Erosion of a pollination mutualism along an environmental gradient in a south Andean treelet,Embothrium coccineum (Proteaceae)

The limit of a species’ distribution can be determined biotically if an environmental gradient causes the loss of critical mutualists such as pollinators. We assessed this hypothesis for Embothrium coccineum, a self‐incompatible red‐flowered treelet growing along a strong west‐east precipitation gradient from rainforest to forest‐steppe ecotone in the rain shadow of the southern Andes in northwestern Patagonia. For 16 populations along this gradient, we quantified composition of the pollinator assemblage, pollination efficiency and limitation, and reproductive output. The treelet has a generalized pollination system, but the hummingbird Sephanoides sephaniodes was the most effective pollinator. The relative importance of this hummingbird as a flower visitor within populations influenced pollen transfer and fruit set more strongly than local precipitation. As hummingbirds and other pollinators, including passerine birds and nemestrinid flies, were replaced by bees towards the dry eastern range limit, pollen limitation increased and reproduction eventually failed. These results support the hypothesis that pollinators can act as important biotic filters influencing plant distribution, and warn against predictions of geographical range shifts based solely on climatic variables.     

Foraging behaviour of bird pollinators on Embothrium coccineum (Proteaceae) trees in forest fragments and pastures in southern Chile

Abstract We investigated the effects of forest patch size on the behaviour of birds feeding on the flower nectar of the proteaceous tree Embothrium coccineum J. R. et G. Forster, which is typically restricted to forest edges in agricultural landscapes in southern Chile. We quantified reproductive parameters of trees (no. inflorescences per branch, total and open flowers per inflorescence) in forest fragments varying from 1 ha (small), to 20 ha (medium) and to >150 ha (large), and in remnant trees in pastures. Visits to flowers by nectar‐feeding birds were recorded during 30‐min observation periods, spread throughout the day during two flowering seasons, November 1992 and 1993 (n = 242 periods overall). Aggressive encounters among flower visitors were recorded in 1992. We expected less visits to trees in pastures and small forest patches because abundances of Embothrium's main pollinators, the flycatcher Elaenia albiceps and the hummingbird Sephanoides sephaniodes, decreased in smaller patches. We found, however, that pollinator visiting rates were negatively correlated with forest patch area and were highest for pasture trees. This trend was largely due a decline in the number of visits by E. albiceps, the main flower visitor, in larger patches. Hummingbird visits did not change with forest patch size. Lower visitation rates to flowering trees in larger forest fragments seemed to be a consequence of territorial defence by E. albiceps and were unrelated to differences in floral display.     

Floral trait evolution associated with shifts between insect and wind pollination in the dioecious genus Leucadendron (Proteaceae)

Transitions between animal and wind pollination have occurred in many lineages and have been linked to various floral modifications, but these have seldom been assessed in a phylogenetic framework. In the dioecious genus Leucadendron (Proteaceae), transitions from insect to wind pollination have occurred at least four times. Using analyses that controlled for relatedness among Leucadendron species, we investigated how these transitions shaped the evolution of floral structural and signaling traits, including the degree of sexual dimorphism in these traits. Pollen grains of wind‐pollinated species were found to be smaller, more numerous, and dispersed more efficiently in wind than were those of insect‐pollinated species. Wind‐pollinated species also exhibited a reduction in spectral contrast between showy subtending leaves and background foliage, reduced volatile emissions, and a greater degree of sexual dimorphism in color and scent. Uniovulate flowers and inflorescence condensation are conserved ancestral features in Leucadendron and likely served as exaptations in shifts to wind pollination. These results offer insights into the key modifications of male and female floral traits involved in transitions between insect and wind pollination.     

