevity, and sexual selection models.     

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.     

Canrightiopsis,a new Early Cretaceous fossil with Clavatipollenites-type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae

Canrightiopsis with three species (C. intermedia, C. crassitesta, C. dinisii) is described from the Early Cretaceous of Portugal based on small, one-seeded berries. The fruits are derived from bisexual flowers with three stamens borne on one side of the ovary. There are no traces of a perianth. Pollen is of the Clavatipollenites-type, monocolpate, semitectate, reticulate-columellate with heterobrochate reticulum and muri with beaded supratectal ornamentation. The ovary is unilocular with a single pendant, orthotropous and bitegmic ovule. The seed is endotestal. The endotesta consists of one layer of palisade-shaped crystal cells with fibrous infillings. The fruit wall has resin bodies or cavities from presumed ethereal oil cells sometimes seen as stomata-like structures on the fruit surface. A phylogenetic analysis resolves Canrightiopsis as a close relative of extant Chloranthaceae, particularly close to extant Chloranthus and Sarcandra. All three taxa share the one-sided position of the stamens on the ovary. An evolutionary sequence from fossil Canrightia to fossil Canrightiopsis and extant Chloranthus and Sarcandra is suggested by loss of perianth, reduction in number of ovules and stamens and displacement of stamens to one side of the ovary. Canrightiopsis also shares several critical features with extant Ascarina including monoaperturate pollen and beaded supratectal ornamentation of the pollen wall.     

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.     

The pollination biology and breeding system of Monarda fistulosa (Labiatae)

Summary Successful cross-pollination of Monarda fistulosa is the result of a complex interaction among flower opening, the pollen-bearing areas of the pollinators and/or their behavior, and the maturation of the stigmas. The flowers open continuously from 0800–2000 h providing a temporally predictable rich source of nectar and pollen. Recently opened flowers may reduce the ability of bees to discriminate between resource rich and poor patches and encourage systematic foraging within patches. The continuous opening of flowers coupled with protandry also results in some flowers of most capitula being in the staminate and others in the pistillate phase. Autogamy is highly unlikely due to strong protandry and the spatial separation of anthers and stigmas. Geitonogamy, at least that mediated by Bombus is unlikely because the pollen is spread over a relatively large area of the wings, which reduces the likelihood of a stigma contacting just deposited pollen. Because pollen is transferred from the much smaller coxal area of Anthophora and other bees that mistake the stigmas of early pistillate phase flowers for stamens some geitonogamy seems inevitable. However, the delayed receptivity of young stigmas to self-pollen decreases the likelihood of self-pollen germinating on such stigmas. Older stigmas are equally receptive to self- and cross-pollen and the number of pollen grains germinating and pollen tubes reaching the base of the style increases with flower age.     

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.     

Neither insects nor wind: ambophily in dioecious Chamaedorea palms (Arecaceae)

Pollination of Neotropical dioecious trees is commonly related to generalist insects. Similar data for non‐tree species with separated genders are inconclusive. Recent studies on pollination of dioecious Chamaedorea palms (Arecaceae) suggest that species are either insect‐ or wind‐pollinated. However, the wide variety of inflorescence and floral attributes within the genus suggests mixed pollination mode involving entomophily and anemophily. To evaluate this hypothesis, we studied the pollination of Chamaedorea costaricana, C. macrospadix, C. pinnatifrons and C. tepejilote in two montane forests in Costa Rica. A complementary morphological analysis of floral traits was carried out to distinguish species groups within the genus according to their most probable pollination mechanism. We conducted pollinator exclusion experiments, field observations on visitors to pistillate and staminate inflorescences, and trapped airborne pollen. A cluster analysis using 18 floral traits selected for their association with wind and insect pollination syndromes was carried out using 52 Chamaedorea species. Exclusion experiments showed that both wind and insects, mostly thrips (Thysanoptera), pollinated the studied species. Thrips used staminate inflorescences as brood sites and pollinated pistillate flowers by deception. Insects caught on pistillate inflorescences transported pollen, while traps proved that pollen is wind‐borne. Our empirical findings clearly suggest that pollination of dioecious Chamaedorea palms is likely to involve both insects and wind. A cluster analysis showed that the majority of studied species have a combination of floral traits that allow for both pollination modes. Our pollination experiments and morphological analysis both suggest that while some species may be completely entomophilous or anemophilous, ambophily might be a common condition within Chamaedorea. Our results propose a higher diversity of pollination mechanisms of Neotropical dioecious species than previously suggested.     

