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.     

广东地区外来种五爪金龙的传粉生物学研究（英文）

五爪金龙(Ipomoea cairica)是一种匍匐或攀援生长的入侵杂草。2006-2007年作者在广东地区开展了对该入侵种的传粉生物学研究。五爪金龙在广东全年均可开花,花序为聚伞花序,花冠为紫色或蓝紫色喇叭状。开花时间在早晨4:30-5:20之间,下午17:40闭合,持续大约12h。每朵花平均分泌花蜜量为5.82±2.47μL(n=10),花蜜平均含糖量为41.03±6.52%,氨基酸含量为0.042mg/mL。开花过程中花粉活性和柱头可受性均保持较高水平(>90%)。花粉/胚珠比率(P/O)为4,778.18±264.58(n=10)。木蜂(Xylocopasp.)是其有效的传粉昆虫,蝇类和蝶类为随机访问者。开花过程中柱头与花药相接触,有利于自花授粉。然而,繁育系统的研究表明五爪金龙是自交不亲和植物,其种子仅能通过异花授粉获得。     

广东地区外来种五爪金龙的传粉生物学研究

五爪金龙(Ipomoea cairica)是一种匍匐或攀援生长的入侵杂草。2006–2007年作者在广东地区开展了对该入侵种的传粉生物学研究。五爪金龙在广东全年均可开花, 花序为聚伞花序, 花冠为紫色或蓝紫色喇叭状。开花时间在早晨4: 30–5: 20之间, 下午17: 40闭合, 持续大约12 h。每朵花平均分泌花蜜量为5.82 ± 2.47 μL (n = 10), 花蜜平均含糖量为41.03 ± 6.52%, 氨基酸含量为0.042 mg/mL。开花过程中花粉活性和柱头可受性均保持较高水平(>90%)。花粉/胚珠比率(P/O)为4,778.18 ± 264.58 (n = 10)。木蜂(Xylocopa sp.)是其有效的传粉昆虫, 蝇类和蝶类为随机访问者。开花过程中柱头与花药相接触, 有利于自花授粉。然而, 繁育系统的研究表明五爪金龙是自交不亲和植物, 其种子仅能通过异花授粉获得。     

毛菍传粉生物学研究

为探讨毛菍(Melastoma sanguineum Sims.)传粉生物学特性,对海南霸王岭国家级自然保护区林缘、林内和林隙3个生境内毛菍的花期、花部形态、访花昆虫等进行观察,同时对其繁育系统相关指标进行了测定。结果表明：毛菍无花蜜分泌;花瓣对访花昆虫有吸引作用;访花昆虫主要有蜂类、食蚜蝇类和丽金龟类;主要传粉昆虫有木蜂、熊蜂,其中木蜂传粉活动最为活跃,为最有效传粉昆虫。毛菍具典型的异型雄蕊,长、短雄蕊在传粉过程中具有明确的功能分化,推测这是毛菍保证传粉成功的策略。毛菍为自交亲和的异交种,需要昆虫传粉;人工自交和异交均具有较高的坐果率;不存在无融合生殖、主动自花授粉和滞后自交的生殖保障现象;其繁育系统是兼性自交。昼夜温差较小和相对湿度较高的林内和林隙生境有利于延长群体花期;光照充足、温度相对较高的林缘生境,花朵发育较快,柱头可授性的时间稍早。花粉、传粉者和环境条件是制约毛菍野外传粉效率的主要因素。     

Pollination Ecology and Effect of Nectar Removal in Macleania bullata (Ericaceae)1

The floral syndrome of Macleania bullataYeo (Ericaceae) reflects its adaptation to hummingbird pollination. Its flowers, however, are subject to high levels of nectar robbing. I examined the floral visitor assemblage of M. bullata in a tropical montane wet forest in southwestern Colombia, focusing on the behavior of the visitors. I also tested for the presence of nocturnal pollination and the effects of nectar removal on new nectar production. The principal floral visitors were the nectar robbing hummingbirds Ocreatus underwoodii (19.1% of visits) and Chlorostilbon mellisugus (18.9%). Only two species of long–billed hummingbirds visited the flowers of M. bullata as “legitimate” pollinators: Coeligena torquata (14.7% of visits) and Doryfera ludoviciae (14.3%). The remaining visits constituted nectar robbing by bees, butterflies, and other species of hummingbirds. Nocturnal pollination took place, although fruit set levels were 2.4 times higher when only diurnal pollination was allowed as opposed to exclusively nocturnal pollination. Nectar robbers removed floral nectar without pollinating the flower. Treatments of experimental nectar removal were carried out to examine if flowers synthesize more nectar after nectar removal. Nectar removal increased the total volume of nectar produced by each flower without affecting sugar concentration. Thus, nectar robbing can impose a high cost to the plants by forcing them to replace lost nectar.     