Patterns of species turnover in plant‐pollinator communities along a precipitation gradient in Patagonia (Argentina)

Recent studies have assessed the influence of different types of gradients (e.g. altitudinal, latitudinal and temporal, among others) on the structure and function of community‐level plant‐pollinator webs. Although the importance of humidity as a major driver of species‐richness gradients worldwide has been stressed by recent reviews, no studies have been specifically designed to address the influence of precipitation gradients on pollination webs. In the present paper we describe for the first time the turnover of species of plants and their associated flower visitors between eight communities located along a steep precipitation gradient in north‐west Patagonia, Argentina. Our results show that: (i) there is a high spatial turnover of plant communities and their associated pollinators; (ii) this turnover is strongly related to precipitation changes across the region; and (iii) the turnover rate is similar for plants and pollinators. Our results support the view that the precipitation gradient is a significant factor associated with the regional turnover of plants and their pollinators in the temperate forests of southern South America.     

Long‐term ‘islands’ in the landscape: low gene flow,effective population size and genetic divergence in the shrub Hakea oldfieldii (Proteaceae)

The genetic structure of disjunct populations is determined by founding genetic properties, demographic processes, gene flow, drift and local selection. We aim to identify the genetic consequences of natural population disjunction at regional and local scales in Hakea oldfieldii using nuclear and plastid markers to investigate long‐term effective population sizes and gene flow, and patterns of diversity and divergence, among populations. Regional divergence was significant as shown by a consistent pattern in principal coordinates, neighbor‐joining and Bayesian analyses, but divergence at the local level was also significant with localized distribution of plastid haplotypes and populations clustering separately in Bayesian analyses. Historical, recent and first‐generation gene flow was low, suggesting that recent habitat fragmentation has not reduced gene migration significantly. Genetic bottlenecks were detected in three populations. Long‐term effective population size was significantly correlated with the number of alleles/locus and observed heterozygosity, but not with census population size, suggesting that the loss of diversity is associated with long‐term changes rather than recent fragmentation. Inbreeding coefficients were significant in only three populations, suggesting that the loss of diversity is linked to drift and bottlenecks associated with demographic processes (local extinction by fires) rather than inbreeding. Historical disjunction as a result of specific ecological requirements, contraction of habitats following drying during the Pleistocene, low gene flow and changes in population size are likely to have been important forces driving divergence through isolation by distance and drift. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 319–334.     

Beyond the pollination syndrome: nectar ecology and the role of diurnal and nocturnal pollinators in the reproductive success of Inga sessilis (Fabaceae)

Inga species present brush‐type flower morphology allowing them to be visited by distinct groups of pollinators. Nectar features in relation to the main pollinators have seldom been studied in this genus. To test the hypothesis of floral adaptation to both diurnal and nocturnal pollinators, we studied the pollination ecology of Inga sessilis, with emphasis on the nectar secretion patterns, effects of sequential removals on nectar production, sugar composition and the role of diurnal and nocturnal pollinators in its reproductive success. Inga sessilis is self‐incompatible and pollinated by hummingbirds, hawkmoths and bats. Fruit set under natural conditions is very low despite the fact that most stigmas receive polyads with sufficient pollen to fertilise all ovules in a flower. Nectar secretion starts in the bud stage and flowers continually secreting nectar for a period of 8 h. Flowers actively reabsorbed the nectar a few hours before senescence. Sugar production increased after nectar removal, especially when flowers were drained during the night. Nectar sugar composition changed over flower life span, from sucrose‐dominant (just after flower opening, when hummingbirds were the main visitors) to hexose‐rich (throughout the night, when bats and hawkmoths were the main visitors). Diurnal pollinators contributed less than nocturnal ones to fruit production, but the former were more constant and reliable visitors through time. Our results indicate I. sessilis has floral adaptations, beyond the morphology, that encompass both diurnal and nocturnal pollinator requirements, suggesting a complementary and mixed pollination system.     

