Pollination biology of the Galápagos endemic, Tournefortia rufo-sericea (Boraginaceae)

Tournefortia rufo-sericea is an endemic member of the Galápagos angiosperm flora. Although not uncommon within the archipelago, its status is presently listed as vulnerable and, as such, a complete knowledge of its pollination biology may prove useful in preventing its decline. Pollination experiments, flower-visitor observations, nectar and fluorescence studies, as well as pollen : ovule ratio and pollen size studies were included in this investigation. The small, white flowers of this species set fruit via open pollination (81%), autonomous self-pollination (80%), diurnal pollination (80%), and nocturnal pollination (85%). Ants are the most common visitor to the flowers of this species, primarily during the day, whereas beetles and moths make visits at night. A small amount of nectar is presumably produced, as moths are often seen probing the corollas. Experiments with fluorescent dust did not support interflower pollen movement. The pollen : ovule ratio was 4972 : 1, which suggests xenogamy, and the mean pollen size was 21 µm. Despite the pollen : ovule ratio, it appears that this species exhibits a breeding strategy of facultative autogamy, in which the majority of flowers set fruit via autonomous self-pollination, but limited outcrossing may occur when pollinators are available. Conservation efforts for T. rufo-sericea should focus on the control or eradication of exotic species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 21–31.     

Partial self‐incompatibility in the polyploid endemic species Scalesia affinis (Asteraceae) from the Galápagos: remnants of a self‐incompatibility system?

Different pollination treatments of capitula were used to examine the breeding system of individuals of the tetraploid endemic species Scalesia affinis from the Galápagos Islands. All types of crossings resulted in approximately 35 achenes per capitulum, but in actively and passively self-pollinated capitula these were mostly without embryos. Among self-pollinated individuals a large variation was found in the production of embryos. Some individuals failed completely to set filled achenes while others produced few or many. Additionally, we found a clear difference in female phenology of florets in self-pollinated capitula compared with florets in cross-pollinated capitula. Female florets in self-pollinated capitula remained receptive for longer time. These results suggest that Scalesia affinis is partly self-incompatible. Outcrossing is assured by the endemic carpenter bee, Xylocopa darwini , which proved to be an important pollinator of Scalesia affinis . Nevertheless, geitonogamy seemed to be considerable in the focal population, for which reason open pollinated capitula produced fewer achenes than controlled outcrossed capitula.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 93–101.     

How flower biology and breeding system affect the reproductive success of the narrow endemic Polygala vayredae Costa (Polygalaceae)

Polygala vayredae is a narrow endemic species from the oriental pre-Pyrenees. Despite its conservation status and rarity, no information is available on its reproductive biology. As the flower is the structure directly involved in pollinator attraction, its morphological and functional traits have major effects on the reproductive success of the plant. In this work, the flower biology and breeding system of P. vayredae were studied to evaluate how they affect the reproductive outcome in natural populations. Flower morphology, flower rewards, and male and female functioning throughout the lifespan of the flower were assessed. Pollination experiments, involving pollinator exclusion and pollen from different sources, were conducted, and the pollen ovule index was determined. Female fitness and the occurrence of pollen limitation were assessed in three natural populations over 2 years by observing the presence of pollen on the stigma, pollen tube development, and fruit production. Polygala vayredae flowers are elaborate and long-lived with nectar rewards. The floral traits are well adapted to xenogamy and entomophily, which are in accordance with the observed breeding system and auto-incompatibility system. No mechanism of reproductive assurance was observed and P. vayredae strictly depends on pollinators to set fruit. Low fruit production was observed in the studied populations, which was largely the result of scarce, unreliable, and/or inefficient pollinators and poor pollen quality. In addition, available resources may be a limiting factor. The reproductive strategy of P. vayredae prevents inbreeding depression by a self-incompatibility system, which in years of scarce pollinators is overcome by the plant habit.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 67–81.     

