Comments on flower evolution and beetle pollination in the generaAnnona andRollinia (Annonaceae)

Dynastid scarab beetle pollination appears basic within the genusAnnona. Those species ofAnnona which are more morphologically derived, as well as allRollinia spp. possess reduced floral chambers and attract small beetles likeNitidulidae orStaphylinidae. Pollination of the primitive species ofAnnona byDynastinae would imply that the genus had not evolved before the Tertiary. The fossil record is in congruence with this hypothesis. Once again it is stressed that the cantharophilous syndrome, as it is found in theAnnonaceae, Magnoliaceae, Eupomatiaceae andCalycanthaceae, with beetles being exclusive pollinators, is a secondary and derived condition and obviously different from the expected basic entomophily of the original angiosperms.     

Beetle pollination of Shorea parvifolia (section Mutica, Dipterocarpaceae) in a general flowering period in Sarawak, Malaysia

Pollination ecology of an emergent tree species, Shorea (section Mutica) parvifolia (Dipterocarpaceae), was studied using the canopy observation system in a lowland dipterocarp forest in Sarawak, Malaysia, during a general flowering period in 1996. Although the species has been reported to be pollinated by thrips in Peninsular Malaysia, our observations of flower visitors and pollination experiments indicated that beetles (Chrysomelidae and Curculionidae, Coleoptera) contributed to pollination of S. parvifolia in Sarawak. Beetles accounted for 74% of the flower visitors collected by net-sweeping, and 30% of the beetles carried pollen, while thrips accounted for 16% of the visitors, and 12% of the thrips carried pollen. The apical parts of the petals and pollen served as a reward for the beetles. Thrips stayed inside the flower almost continuously after arrival, and movements among flowers were rare. Fruit set was significantly increased by introduction of beetles to bagged flowers, but not by introduction of thrips. Hand-pollination experiments and comparison of fruit set in untreated, bagged, and open flowers suggested that S. parvifolia was mainly outbreeding.     

Flowering mechanisms,pollination strategies and floral scent analyses of syntopically co‐flowering Homalomena spp. (Araceae) on Borneo

In this study, the flowering mechanisms and pollination strategies of seven species of the highly diverse genus Homalomena (Araceae) were investigated in native populations of West Sarawak, Borneo. The floral scent compositions were also recorded for six of these species. The selected taxa belong to three out of four complexes of the section Cyrtocladon (Hanneae, Giamensis and Borneensis). The species belonging to the Hanneae complex exhibited longer anthesis (53–62 h) than those of the Giamensis and Borneensis complexes (ca. 30 h). Species belonging to the Hanneae complex underwent two floral scent emission events in consecutive days, during the pistillate and staminate phases of anthesis. In species belonging to the Giamensis and Borneensis complexes, floral scent emission was only evident to the human nose during the pistillate phase. A total of 33 volatile organic compounds (VOCs) were detected in floral scent analyses of species belonging to the Hanneae complex, whereas 26 VOCs were found in samples of those belonging to the Giamensis complex. The floral scent blends contained uncommon compounds in high concentration, which could ensure pollinator discrimination. Our observations indicate that scarab beetles (Parastasia gestroi and P. nigripennis; Scarabaeidae, Rutelinae) are the pollinators of the investigated species of Homalomena, with Chaloenus schawalleri (Chrysomelidae, Galeuricinae) acting as a secondary pollinator. The pollinators utilise the inflorescence for food, mating opportunities and safe mating arena as rewards. Flower‐breeding flies (Colocasiomyia nigricauda and C. aff. heterodonta; Diptera, Drosophilidae) and terrestrial hydrophilid beetles (Cycreon sp.; Coleoptera, Hydrophilidae) were also frequently recovered from inflorescences belonging to all studied species (except H. velutipedunculata), but they probably do not act as efficient pollinators. Future studies should investigate the post‐mating isolating barriers among syntopically co‐flowering Homalomena sharing the same visiting insects.     

A new species of Annona (Annonaceae) from the northeastern Guayana Shield

Annona prevostiae, a new species from the northeastern margin of the Guayana Shield, is described and illustrated.     

Beetle pollination of peacock moraeas (Iridaceae) in South Africa

The peacock moraeas are South African geophytes withIris-like flowers that are characterized by broad outer tepals with iridescent spots. Three of the seven species were studied and found to be exclusively visited and pollinated by beetles. The primary pollinators were hopliine (Scarabaeidae) beetles in the generaPeritrichia, Monochelus, Anisonyx andAnisochelus. These beetles visit flowers for nectar, pollen and mating.Peritrichia rufotibialis was the primary pollinator ofMoraea villosa, P. abdominalis was the primary pollinator ofM. tulbaghensis, whileMonochelus sp. nov. was the primary pollinator of the third species,M. neopavonia. High levels of fruit set attest to the effectiveness of these beetles as pollinators. Average fruit set ranged from 83.5 ± 34.8% (M. villosa) to 97.2 ± 11.6% (M. tulbaghensis). Average seed set in the various populations was more variable, but was also quite high, ranging from 77.3 ± 40.8 (M. villosa) to 342.2 ± 115.0 (M. neopavonia) seeds per capsule. Although hopliine beetles are abundant and diverse in southern Africa and commonly visit flowers, this is the first well-documented report of pollination by these beetles.     

