Long-proboscid Fly Research Articles

Evidence for moth pollination in a rhinomyiophilous Erica species from the Cape Floristic Region of South Africa.

Contrasting pollination syndromes in closely related species suggest that floral trait divergence is associated with differences in pollination system, but empirical observations are required to confirm syndrome-based predictions. We present a comparative study of two closely related Erica species with contrasting pollination syndromes from the Cape Floristic Region of South Africa. Ericacylindrica has narrowly tubular pale and strongly scented flowers and is known to be hawkmoth-pollinated. The closely related Ericainfundibuliformis has bright flower colours and appears to lack scent, traits that are suggestive of pollination by long-tongued nemestrinid flies (rhinomyiophily). Floral trait measurements revealed that both species exhibit predominantly upright flower orientation and elongated floral tubes, although tube length of E.infundibuliformis is consistently greater than that of E.cylindrica. For both species, petals are brighter than floral tube surfaces, but flowers of E.cylindrica lack the strong UV reflectance found in E.infundibuliformis. Nectar of E.infundibuliformis is more concentrated and produced in larger volumes. Scent composition, but not evening scent emission rates, differed between the species: scent of E.cylindrica is dominated by aromatic compounds, whereas scent of E.infundibuliformis is dominated by (E)-ocimene and other terpenoid compounds and is emitted at higher rates during the day than the evening. Pollinator observations contradicted trait-based predictions: although a single nemestrinid fly captured in the vicinity of E.infundibuliformis did carry Erica pollen, almost all other diurnal flower visitors were nectar-robbing Hymenoptera which did not carry Erica pollen. Contrary to predictions, at two sites and over two flowering seasons, flowers were consistently visited in the evenings by several species of settling moths and hawkmoths which carried pollen, almost exclusively of Erica, on their proboscides. Our findings thus suggest that, despite objective differences in key floral traits between the closely related hawkmoth-pollinated E.cylindrica and E.infundibuliformis, moths are also important pollinators of E.infundibuliformis. A bimodal pollination system involving predominant pollination by moths and occasional visits by long-proboscid flies could partially reconcile findings with predictions. Our study further suggests that hawkmoth pollination may be more widespread in both Erica and the broader Cape flora than has hitherto been assumed and emphasises the importance of nocturnal pollinator observations.

Pollen transfer efficiency in Erica depends on type of pollinator.

Pollen transfer efficiency (PTE; the proportion of pollen removed from flowers that reaches conspecific stigmas) is expected to vary with the type of pollinator and flower morphology, and to influence male siring success. Many species in the genus Erica are pollinated by bees (which consume pollen and should thus lower PTE) but during its radiation in the Cape, several independent shifts to both sunbird and long-proboscid fly (LP fly) pollinators, which do not consume pollen have taken place. Improvements in PTE could be one of the factors driving these pollinator shifts. PTE data for 15 Erica species (five for each of the three pollinator types) were collected and compared in relation to type of pollinator and anther exsertion. LP fly- and bird-pollinated species had higher PTE in comparison with bee-pollinated species. Species with inserted anthers had higher PTE than those with exserted anthers. This suggests that sunbirds and LP flies are more efficient pollinators than bees. Additionally, the study suggests that insertion of anthers within the corolla tube can reduce pollen losses.

Flower orientation and corolla length as reproductive barriers in the pollinator-driven divergence of Erica shannonea and Erica ampullacea.

A variety of reproductive barriers can enable reproductive isolation and stable coexistence of plant species. Differing floral traits might play an important role in reproductive isolation imposed by pollinators. Such shifts in pollinator use have been hypothesized to contribute to the radiation of Erica (Ericaceae) in the Cape Floristic Region, South Africa. The sister species Erica shannonea and Erica ampullacea co-occur and overlap in flowering phenology. Both have unscented long-tubed flowers consistent with adaptations for pollination by long-proboscid flies (LPFs), but differences in flower orientation and corolla tube length are indicative of a shift in pollinator species. We conducted controlled pollination experiments and pollinator observations to determine the breeding system and pollinators of the two species. Both species are self-incompatible and require pollinator visits for seed production, suggesting that pollinators could strongly influence flower evolution. The horizontally orientated flowers of E. shannonea were found to be pollinated by Philoliche rostrata (Tabanidae), which has a long, fixed forward-pointing proboscis, while the vertically upright orientated flowers of E. ampullacea were found to be pollinated by Prosoeca westermanni (Nemestrinidae), which has a shorter proboscis that can swivel downwards. The nemestrinid fly's proboscis is too short to access the nectar in the relative long-tubed flowers of E. shannonea and the tabanid fly's proboscis cannot swivel down to access the upright flowers of E. ampullacea. Consequently, these traits are likely to act as reproductive barriers between the two Erica species and thereby might have contributed to speciation and enable stable coexistence.

Biomechanics of nectar feeding explain flower orientation in plants pollinated by long-proboscid flies.

The function of flower orientation is much debated, with adaptation to pollinator mouthparts being a particularly compelling explanation, but also one that has lacked empirical support from broad-scale comparative studies. The two families of long-proboscid fly pollinators show similar hovering behaviour while feeding on nectar but differ in the biomechanics of their proboscides which can be up to 80mm in length: Tabanidae have a fixed forward-pointing proboscis while Nemestrinidae can swivel their proboscis downwards. We predicted that this difference has implications for the evolution of flower orientation. We established the flower angles of 156 South African plant species specialised for pollination by long-proboscid flies. Using a phylogenetically corrected analysis, we found that flowers pollinated by Tabanidae tend to be horizontally orientated, while those pollinated by Nemestrinidae tend to be more variable in orientation and more often vertically orientated. These results confirm the importance of pollinator biomechanics for the evolution of floral traits and highlight a potential mechanism of reproductive isolation between sympatric plant species pollinated by different long-proboscid fly families.