Reproductive biology ofWahlenbergia (Campanulaceae) endemic to Robinson Crusoe Island (Chile)

The reproductive biology ofW. berteroi, W. fernandeziana, and a putative hybrid betweenW. fernandeziana andW. grahamiae, endemic to Robinson Crusoe Island (Juan Fernández archipelago, Chile) was studied. Flowers are hermaphroditic, protandrous, offer nectar, and exhibit secondary pollen presentation involving pollen collecting hairs on the style. These features imply allogamy and biotic pollination. However, male and female phases overlap and no effective pollinators were observed. Experimental data indicate these taxa are self-compatible and facultatively autogamous, a conclusion also suggested by the pollen/ovule ratios. Selfing is accomplished when the stigmatic lobes reflex and touch the style, except forW. berteroi where they do not reflex completely. Autogamy is accomplished in the latter when pollen grains deposited on the inner surface of the corolla throat by the pollen brush are gathered by stigmatic lobes when shaken by wind. The degree of autogamy, and perhaps self-compatibility, seems to be inconstant, as implied by the variable natural seed set (overall range 21–188 seeds per fruit). A mixed mating system — primarily outcrossing/entomophilous, but also autogamous — must have been present in the continental ancestors of these taxa. Autogamy promoting self-fertilization is important now — on an island with scarce pollinators — and in the past — when the first founders arrived.     

Reproductive biology and responses to threats and protection measures of the total population of a Critically Endangered Galápagos plant,Linum cratericola (Linaceae)

We describe a reproductive and population dynamic study of the total population of a Critically Endangered plant over 10 years, during which part of the population was protected from the main threat, feral ungulates. Linum cratericola (Linaceae) was first discovered in 1966 at two sites on Floreana Island, Galápagos. It has since disappeared from one site and survives in three groups of plants at the other, where it has been threatened with extinction by introduced ungulates and invasive plants. Population size, and growth rate and mortality of individual plants, have been monitored since 1997. The population appears highly responsive to changes in threat levels: the three plant groups increased rapidly following protection by fencing and ungulate control, but temporarily declined when feral goat pressure increased and during dry periods. Natural factors that may contribute to population limitation include dry years, grazing by native snails and competition from native vascular plants and cryptogams. Linum cratericola has a single flowering period per year and produces abundant seed with 28% germination after scarification, but with no obvious adaptations for long‐distance dispersal. Potential pollinators included the butterfly Leptotes parrhasioides, the hoverfly Toxomerus crockeri and the carpenter bee Xylocopa darwinii, all endemic to Galápagos. The continued survival of L. cratericola in the wild depends on effective protection from introduced herbivores and invasive plants. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 161 , 89–102.     

红豆杉的胚珠发育,传粉滴形成和传粉过程

观察了红豆杉（Ｔａｘｕｓ ｃｈｉｎｅｎｓｉｓ （Ｐｉｌｇ．）Ｒｅｈｄ）的花粉形态和水合特性及胚珠发育、传粉滴形成与传粉过程。成熟花粉为单细胞,无气吓,形状不规则,外壁表面具大量乌氏体。花粉水合时,内壁膨胀,外壁开裂。通常情况下,外壁保留在水滴或传粉滴的表面,而花粉的其他部分进入水滴或传粉滴内。在８月下旬,可观察到下弯的雌性生殖芽。下弯这一特性是雌性生殖芽区别于营养芽的重要特征。这一时期的雌性生殖芽     

Population genetic structure and levels of inbreeding depression in the Mediterranean island endemic Cyclamen creticum (Primulaceae)

The variation and evolution of reproductive traits in island plants have much attention from conservation and evolutionary biologists. However, plants on islands in the Mediterranean region have very little attention. In the present study, we examine the floral biology and mating system of Cyclamen creticum , a diploid perennial herb endemic to Crete and Karpathos. Our purpose is to quantify (1) variation and covariation of floral traits related to the mating system, (2) the ability of the species to self in the absence of pollinators and its relative performance on selfing and outcrossing and (3) generic diversity within and among populations. Pollen/ovule ratios were indicative of a xenogamous species. A controlled pollination experiment showed that the species is self-compatible but is unable to set seed, in the absence of pollinators, probably due to stigma-anther separation. A multiplicative estimate of inbreeding depression based on fruit maturation, seed number and percentage seed germination gave δ= 0.38 Population genetic diversity was high, 54.76% polymorphic loci, a mean of 1.78 alleles per locus and a mean observed heterozygosity of 0.053. F -statistics nevertheless indicated high inbreeding rates (mean F ^is= 0.748) in natural populations, and low levels of population differentiation (mean F^is= 0.168). C. creticum thus appears to have a mixed-mating system with high levels of (pollinator) mediated inbreeding (either by facilitated selfing, geitonogamy or biparental inbreeding) in natural populations.     