Reproductive biology ofPolyalthia littoralis (Annonaceae)

Flowering phenology of some annonaceous trees and reproductive biology ofPolyalthia littoralis (Annonaceae) were studied. The trees showed various types of flowering phenology within the family. Among them,P. littoralis had hermaphroditic and protogynous flowers, and exhibited continuous flowering throughout the year. Bagged flowers set fruits and seeds comparable to the control. The observations of meiotic stages and the results of castrated tests indicated no possibility of apomictic reproduction. The abscised anthers attached to torus by the extended spiral thickenings, and accomplished self-pollination. Outbreeding possibly occurs, but the following self-pollination guarantees seed set.     

The pollination ofStenorrhynchos lanceolatus (Aublet) L. C. Rich. (Orchidaceae: Spiranthinae) by hummingbirds in southeastern Brazil

Hummingbird pollination is documented for a natural population ofStenorrhynchos lanceolatus Aublet. L. C. Rich. occurring in Rio de Janeiro State, southeastern Brazil. At the study site the plants are pollinated byPhaethornis eurynome (Phaethorninae),Thalurania glaucopis (females only) andLeucochloris albicollis (Trochilinae). The plants offer nectar as a reward and the pollinaria become stuck to the surface of the hummingbird's bill while it is probing the flowers. The orchid population received a few (0–4) hummingbird visits per day, with about 83% of the flowers being pollinated. In spite of the low frequency of visits, the granular structure of the pollinarium plus the behaviour of the most frequent pollinators, which tend to visit all the fresh-looking flowers of each inflorescence, a very high fruiting success was promoted. Experimental evidence suggests that the pollinaria may remain up to 6.30 hours on the hummingbird's bill, enhancing the chances of cross-pollination and long-distance pollen flow.     

Sporoderm characters ofTetrameranthus duckei (Annonaceae) and their systematic implications

Ultrastructural and micromorphological investigations of the sporoderm ofTetrameranthus duckei pollen confirm the interpretation of this genus as being isolated within theAnnonaceae.     

Seed development and function inArtabotrys hexapetalus (Annonaceae)

Until now seeds ofAnnonaceae were characterized as mesotestal only. The seed ofArtabotrys hexapetalus, however, is meso- and endotestal. An outer mechanical layer which surrounds the seed as a lignified fibrous tissue is derived from the mesotesta. A complex inner mechanical layer develops from a partially multi-layered endotesta built up by crystal-containing stone cells. The multi-layered endotesta forms a prominent seed plug in the micropylar region which is prolonged along both sides of the perichalaza as inner walls. The endotesta is one-layered on the sides of the seed and participates in rumination. In addition the endotesta may serve for deposition of end-products of metabolism. The complex growing process of the perichalaza and its surrounding tissues is described in detail. The perichalazal pad of tanniferous cells forms an U-shaped septum, in conjunction with the endotesta, dividing the seed into two chambers. During seed development it functions as transmitter of nutrients from the outer chamber filled with starch grains to the nucellus, endosperm and embryo contained in the inner one. During germination this pad probably serves for the uptake of water. — At the initial phases of germination the seed dehisces into two valves along the raphal and antiraphal side. Later on an additional parenchymatous operculum covering the seed apex disintegrates and the endotestal plug fixed by its two prolongations splits along a preformed fracture line into two parts to release the seedling. Rudiments of an aril are recognizable in young seeds only. — The data obtained fromArtabotrys hexapetalus are discussed and compared with published information on seeds in other annonaceous taxa. For systematic considerations the necessity to define the origin of the annonaceous seed plug from one or the other integument is emphasized as it may prove to be an important differential character withinAnnonaceae.     

Movement patterns of Cetonia beetles (Scarabaeidae) among flowering Viburnum opulus (Caprifoliaceae)

The movement patterns of flower-visiting Cetonia (Coleoptera, Scarabaeidae) were studied in central Sweden over 4 years, providing the first quantitative study of beetle pollination behaviour conducted in a temperate zone. The beetles were marked individually and tracked throughout their visits to Viburnum opulus L. (Caprifoliaceae), a partly beetle-pollinated shrub displaying large umbel-like, creamy-white blossoms. Beetle abundance differed greatly between study years. Of marked beetles observed on V. opulus flowers, an average of 26% returned each year. The beetles performed frequent inter-plant flights, and showed fidelity to particular V. opulus individuals at the study site. Furthermore, they preferred V. opulus to other plants flowering concomitantly and showed considerable constancy in this habit. Movements were mostly between individuals from unshaded locations with high inflorescence and flower number. Normally, the beetles flew on average about 4 times the horizontal nearest-neighbour distance between flowering V. opulus (c. 18 m). From these observations it is concluded that Cetonia beetles are powerful long-distance dispersal agents for V. opulus pollen, perhaps leading to an enhanced gene flow not possible with the plant's other pollinators.     