Climatic influences on fire regimes along a rain forest-to-xeric woodland gradient in northern Patagonia, Argentina

ABSTRACT. Influences of annual climatic variation on fire occurrence were examined along a rainfall gradient from temperate rainforest to xeric woodlands in northern Patagonia, Argentina. Fire chronologies were derived from fire scars on trees and related to tree-ring proxy records of climate over the period 1820–1974. Similarly, fire records of four Patagonian national parks for the period 1940–1988 were compared to instrumental weather data. Finally, the influences of broad-scale synoptic weather patterns on fire occurrence in northern Patagonia were explored.
Fire in Nothofagus rainforests is highly dependent on drought during the spring and summer of the same year in which fires occur and is less strongly favoured by drought during the spring of the previous year. The occurrence of fire in dry vegetation types near the steppe ecotone is less dependent on drought because even during years of normal weather fuels are thoroughly desiccated during the dry summer. In xeric Austrocedrus woodlands, fire occurrence and spread are promoted by droughts during the fire season and also appear to be favoured by above-average moisture conditions during the preceding 1 to 2 growing seasons which enhances fuel production. Thus, in the xeric woodlands fire is not simply dependent on drought but is favoured by greater climatic variability over time scales of several years.
Fire activity in northern Patagonia is greatly influenced by the intensity and latitudinal position of the subtropical high pressure cell of the southeast Pacific. Greater fire activity is associated with a more intense and more southerly located high pressure cell which blocks the influx of Pacific moisture into the continent. Although long-term changes in fire occurrence along the rainforest-to-xeric woodland gradient have been greatly influenced by human activities, annual variation in fire frequency and extent is also strongly influenced by annual climatic variation.     

Post‐pollination process in a partially self‐compatible distylous plant,Primula sieboldii (Primulaceae)

Pollen germination and pollen‐tube growth under natural conditions were observed in a population of a distylous species, Primula sieboldii, in which partial self‐compatibility has been demonstrated in some long‐styled genets. We observed post‐pollination processes microscopically in styles collected after self‐morph and inter‐morph hand pollination (with standardized pollen load on the stigmas) in four genets each from the following three ‘genet types’: self‐incompatible long‐styled (SI), partially self‐compatible long‐styled (SC) and self‐incompatible short‐styled morph genets. Irrespective of the genet type, pollen germination began within 24 h after pollination and tubes of pollen reached to the style base with 48–96 h after inter‐morph pollination. Although pollen tubes germinated after self‐pollination in the SC genets, the number of germinated pollen tubes was significantly lower than in the case of inter‐morph pollination. Few pollen tubes germinated after self‐pollination of the SI or short‐styled genets. In SC genets, the rate of pollen‐tube growth did not differ between self‐morph and inter‐morph pollination (～1.9 mm/day). Therefore, differences in self‐compatibility between SC and SI genets in P. sieboldii are likely to be attributable to differential pollen germination rates rather than to differential pollen‐tube growth rates.     

The bracts of the alpine ‘glasshouse’ plant Rheum alexandrae (Polygonaceae) enhance reproductive fitness of its pollinating seed‐consuming mutualist

Environmental conditions are known to affect the reproductive fitness of insects, as do host plants. Although the highly specialized bracts of Himalayan ‘glasshouse’ plants are thought to be an adaptive morphology that allows the plant to reproduce in harsh alpine environments, little information appears to exist concerning the benefit of these bracts for the pollinating insects. In this study, we examine whether the semi‐translucent bracts of Rheum alexandrae, a giant herb endemic to the alpine zone of the Hengduan Mountains in south‐western China, enhance the reproductive fitness of its pollinator. Rheum alexandrae depends mainly on mutualistic seed‐consuming Bradysia flies for pollination. Bracts increased interior temperature on sunny days, acted as a buffer against fluctuating air humidity, greatly decreased the intensities of ultraviolet B/C radiation and provided shelter from strong winds. Bract removal significantly decreased adult oviposition and offspring performance of pollinators during flowering and fruiting, respectively. Our results indicate that the bracts of R. alexandrae enhance the reproductive fitness of its pollinating seed‐consuming mutualist in alpine environments, which may be attributed to the positive effects of the bracts on the interior microenvironment. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 349–359.     