Erdtmanitheca portucalensis,a new pollen organ from the Early Cretaceous (Aptian–Albian) of Portugal with Eucommiidites-type pollen

A new lignitised, slightly compressed pollen organ, Erdtmanitheca portucalensis, with affinities to extinct Erdtmanithecales from the Early Cretaceous (Aptian–Albian) of Vale de Água (Lusitanian Basin, western Portugal), is described. The pollen organ is composed of loosely arranged microsporophylls radiating from a central core. The estimated number of microsporophylls is about 100–150. The microsporophylls are sessile and ellipsoidal to barrel-shaped with a flattened or slightly apically depression containing about ten narrow sporangia. The sporangia enclose abundant well-preserved pollen grains of Eucommiidites-type. Pollen grains found in situ are elliptical in equatorial outline, about 16.0–27.2 μm long and 11.9–16.4 μm wide. The main (distal) colpus is long with expanded rounded ends. It is flanked by two subsidiary colpi in an almost equatorial position. The surface of the pollen wall is psilate and occasionally punctate. The ektexine is composed of a distinct tectum, granular infratectal layer and a thin foot layer. The endexine is thick and laminar. The new Early Cretaceous Portuguese pollen-organ is similar in several respects to that of Erdtmanitheca texensis described from the Late Cretaceous of Texas, USA. The new fossil species further documents the importance of the Bennettitales-Erdtmanithecales-Gnetales group in the Early Cretaceous floras of Portugal extending the stratigraphic and geographical distribution of the genus with regard to systematic and phylogenetic significance of the Eucommiidites-producing plants that may have been co-occurring with the Early Cretaceous diversification of angiosperms. It is ascertained that perforate tectum occurs in pollen grains with a well-developed foot layer as well as in pollen grains in which a foot layer is poorly developed or lacking, and that pollen features do not support a separation of the Erdtmanithecales seeds and pollen organs.     

POLLINATION ECOLOGY OF A SEAGRASS,THALASSIA TESTUDINUM (HYDROCHARITACEAE), IN ST. CROIX

The reproductive ecology of a dioecious Caribbean seagrass, Thalassia testudinum was studied in St. Croix, U.S. Virgin Islands. Flowering in Thalassia coincided with spring tides with anthesis and anther dehiscence in staminate plants occurring at night. As predicted by search theory, pollen is dispersed in negatively buoyant rafts of pollen grains (that superficially resemble strings of frog spawn) which are bound by a slime of apparent thecal origin. Dispersal of pollen is submarine and approximately two-dimensional along the plane of the substratum. The stigmas on pistillate plants are linear, stiff, and densely papillate; they protrude from the mouth of the subtending leaf. Floral ratios of staminate to pistillate flowers were 60 to 1 with an average distance of 4.0 cm from pistillate flower to the closest staminate flowers. Populations included a high proportion of short shoots with numerous (> 100) leaf scars indicating ages for these axes of up to 3–4 years. Nonflowering shoots can be sexed and frequency of flowering can be assessed from scars of old inflorescences. Preliminary results suggest that there is no regular pattern of flowering, since the interval between flowering events is variable.     

LOSS OF FLORAL ORGANS IN ATELEIA (LEGUMINOSAE: PAPILIONOIDEAE: SOPHOREAE)

Ateleia herbert-smithii is unique among legumes in being a wind-pollinated tree; carpellate and staminate flowers are restricted to different trees. Development of the two floral morphs, however, is essentially the same except for smaller carpels in functionally staminate flowers and failure of pollen formation in the anthers of functionally carpellate flowers. The floral development of Ateleia herbert-smithii is highly atypical among papilionoids and the tribe Sophoreae. Order of organ initiation is: sepals, solitary petal, carpel, and lastly all stamens in erratic order. Sepal order is unidirectional from the abaxial side, the normal pattern for papilionoids. Only one petal, the vexillum or standard, is initiated. Subsequent initiation is completely different from the usual unidirectional pattern of most papilionoids. A meristem ring forms, delimiting the solitary carpel centrally. Ten stamen primordia are initiated on the meristem ring, first laterally, then adaxially, and lastly abaxially. There is a tendency for antesepalous stamens to form before the antepetalous ones. The loss of four of the five petals is thought to alter drastically the subsequent organogeny as to position of organs and their order of initiation. Carpel initiation in Ateleia is precocious, but not uniquely so among legumes.     

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.