The pollination syndrome ofDeplanchea tetraphylla (Bignoniaceae)

The reproductive structures ofDeplanchea tetraphylla (Bignoniaceae) exhibit a significant number of unusual features: inflorescence with an apical platform; flowers yellow, short-tubed, strongly zygomorphic; mouth closed through lateral compression; stamens and style long-exserted, erect or slightly reclined; nectar dark brown, exposed in the spoon-shaped lowermost corolla lobe and apparently acting also as a visual cue. These features suggest a highly elaborate syndrome for bird pollination: the birds (probably lorikeets) perch on the inflorescence platform and bend downwards to take up the exposed nectar, thus touching the exserted anthers and stigmas with the throat or breast. The likely evolution of this syndrome by additive steps, effecting a change from head up to head down position of the pollinator, can be traced from the floral structure of the remaining four species of the genus.     

Octad pollen formation in Cymbopetalum (Annonaceae): the binding mechanism

Our recent study of tetrad pollen formation in Annona (Annonaceae) revealed that after meiosis the callose-cellulose envelope forms a special conjugation with individual microspores and the forthcoming callose digestion is incomplete. The undigested part forms a central binder holding the four microspores of the tetrad together. This process causes the microspores to rotate 180 degrees. In this paper we describe pollen formation in another annonaceous genus, Cymbopetalum, in which the pollen is shed in octads, through use of light microscopy, epifluorescence microscopy, and TEM. In Cymbopetalum, two meiocytes, connected by abundant cytomictic channels, are produced in each sporangium. Octad pollen formation in Cymbopetalum is shown to be comparable to the synchronized formation of two connected Annona tetrads, which then integrate into a single octad. Unique features of Annona polyad formation, e.g. special binding between the callose-cellulose envelopes and microspores, incomplete callose digestion, and microspore rotation, also occur in Cymbopetalum. In addition, formation of the Cymbopetalum octad involves development of a cushion-like structure that binds the distal pronexine of all eight microspores, and there is the production of intine protrusions. The evolutionary origin of the callose-cellulose binding mechanism within the family is discussed.     

Hydrophilous pollination and reproductive morphology in the seagrassPhyllospadix scouleri (Zosteraceae)

In the seagrassPhyllospadix scouleri (Zosteraceae) floral aggregates (spadices) occur on short lateral axes produced by subordinate vegetative shoots. The filamentous pollen is both dispersed on the surface of the sea as well as below the surface. In surface pollination, snowflake-like search vehicles (pollen rafts) float and collide with any rigid female inflorescences that emerge from the water surface. In submarine pollination, collinear bundles of pollen grains are dispersed. Analysis of seed set indicates ovule position within the inflorescence to affect likelihood of fertilization in submarine, but not surface-pollinated inflorescences. Agamospermy appears to be unlikely, but the strongly female-biased shoot sex ratio remains to be explained.     

Size-dependent pollination efficiency in Anchusa officinalis (Boraginaceae): causes and consequences

Summary Bumblebees foraging on the self-incompatible Anchusa officinalis fly between near neighbour plants and between near neighbour inflorescences within plants. Although many-flowered plants attracted most bumblebees these plants received fewer visits on a per flower basis than smaller plants, and each bumblebee visited a smaller proportion of the flowers. The calculated effective visitation rate per flower was highest on plants of an intermediate size. If pollen-carryover was assumed to be limited the most efficient plant was predicted to be smaller since the proportion of fertilized flowers per bumblebee visit is expected to decrease further on the largest plants in relation to the total flower number. These predictions were tested by measuring fruit-set in the field. The percentage fruit-set decreased with plant size at all sizes that were investigated. That the most efficient plant was small indicates that pollen-carryover was indeed limited. However, the low percentage fruit-set associated with large size did not present a serious problem since the total estimated seed production per plant still increased with size. Selection favoring smaller plants may be low or absent in Anchusa.     