Fire and flood: Soil‐stored seed bank and germination ecology in the endangered Carrington Falls Grevillea (Grevillea rivularis,Proteaceae)

Abstract Seed germination is dependent on the interaction between the dormancy state of a seed and the presence of favourable environmental conditions. Thus, the spectacular pulse of seedling recruitment in many Australian vegetation communities following disturbances such as fire can be attributed to changes in microsite conditions and/or the dormancy‐breaking effect of the disturbance on accumulated seed banks. Grevillea rivularis is a threatened species endemic to the area immediately above Carrington Falls in the NSW Southern Highlands. Most of the population is confined to the riparian vegetation zone in woodland and heath, and is therefore subject to periodic disturbance from fire and flood. For this species, a pulse of seedling recruitment has been recorded after fire, flood and mechanical soil disturbance. The aims of this study were to examine the density and vertical distribution of the soil‐stored seed bank and to investigate the role of heat and scarification as cues for germination of fresh and soil‐stored seed. There was a large seed bank under the canopies of established individuals (194 ± 73 seeds m^?2) and most seeds were found in the 0–2 cm and leaf‐litter layers of the soil profile. The germination response of soil‐stored and fresh seed was examined using a hierarchical series of laboratory experiments. Seeds of G. rivularis showed marked dormancy polymorphism. Thirty‐six percent of soil‐stored seed germinated without treatment, whereas no untreated fresh seeds germinated. Scarification or heating caused significant germination of dormant soil‐stored seed, but only scarification resulted in germination of dormant fresh seeds. These results highlight important differences in the dormancy state of soil‐stored and fresh seed. Thus, being a riparian species in a fire‐prone environment, the dormancy mechanisms in seeds of G. rivularis suit this species to disturbance by both fire and flood.     

Reproductive biology and pollination of Utricularia reniformis A.St.‐Hil. (Lentibulariaceae)

Utricularia reniformis is an endemic Brazilian carnivorous plant, most common in high‐altitude grasslands. Knowledge of the reproductive biology of U. reniformis is essential for planning conservation strategies, but it is currently poorly understood. Thus, we studied the floral morphology, floral biology, breeding system and pollination of this species. U. reniformis produces and stores nectar in the flower spur, a classic feature of bee‐pollinated flowers, and we recorded Xylocopa sp. and Bombus sp. as pollinators. Moreover, although it is self‐compatible it is an obligate animal‐pollinated species, as the sensitive stigma avoids self‐pollination. However, in natural conditions reproductive success is low due to the rarity of visits from pollinators. We suggest that the low reproductive success caused by a deficit of pollinators may affect gene flow, causing loss of genetic diversity in U. reniformis.     

TheMoegistorhynchus longirostris (Diptera: Nemestrinidae) pollination guild: long-tubed flowers and a specialized long-proboscid fly pollination system in southern Africa

A guild of 20 late spring- and early summer-flowering species ofIridaceae, Geraniaceae andOrchidaceae is pollinated partly or exclusively by the long-proboscid flyMoegistorhynchus longirostris (Nemestrinidae). This large-bodied fly, active in late spring and early summer, has mouthparts 40–70 (90) mm long and forages for nectar from a variety of species. These plants share a suite of convergent floral features including a straight or weakly curved floral tube usually 50–70 mm long but sometimes to 90 mm, relatively short petals or tepal lobes coloured white, cream or salmon with reddish nectar guides, and often violet or red anthers and pollen. Flowers of most species with these characteristics are zygomorphic with the stamens either arcuate (mostIridaceae) or declinate (Geraniaceae and someIridaceae). The flowers are odourless and typically secrete large amounts of nectar of relatively constant sugar concentration, mostly 24–29%, with a high sucrose:hexose ratio. Guild members utilize five separate sites of pollen deposition on the body of the fly, typically utilizing different deposition sites when two or more co-occur, indicating strong selection to aviod pollen contamination.M. longirostris is restricted to the west coast of southern Africa and at least 8 species appear to depend exclusively on the insect for pollination. The remaining species in the guild are pollinated by one or both of the long-proboscid fliesPhiloliche gulosa andP. rostrata (Tabanidae) over other parts of their range. Species and races pollinated entirely byM. longirostris have longer floral tubes which makes nectar unavailable to other insects, including other species of long-proboscid fly. The only insect with mouthparts long enough to forage effectively on these long-tubed flowers isM. longirostris and this fly must be considered a keystone species in the ecosystems in which it occurs.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