Pollination biology of a night‐flowering Galápagos endemic,Ipomoea habeliana (Convolvulaceae)

Ipomoea habeliana is an endemic, night‐flowering member of the Galápagos flora. Pollination experiments, flower‐visitor observations, nectar sampling, pollen transfer, and pollen to ovule ratio and pollen size studies were included in this project. The large, white flowers of this species set fruit via open pollination (55%), autonomous autogamy (51%), facilitated autogamy (91%), cross‐pollination (80%), diurnal open pollination (60%) and nocturnal open pollination (60%). Fruit set is pollen‐limited. Ants, beetles, crickets and hawk moths regularly visit the flowers. Ants are the most frequent visitors, but hawk moths are the only effective pollinators. Nectar is available throughout the night, but is most abundant early in the evening when hawk moth visits are most frequent. Experiments with fluorescent dust demonstrate intra‐ and inter‐plant pollen movement by hawk moths. Although this species is adapted for hawk moth pollination, it readily sets fruit via autonomous autogamy when no visits are made. Thus, it is concluded that it is facultatively xenogamous. Additional support for this conclusion is provided by the pollen to ovule ratio of 1407 and by the fact that the plants grow in a region that has few or no faithful pollinators. Conservation efforts for I. habeliana should include hand pollinations, which could significantly increase seed set. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 11–20.     

Reproductive biology of the invasive exotic shrub, Rhododendron ponticum L. (Ericaceae)

The reproductive biology of an exotic species will affect its ability to become naturalized and invasive in non-native habitats. Rhododendron ponticum is an ecologically damaging exotic weed in the British Isles, which spreads predominantly by seed. I investigated how inbreeding and outcrossing affect seed production and germination in a wild population of this species in Ireland. Experimental manipulations revealed low fruit and seed set when insects were excluded from flowers, suggesting that this species has limited capability for spontaneous autogamy. Hand-pollination treatments showed that, although flowers are self-compatible (with self and same plant pollen), higher levels of seed set occur following outcrossing (xenogamy). There was no significant difference in rate of germination of seeds from inbred or outcrossed treatments. The addition of xenogamous pollen to open flowers did not increase fruit or seed set, suggesting that flowers in this population are not pollen limited: native generalist pollinators, mainly bumblebees ( Bombus spp.), are providing an adequate pollinator service. This work demonstrates that outcrossing increases seed set and pollinators are required to facilitate this. Hence, generalist native pollinators can promote invasion by exotic plants. Native pollinators can clearly play an important part in alien species invasion.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 373–381.     

The bark and ambrosia beetles (Curculionidae, Scolytinae) of Cocos Island, Costa Rica and the role of mating systems in island zoogeography

Cocos Island is a small oceanic island midway between Costa Rica and the Galápagos Archipelago; about 2 Myr in age, it is the only tropical oceanic island in the eastern Pacific with tropical wet forest. We identified several hundred bark beetle specimens collected during recent expeditions by INBio, the National Biodiversity Institute of Costa Rica, and re-examined all specimens from earlier collections. We report 19 species in ten genera, seven or eight of which are endemic, making scolytines the largest group of beetles known from the island. We describe as new Pycnarthrum pseudoinsulare , Xyleborinus cocoensis , and Xyleborus sparsegranulosus , resurrect Xyleborus bispinatus as separate from X. ferrugineus , and report six other species as new to Cocos Island. Three-quarters of the scolytines reproduce by brother–sister mating, and we argue that inbreeders are superior island colonists because they are less affected than are outbreeders by problems of mate location and inbreeding depression. The fauna and flora of Cocos Island arrived by dispersal and human transport. We examine natural colonization patterns for the fauna, using the distributions of the relatives of island endemics: most colonization came from the Americas, but the closest relatives to some endemics are found on Caribbean or Galápagos islands. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 729–743.     