Bird pollination in South African species ofSatyrium (Orchidaceae)

Field observations showed that three South African orchid species,Satyrium carneum, S. coriifolium andS. princeps, are pollinated by sunbirds. Foraging sunbirds insert their bills into the labellum chamber of the flowers and suck nectar from the labellum spurs with their tongues. The column overarches the entrance to the labellum and pollinaria become affixed to the upper mandible of the bill. Birds often attempt to remove pollinaria by rubbing their bills against a branch, but are mostly unsuccessful due to the large plate-like viscidia which attach the pollinaria very firmly to the bill. Other modifications for bird pollination in theseSatyrium species may include the red, carmine or orange colour of the flowers, sturdy inflorescence stems used for perching and large amounts of dilute nectar in the spurs. EachSatyrium species was pollinated by several species of sunbird, and despite some differences in flowering time, occasional hybrids occur at a site whereS. carneum andS. coriifolium share the same sunbird pollinators.     

New and noteworthyHabenaria spp. (Orchidaceae) from New Guinea and adjacent islands

5 new taxa ofHabenaria, namelyH. bougainvillae, H. elongata R. Br. var.leptophylla, H. ensigera, H. rechingeri andH. trichoglossa, are described and illustrated, with reference to affinities to related Australian and Indo-malayan species. The occurrence in New Guinea of severalHabenaria spp. typical for a savanna-like vegetation, led to look more thoroughly at these taxa:H. elongata R. Br. andH. ochroleuca R. Br., considered so far to be endemic in Northern Australia, andH. khasiana Hook. f., hitherto only known from southeastern Asia.Studies in the subtribeHabenariinae Bentham (Orchidaceae), 2.—Part 1: Candollea34, 357 (1979).Dedicated to Hofrat Prof. DrKarl Heinz Rechinger on the occasion of his 80th birthday. — On April 30, in the year 1927, the author and his brotherOtto Renz metKarl Heinz Rechinger on a small steam-boat in a stormy Aegaen Sea, travelling from Piraeus to the Northern Sporades Islands:Karl Heinz with destination to Chelidromia, the author andO. Renz to Skopelos. By this lucky chance a lasting friendship began.     

Contrasting pollination ecology of Disepalum species (Annonaceae): evolutionary loss of the floral chamber and partial breakdown of protogyny associated with a shift in pollination system

Disepalum comprises two monophyletic sister subgenera, Enicosanthellum and Disepalum, with strikingly different floral morphologies: the former has two whorls of unfused sepals, forming a partially enclosed floral chamber, whereas the latter possesses a single whorl of congenitally fused petals and lacks a floral chamber. The pollination ecologies of representative species are reported, including assessments of floral phenology, pollinators and floral thermogenesis. Disepalum pulchrum (subgenus Enicosanthellum) has hermaphroditic flowers with a pollination chamber and is protogynous with prolonged anthesis; it is pollinated by nitidulid beetles and drosophilid flies. Disepalum anomalum (subgenus Disepalum) is also hermaphroditic with prolonged anthesis, but has incomplete protogyny due to overlapping pistillate and staminate phases; it is pollinated by meliponine bees, which are attracted by the pollen, but which are only able to transfer pollen to receptive stigmas during the overlap phase. Alternative evolutionary hypotheses are evaluated, including the possibility that the ancestor of the subgenus Disepalum lineage may have experienced a profound genetic mutation, possibly involving genes responsible for organ merism and fusion, resulting in the loss of the pollination chamber and hence favouring different floral visitors. The breakdown in protogyny required for effective pollination is likely to have had significant ramifications on population genetic diversity.     

Insect-induced wind pollination of the palm Chamaedorea pinnatifrons and pollination in the related Wendlandiella sp.

In Peruvian Amazonia pollination ecology of two palms, Chamaedorea pinnatifrons (Jacq.) Oerst. and Wendlandiella sp. was studied from October 1988 to January 1990. Both palms are dioecious understory species that form locally dense populations.In C. pinnatifrons both sexes flower synchronously during the dry season. Prior to anthesis, the pendulous male inflorescence is inhabited by numerous thrips (Thysanoptera) and Ptiliidae (Coleoptera). Staminate flowers open by a small basal slit between the petals. At anthesis pollen is shed and the movements of the insects inside the flowers trigger pollen release in small clouds. Thus, the powdery pollen becomes airborne and finally air currents act as a vector, carrying pollen to the inconspicuous female plants, which usually are not visited by insects. The term insect induced wind pollination is suggested for this pollination mode. Wendlandiella flowers during four months in the dry season. Male and female plants were not visited by insects. The dry condition of the pollen indicates that anemophily is the pollination mode in Wendlandiella. Fructification is rare but the plants show intense vegetative propagation.The significance of anemophily in the tropical lowland rain forest is discussed.     

A new species of Sinningia (Gesneriaceae) from northeastern Brazil

Sinningia nordestina is described and illustrated. Morphological and molecular characters are discussed. It is a new species endemic to northeastern Brazil and is distinguished from other members of the genus by its vestigial tubers, ascending pedicels with pendent flowers, small corollas, and deviating flowering period.