‘Black’ and ‘white’ death: burials in a time of Ebola in Freetown,Sierra Leone

The article examines experiences of the 2014‐15 Ebola crisis in Freetown, Sierra Leone, through an analysis of the performance of burials. While most of the city's residents had no contact with the virus, ‘Ebola’ was inescapable, owing to the onerous state of emergency regulations imposed by national and international authorities. All burials, regardless of the cause of death, were to be performed by newly established official teams operating according to unfamiliar biomedical and bureaucratic protocols. Burials became emblematic of the crisis through presenting a conflict between local practices and novel procedures, which was coded locally in a complex racial language of ‘black’ and ‘white’, recalling a long regional history of violent integration into the Atlantic World. Building on long‐standing anthropological discussion on the relationship between ‘good’ death and social order, the article explores how burials became sites around which opposing ‘orders’ were experienced, negotiated, and reconciled in locally meaningful ways.     

Pollen tube growth in the self‐compatible sweet cherry genotype, ‘Cristobalina’, is slowed down after self‐pollination

Sweet cherry is a self‐incompatible fruit tree species in the Rosaceae. As other species in the family, sweet cherry exhibits S‐RNase‐based gametophytic self‐incompatibility. This mechanism is genetically determined by the S‐locus that encodes the pollen and pistil determinants, SFB and S‐RNase, respectively. Several self‐compatible sweet cherry genotypes have been described and most of them have mutations at the S‐locus leading to self‐compatibility. However, ‘Cristobalina’ sweet cherry is self‐compatible due to a mutation in a pollen function modifier that is not linked to the S‐locus. To investigate the physiology of self‐compatibility in this cultivar, S‐locus segregation in crosses involving ‘Cristobalina’ pollen, and pollen tube growth in self‐ and cross‐pollinations, were studied. In the crosses with genotypes sharing only one S‐haplotype, the non‐self S‐haplotype was inherited more frequently than the self S‐haplotype. Pollen tube growth studies revealed that the time to travel the whole length of the style was longer for self‐pollen tubes than for cross‐pollen tubes. Together, these results suggest that ‘Cristobalina’ pollen tube growth is slower after self‐pollination than after cross‐pollination. This reproductive strategy would allow self‐fertilisation in the absence of compatible pollen but would promote cross‐fertilisation if cross‐compatible pollen is available, a possible case of cryptic self‐incompatibility. This bet‐hedging strategy might be advantageous for an ecotype that is native to the mountains of the Spanish Mediterranean coast, in the geographical limits of the distribution of this species. ‘Cristobalina’ blooming takes place very early in the season when mating possibilities are scarce and, consequently, self‐compatibility may be the only possibility for this genotype to produce offspring.     

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.     

A new pollination system: dung-beetle pollination discovered in Orchidantha inouei (Lowiaceae, Zingiberales) in Sarawak, Malaysia

Lowiaceae, a family of the Zingiberales, comprise 11 species in the single genus Orchidantha. Here we present the first report on the pollination of Lowiaceae and describe a new system of dung-beetle pollination from Sarawak, Borneo. Orchidantha inouei has a zygomorphic flower located just above the ground. Observations revealed that the plant is visited frequently and is pollinated by scarabaeid dung beetles, mainly members of the genus Onthophagus. All four species of Onthophagus collected on O. inouei have also been caught using traps baited with dung or carrion in Borneo. Onthophagus was presumably attracted to the dung-like odor of the flower. Pollination of O. inouei is different from other examples of beetle pollination in that its flower provides neither reward nor protected space. Dung beetles are excellent at following a particular dung scent. Orchidantha is the only genus that includes species lacking floral nectar. It is interesting that this deception pollination using dung beetles was found in Zingiberales, in which all known species have mutual and specialized relationships with their long-distance, but costly, pollinators-bees, birds, and bats.