On the mechanism of floral shifts in speciation: gained pollination efficiency from tongue- to eye-attachment of pollinia in Platanthera (Orchidaceae)

This work explores the mechanism of floral shifts that may result in speciation. The model organisms chosen were the moth-pollinated pollinator-limited orchid species Platanthera bifolia and P. chlorantha . P. bifolia exhibits tongue-attachment of pollinia on pollinators, a character state that has been found previously to be ancestral. The close relative P. chlorantha exhibits the derived state of eye-attachment of pollinia on pollinators. We reasoned that differences between the species in pollination efficiency could give insights into the mechanism of floral shifts and thus cladogenesis. Four populations per species were investigated. In three populations, where the species were growing intermixed and were sharing pollinators, there was significant difference in the pollen export and import efficiency per visit-night (night with pollen export and/or import) between the two species. P. bifolia exported pollinia more efficiently but imported pollen less efficiently than did P. chlorantha . Pollen import was 1.7–4 times faster in P. chlorantha (eye-attachment) than it was in P. bifolia (tongue-attachment). P. chlorantha had a lower risk of interference between pollen import and export. An increase in fitness through greater speed and efficiency of pollen import due to an enlargement of the stigmatic surface and a reduction in the risk of sexual interference may therefore be mechanisms of the floral shift from tongue- to eye-attachment of pollinia on pollinators.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 481–495.     

Flowering in a bamboo, Melocanna baccifera (Bambusoideae: Poaceae)

Melocanna baccifera (Roxburgh) Kurtz ex Skeels, a species of bamboo introduced to Sri Lanka from India, flowered and set fruit during 2001–2002. Culms that flowered and set fruit died. The incidence of flowering is significant in that flowering took place close to the predicted mast flowering in 2007. At the onset of flowering, inflorescences were predominantly staminate. But later in 2002, bisexual and pistilate flowers also developed leading to fruiting. Both protandry and protogyny were observed in the bisexual florets. Floral characters indicated that the species was mainly out-crossing. Although anther dehiscence released pollen and stigmas were exerted, pollination of stigmas was inefficient. The few stigmas that were naturally pollinated showed limited pollen tube growth. However, fruit set took place. Fruits were very rarely seen to germinate naturally. Many fruits were devoid of an embryo, indicating that parthenocarpy and/or apomixis may have taken place. Excised embryos germinated in vitro .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 287–291.     

A preliminary palynological characterization of Spanish thyme honeys

Pollen was analysed in 25 thyme honey samples from Spain. The honey samples were provided by professional beekeepers, and pollen was obtained by centrifugation. A minimum of 8% of Thymus sp. pollen is considered sufficient to typify a honey as unifloral when considering pollen grains from anemophilous and nectarless plants. The quantitative analysis showed that nectar is the main honey source in the samples studied. The qualitative analysis of the samples revealed the presence of 53 taxa belonging to 27 families. The Spanish thyme honeys of the studied region are characterized by their medium to high numbers of grains of pollen (NGP; mean = 155 000) and their medium to high content of honeydew indicator elements (HDE; mean = 8300). Genista type (present in 100% of the samples), Echium plantagineum and Reseda luteola (90%), Eucalyptus type (80%), Calendula type and Olea europaea (75%), and Cistus monspeliensis (70%) may be mentioned among the characteristic accompanying species of this honey type.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 323–330.     

Speciation in Darwin's darklings: taxonomy and evolution of Stomion beetles in the Galápagos Islands, Ecuador (Insecta: Coleoptera: Tenebrionidae)