The reproductive biology of the Mediterranean endemic Cyclamen balearicum Willk. (Primulaceae)

Knowledge of the reproductive biology of endemic plants improves our understanding of how mating system may be related to patterns of species abundance and provides a basis for the development of rational conservation programmes. In this paper we present natural population data on the floral biology and reproductive ecology of the endemic Mediterranean species Cyclamen balearicum Willk. This is a long-lived, diploid perennial herb which occurs in southern France in five fragmented and isolated regions and on the Balearic Islands of Mallorca, Menorca, Ibiza, Cabrera and Draponera. Our observations indicate a particularly scarce pollinator activity (rare syrphid visits) and dispersal by ants over small distances. A controlled pollination experiment in a natural population showed that in southern France C. balearicum is fully self-compatible and that selfing is autonomous and probably delayed (i.e. following opportunities for outcrossing). The proximity of stigmas and anthers will favour autonomous selfing. The high pollen/ovule ratio indicates nevertheless that C. balearicum has a mixed mating system. Patterns of variation in stigma-anther separation and pollen production per flower suggest that not only has the current mating system of the species evolved from an outcrossing ancestor but that due to the fragmentation and isolation of populations greater levels of selfing have evolved in southern France (and to an intermediate degree on Ibiza and Menorca). On the island of Mallorca where larger continuous belts of C. balearicum habitat still exist the species has floral traits indicating a more outcrossed mating system. To our knowledge this is the first paper to document such trends in floral traits in the endemic component of the Mediterranean flora.     

Cryptic diversity in Black rats Rattus rattus of the Galápagos Islands,Ecuador

Human activity has facilitated the introduction of a number of alien mammal species to the Galápagos Archipelago. Understanding the phylogeographic history and population genetics of invasive species on the Archipelago is an important step in predicting future spread and designing effective management strategies. In this study, we describe the invasion pathway of Rattus rattus across the Galápagos using microsatellite data, coupled with historical knowledge. Microsatellite genotypes were generated for 581 R. rattus sampled from 15 islands in the archipelago. The genetic data suggest that there are at least three genetic lineages of R. rattus present on the Galápagos Islands. The spatial distributions of these lineages correspond to the main centers of human settlement in the archipelago. There was limited admixture among these three lineages, and these finding coupled with low rates of gene flow among island populations suggests that interisland movement of R. rattus is rare. The low migration among islands recorded for the species will have a positive impact on future eradication efforts.     

Resource limitation and pollen source (self and outcross) affecting seed production in two louseworts, Pedicularis siphonantha and P. longiflora (Orobanchaceae)

Two animal-pollinated hermaphrodite plants, Pedicularis siphonantha and P. longiflora , have been used to investigate factors limiting seed production in natural populations. To evaluate the potential seed abortion due to resources limitation, seed development has been observed and seed count conducted twice. Seed production per capsule has been compared when flowers have been removed and in a control group. Open pollination has been investigated and pollen supplementation undertaken to estimate the possibility of pollen limitation. Results show that seed abortion is frequent. Stigmatic pollen load is significantly higher than ovule number per ovary under open pollination for both species. Additional self and outcross pollen did not affect seed production. Flower removal significantly increases seed production per capsule, which indicates that seed production of the studied species is limited by available resources. To detect differences in seed production between flowers pollinated by self and outcross pollen, hand pollination of bagged flowers has also been conducted in natural populations of the two Pedicularis species. Compared with open pollination, hand-pollinating self-pollen decreases, while outcross pollen increases seed production per capsule. Such results suggest that inbreeding depression in the two self-compatible species may also result in partial seed abortion under open pollination if mixed pollen is deposited on the stigma. Our results also suggest that pollen interference plays an important role in low female fertility in the two Pedicularis species.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 83–89.     