Stomion is one of three speciose and taxonomically difficult genera of flightless Darwin's darkling beetles of the Galápagos Islands. The previously named species and subspecies of Stomion are analysed by morphometric and cladistic methods. Eleven species are here recognized, which probably descended from colonization by a single ancestral species. Cladistic analysis yielded two weakly supported clades that are different from previously published morphological and allozyme genotype groups. Phenotypic and genotypic characters may be evolving independently. There is much within-species variation, but there is no compelling evidence of adaptive radiation in Stomion , in contrast to that in the famous Darwin's finches. The analyses show the need for changes in morpho-species taxonomy: S. punctipennis Van Dyke ( syn. nov. ) is elevated to full species from a subspecies of S. galapagoensis Waterhouse. There is no evidence to support the validity of: S. laevigatum santacruzensi Franz ( syn. nov. ), synonomized here under S. l. laevigatum Waterhouse; S. galapagoensis leleupi Kaszab ( syn. nov. ), synonomized here under S. linelli Blair; S. galapagoensis pinzoni Franz ( syn. nov. ), synonomized here under S. obesum Van Dyke. S. genovesa sp. nov. is described from Genovesa Island. Lectotypes are designated for S. galapagoensis Waterhouse, S. laevigatum Waterhouse and S. helopoides Waterhouse .   © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 141 , 135–152.     

Pollination ecology of Arum italicum (Araceae)

The pollination ecology of Arum italicum was studied in south-western France. This plant attracts olfactory dung-breeding flies through deceit. These insects are principally represented by Diptera, all belonging to saprophyte families. The volatilization of the odouriferous compounds, responsible for their attraction, is achieved through the production of heat by the appendix. The insects are trapped for 24 h in order to participate in both sexual phases of the protogynous inflorescence. The male flowers produce three heat events during flowering. These peaks of heat seem to be involved in the spathe movements, since they occur during the opening of the inflorescence and the liberation of the insects. The last male heat event may be linked with the liberation of pollen and its dispersion by stimulating trapped flies. According to their frequency and pollen-load, two Psychoda species appear to be the most efficient pollinators ( P. crassipenis and P. pusilla ). Nevertheless, each of the other attracted species could play a significant role under different spatio-temporal conditions. Experiments on self-pollination have shown that obligate cross-pollination is necessary for A. italicum to set seeds. Moreover, hand- and natural-pollinated plants showed similarly high abortion frequencies suggesting that seed set may be more constrained by resources rather than by pollination limitation.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 205–214.     

Pollination of the lady's slipper Cypripedium henryi Rolfe (Orchidaceae)

The pollination ecology of Cypripedium henryi Rolfe, a slipper orchid endemic to west China, was investigated, and its floral shape, size, colour, and scent were analysed. Examination of the breeding system suggests that the flowers are self-compatible, but need pollen vectors for successful reproduction. The flower is rewardless; over 15 insects belonging to Araneida, Hymenoptera, Diptera, Lepidoptera, and Coleoptera were recorded as flower visitors, but most only alighted or rested on the flower. In the total 32 h of observations over 2 years, female Lasioglossum bees were found to be the most frequent visitors and the only pollinators. They showed a high visitation frequency and, surprisingly, re-visited the same flowers frequently. Cypripedium henryi probably attracts pollinators visiting the flowers through general food deception (odour components, colour, false nectar guides), as well as special structures (slippery labellum, slippery staminode). Although three Lasioglossum species visited the flowers, only L. sauterum Fan et Ebmer was found with pollen. Lasioglossum flavohirtum Ebmer was large and climbed out from the entrance. Morphologically, L. sichuanense Fan et Ebmer could be considered as a potential pollinator, but the collected specimens were found to have no pollen of C. henryi on their bodies. It was speculated that the particular floral scent of C. henryi discouraged the entrance of L. sichuanense bees. Lasioglossum sauterum was matched morphologically to the flower, but not all of the visitations resulted in effective pollinations, as some flowers of C. henryi were frequently re-visited and the pollen mass had been taken away by bees on previous visitations. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 491–499.     

Pollination by fungus gnats in four species of the genus Mitella (Saxifragaceae)

The first example of pollination by fungus gnats in the eudicots is reported. The genus Mitella (Saxifragales) is characteristically produces minute, inconspicuous, mostly dull-coloured flowers with linear, sometimes pinnately branched, petals. To understand the function of these characteristic flowers, we studied the pollination biology of four Mitella species with different floral traits and different sexual expression: dioecious M. acerina , gynodioecious M. furusei var. subramosa , and hermaphroditic M. stylosa var. makinoi and M. integripetala. Flower-bagging experiments showed that wind pollination did not occur in the dioecious and gynodioecious species. Two years of observations of flower visitors at six study sites in Japan revealed that the principal pollinators of all four Mitella were specific species of fungus gnats (Mycetophilidae), which landed on the flowers with their long spiny legs settling on the petals. Characteristically, numerous pollen grains were attached to the fungus gnats in specific locations on the body. Although, on average, 1.3–2.6 fungus gnats visited each inflorescence per day, the fruit set of both bisexual and female flowers exceeded 63%. These results suggest that fungus gnats are highly efficient pollinators of Mitella spp., and that Mitella flowers are morphologically adapted to pollination by fungus gnats.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 449–460.     

Systematic relevance of pollen and orbicule characters in the tribe Hillieae (Rubiaceae)

Pollen and orbicule morphology of 26 species in the tribe Hillieae is described based on observations by light, scanning and transmission electron microscopy. Pollen and orbicule characters are critically evaluated and discussed in the context of existing hypotheses of systematic relationships within the tribe. Pollen is 3-zonocolporate with a perforate, microreticulate, reticulate or eutectate sexine. In the two species of Blepharidium , however, the pollen has one, four or five apertures. These pollen morphological data were incorporated into an existing macromorphological matrix of the group and cladistically analysed. The resulting phylogenies indicate that it may be appropriate to reduce Cosmibuena to a subgenus of the genus Hillia , while Blepharidium should be removed from Hillieae because of the deviating pollen type that is unique to the Rubiaceae. All species investigated produce orbicules, which are mostly spherical and possess an electron-lucent core that is sometimes characteristically flattened.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 303−321.     

The cytological origin of the polyads and their significance in the reproductive biology of Mimosa bimucronata

The mechanism of polyad formation in Mimosa bimucronata (DC.) Kuntze is described and its significance in the reproductive biology is discussed. In this species c . 500 complexes of two coupled pollen mother cells (PMCs) per pollen sac are formed after the last mitotic division of the sporogenous tissue. All the pollen grains of each complex remain attached and form a bitetrad. This mechanism leads to the formation of many polyads per pollen sac. On this basis, we propose that in Mimosa the number of pollen grains per polyad (eight or 16) is determined by the number of cells that remain attached after the last mitotic divisions of the sporogenous tissue. Concerning the biological significance of the polyads in M. bimucronata , we find that the number of pollen grains per polyad is correlated with the number of ovules per ovary, and that the internal dimensions of the pit-shaped stigma are sufficient just for one polyad to be localized in it. Based on the number of polyads per flower this species has a high pollen/ovule ratio characteristic of xenogamous plants, in opposition to that expected for plants with composite pollen dispersal units. In contrast, it has low pollination efficiency, low fruit set, a 1/1 relationship between the pollen grains of a polyad and the number of ovules per ovary, and high seed set per fruit. These facts support the hypothesis that the occurrence of polyads in M. bimucronata would be an adaptation helping to ensure high seed set following a single cross pollination event.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 343–349.     