Fine‐tuning the nested structure of pollination networks by adaptive interaction switching,biogeography and sampling effect in the Galápagos Islands

The structure of pollination networks, particularly its nestedness, contain important information on network assemblages. However, there is still limited understanding of the mechanisms underlying nested pollination network structures. Here, we investigate the role of adaptive interaction switching (AIS), island area, isolation, age and sampling effort in explaining the nestedness of pollination networks across ten Galápagos Islands. The AIS algorithm is inspired by Wallace's elimination of the unfit, where a species constantly replaces its least profitable mutualistic partner with a new partner selected at random. To explain network structures, we first use a dynamic model that includes functional response of pollination and AIS, with only species richness and binary connectance as input (hereafter the AIS model). Thereafter, other explanatory variables (isolation, area, age and sampling effort) were added to the model. In four out of ten islands, the pollination network was significantly nested, and predictions from the AIS model correlated with observed structures, explaining 69% variation in nestedness. Overall, in terms of independent contribution from hierarchical partitioning of variation in observed nestedness, the AIS model predictions contributed the most (37%), followed by sampling effort (28%) and island area (22%), with only trivial contributions from island isolation and age. Therefore, adaptive switching of biotic interactions seems to be key to ensure network function, with island biogeographic factors being only secondary. Although large islands could harbour more diverse assemblages and thus foster more nested structures, sufficient sampling proves to be essential for detecting non‐random network structures.     

Delayed autonomous self‐pollination in two Japanese varieties of Epipactis helleborine (Orchidaceae)

The terrestrial orchid Epipactis helleborine is a morphologically variable species with a wide geographical distribution. It is found throughout Europe and continues eastwards to Siberia, China and Japan. It is usually pollinated by social wasps and displays the morphological characteristics of an outcrossing species. In warm, temperate areas of Japan, E. helleborine often appears in alpine or subalpine regions, and has never been found in low‐altitude forests, except for coastal pine forests. The coastal population of E. helleborine is often classified at the variety level, as E. helleborine var. sayekiana, and the inland populations are known as E. helleborine var. papillosa. It is possible that E. helleborine var. sayekiana possesses a distinctive selfing strategy, as its autonomous self‐pollination has evolved in dry habitats, such as coastal dunes. The present study investigated the pollination biology of E. helleborine var. sayekiana and var. papillosa to detect differences in their reproductive systems. Unexpectedly, both E. helleborine var. papillosa and E. helleborine var. sayekiana were found to possess a self‐pollination strategy and were therefore rarely visited by insects. Self‐pollination occurs at the end of the flowering season and probably acts to ensure pollination even if insect‐mediated pollination fails. Moreover, there are no floral differences between E. helleborine var. papillosa and E. helleborine var. sayekiana. These observations suggest that it is not necessary to distinguish between these two varieties. Furthermore, E. helleborine var. papillosa was pollinated by hoverflies, suggesting that E. helleborine var. papillosa is probably biologically distinct from its mother species, E. helleborine. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 733–743.     

Reproductive ecology of the endemic Lecocarpus pinnatifidus (Asteraceae) in an isolated population in the Galápagos Islands

Lecocarpus pinnatifidus is an endemic member of the Asteraceae occurring on only one island in the Galápagos archipelago. The capitula are large with female ray florets and male disc florets. They are self-compatible but this study suggests fruit set is pollen limited. Visits from Xylocopa darwini and other larger insect pollinators are rare, and small insects seem to be the main pollinators. Small insects carry few pollen grains and most likely mediate self-pollinations. Self-compatibility and seed set after selfing are the most common reproductive strategy in the Galápagos Islands and L. pinnatifidus seemingly fits well into this group.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 171–180.     

Nocturne for an unknown pollinator: first description of a night‐flowering orchid (Bulbophyllum nocturnum)

Bulbophyllum nocturnum , a species of section Epicrianthes from New Britain, is described and illustrated. It is the first known example of an orchid species in which the flowers open after dark and close in the morning. The poorly understood pollination biology of section Epicrianthes, a clade with highly unusual flowers, is discussed. Attention is drawn to the close resemblance between the petal appendages of some species and the fruiting bodies of certain Myxogastria. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 344–350.     

Differential self‐fertilization rates in response to variation in floral traits within inflorescences of Aquilegia buergeriana var. oxysepala (Ranunculaceae)

To assess variation in the proportion of self‐fertilized seeds among flowers within inflorescences and the relationship between floral traits and the rate of self‐fertilization, the proportion of self‐fertilized seeds among individual flowers was estimated using ten microsatellite markers in self‐compatible plants of Aquilegia buergeriana var. oxysepala. Within‐inflorescence variation in floral traits, such as the duration of the male and female phases, flower size, herkogamy and the number of pollen grains and ovules in two natural populations, were investigated. The first flower in an inflorescence produced more seeds and a higher proportion of self‐fertilized seeds than the second flower. The higher proportion of self‐fertilized seeds in the first flowers was accompanied by a higher number of pollen grains and ovules in the bud stage and the female phase. These results indicate that the high proportion of self‐fertilized seeds in the first flowers in an inflorescence may be due to the high number of remaining pollen grains in the female phase. This suggests that variation in floral traits within inflorescences affects seed quality and quantity among flowers within inflorescences.