Flower-visiting bees and the breakdown of the tristylous breeding system of Eichhornia azurea (Swartz) Kunth (Pontederiaceae)

Populations of Eichhornia azurea (Swartz) Kunth (Pontederiaceae) are often composed of three floral morphs that differ in style length and anther position. Hence, E. azurea is described as tristylous. For sexual reproduction the plant requires specialized pollinators able to transfer pollen from flowers with long-, mid-, or short-level anthers to flowers with long-, mid-, or short-styles, respectively. This is because within a floral morph, self-incompatibility is the rule. Populations of E. azurea were studied from six localities in south-eastern Brazil (São Paulo and Minas Gerais). The frequencies of floral morphs, floral morphology and flower-visiting bees were recorded in each population. Ten different bee species were observed visiting plants. Four of them were considered generalist bees and not appropriate for legitimate pollination of the three floral morphs. Another three species were considered opportunists and only three species were considered to be potential pollinators. At the coastal region (Juréia), the plant population lacked the short-styled morph and the mid-styled morph and exhibited a semi-homostylous condition, in which the low-level anthers were adjacent to the stigma. This morph showed high level of self-compatibility. The disappearance of the short-styled morph from this region is likely correlated with the absence of Ancyloscelis gigas Friese, a specialized long-tongued bee that is oligolectic on this plant in other localities.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 77 , 499–507.     

Chemical communication in the sexually deceptive orchid genus Cryptostylis

Pollination by sexual deception is among the most intriguing of orchid pollination syndromes. Odours are well established as the primary stimuli for sexually attracting the male insect pollinators in these orchids. We applied gas chromatography with electroantennographic detection (GC-EAD) to investigate chemical communication between the sympatric, but morphologically distinct, orchids Cryptostylis erecta and C. subulata and their pollinators. Cryptostylis is unusual among sexually deceptive orchid genera in that all five Australian species share the same pollinator, the ichneumonid wasp Lissopimpla excelsa , but hybrids are unknown. We show that volatile odour compounds are not produced in detectable amounts in either species. Floral extracts containing many low-volatility compounds showed considerable differences in composition between C. erecta and C. subulata . By contrast, GC-EAD revealed the male wasp pollinators are electrophysiologically responsive to the same GC peak on two different kinds of GC column in both orchids. This leads us to predict that a single compound is the sexual attractant in all Australian Cryptostylis . The apparent conservation of chemical signals among distinct species contrasts with that of other sexually deceptive orchids that are often morphologically similar but reproductively isolated by their different chemical signals.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 199–205.     

Morphological variation among populations of Lecocarpus (Asteraceae) on the Galápagos Islands

Representative samples were collected from almost all known populations of the endemic Galapagos genus Lecocarpus. Multivariate statistical methods were applied to morphological characters to investigate differentiation among species and populations. In discriminant analysis no misclassifications were made among species. Populations of L. darwinii and L. lecocarpoides were better discriminated than populations of L. pinnatifidus . Principal Component Analysis (PCA) separated species well although intermediate populations occur between L. darwinii and L. lecocarpoides . Clear patterns of within-species differentiation were seen among populations of L. darwinii and L. lecocarpoides , but not among populations of L. pinnatifidus .
Populations of L. lecocarpoides at present grow on separate islands. All populations of L. darwinii are found on San Cristóbal, but this island might have been separated into more islands in the past. Lecocarpus pinnatifidus has probably always been growing on only one island. This suggests that the sea is the major barrier to dispersal of the three species. We find it likely that the degree of reproductive isolation caused by the sea is crucial to the differentiation among populations, and that genetic drift rather than adaptation has been responsible. The analyses cast new light on the identity of classic collections made by Darwin in 1835 and Stewart in 1906. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 523–544.     

A palynological study of the genus Mentha L. (Lamiaceae)

The pollen morphology and exine structure of 10 Mentha L . species were investigated using light microscopy and scanning electron microscopy. The pollen grains of all 11 species were hexazonocolpate with granular membranes and a circular amb, varying in shape from prolate-spheroidal to suboblate. Different colpus shapes were recognized in M . ×  dumetorum . The exine was bireticulate in section Pulegium , and reticulate in section Menthae . A correlation was found between pollen size and chromosome number. The results indicate that the pollen characters of the genus Mentha are valuable for taxonomic applications and may be useful for classification.  © 2008 Uludag University. Journal compilation © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 141–154.