Sending private messages: Floral ultraviolet signals are correlated with pollination syndromes in Erica

The presence of ultraviolet (UV, wavelengths between 300-400 nm) reflectance in insect-pollinated flowers has been linked to pollination efficiency and pollination shifts, but little is known about its prevalence and function in other pollination systems and African species. We chose the genus Erica for studying the prevalence of UV because of its extreme radiation (c. 680 species) in the Cape, South Africa, with a diversity of pollination syndromes. This study quantified the prevalence and brightness of UV reflectance for five Erica pollination syndromes and tested pollinator preferences for UV reflectance in the two groups with the highest prevalence: sunbirds and long-proboscid flies. Our results show that UV colouration is absent or rare in Erica species pollinated by unclassified insects, rodents or wind. About 17 % of bird-pollinated species reflected UV but choice experiments revealed that free-ranging sunbirds showed no preference for UV signals. All sampled long-proboscid fly-pollinated species reflected UV and its experimental removal decreased seed set drastically, suggesting that long-proboscid flies in the Cape strongly prefer or depend on UV and thereby contributed to selecting for the evolution of this signal.

Pollination biology of Erica aristata: First confirmation of long-proboscid fly-pollination in the Ericaceae

Long-proboscid fly (LPF) pollination has been recorded in multiple plant lineages in South Africa. However in the large and morphologically diverse genus Erica, evidence for LPF pollination is limited to predictions based upon floral syndromes as well as anecdotal observation and limited citizen science records. Here we use breeding system experiments, pollinator exclusion, remote camera traps, spectral analysis and field observation in the Klein River Mountains, Western Cape Province, South Africa, to demonstrate this mode of pollination in the range-restricted taxon Erica aristata. This species has long narrow corolla tubes with narrow opening and contrasting white and crimson pink petals. Flowers were visited and pollinated exclusively by one long-proboscid fly species, Prosoeca rubicunda (Nemestrinidae) over four years at this study site. Breeding experiments showed that E. aristata is not self-compatible and pollinator exclusion experiments indicate that pollinator visits were required for seed-set. Low nectar volume and moderately high nectar sugar concentration were consistent with other long-proboscid fly-pollinated plant species in the Cape. This study represents the first confirmed instance of long-proboscid fly-pollination in the genus Erica.

Corolla stickiness prevents nectar robbing in Erica.

Floral stickiness is a rare trait with unknown function, but it is common in the mega-diverse Cape genus Erica (Ericaceae). This study investigated the role of stickiness measured as adhesive strength in Erica as protection against nectar robbing and its correlation with floral traits. We compared the incidence of nectar robbing in flowers of the same species with or without experimentally added stickiness, and amongst communities of co-occurring species with flowers differing in stickiness. Additionally, we tested the relationship between stickiness and pollination syndrome, corolla shape, corolla length and sepal-corolla ratio across the whole genus. Stickiness was correlated with lower floral damage rates within and between species, indicating it functions as an anti-nectar robbing trait. Across the genus Erica, stickiness is most strongly correlated with bird and long-proboscid fly pollination, presumably because of their larger nectar rewards. Stickiness was also correlated with floral traits that are often associated with high risk of being damaged by nectar robbers: narrow-mouthed corollas, long corollas and shorter sepals. These results show that corolla stickiness defends Erica flowers against nectar robbing and thereby potentially improves fitness.

First long-proboscid flies (Diptera: Zhangsolvidae) from the Lower Cretaceous of South Korea

Khramov, A.V., Nam, G.-S. & Vasilenko, D.V. 13 December 2019. First long-proboscid flies (Diptera: Zhangsolvidae) from the Lower Cretaceous of South Korea. Alcheringa 44, 160–168. ISSN 0311-5518 Buccinatormyia gangnami sp. nov. (Zhangsolvidae) is the first record of Diptera from the Lower Cretaceous Jinju Formation of the Korean Peninsula. B. gangnami is described based on six specimens, the largest number of fossils known for a single species of Zhangsolvidae. Four specimens of B. gangnami preserve a long proboscis; mouthparts of this species show a twofold variation in relative length, which may be partially explained by sexual dimorphism. B. gangnami is the third known species of Zhangsolvidae with extremely elongate antennal flagella. The preserved abdominal colouration pattern in these fossils suggests that B. gangnami was probably a wasp mimic. Alexander V. Khramov [a-hramov@yandex.ru], Borissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117997 Moscow, Russian Federation; Gi-Soo Nam [nks33@naver.com], Gongju National University of Education, 27, Ungjin-ro, Gongju-si, Chungcheongnam-do, 32553, South Korea; Dmitry V. Vasilenko* [damageplant@mail.ru], Borissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117997 Moscow, Russian Federation. *Also affiliated with: Cherepovets State University, Lunacharskogo 5, 162600 Cherepovets, Russian Federation.

Intraspecific divergence in floral-tube length promotes asymmetric pollen movement and reproductive isolation.

The causative link between phenotypic divergence and reproductive isolation is an important but poorly understood part of ecological speciation. We studied the effects of floral-tube length variation on pollen placement/receipt positions and reproductive isolation. In a population of Lapeirousia anceps (Iridaceae) with bimodal floral-tube lengths, we labelled pollen of short- and long-tubed flowers with different colour fluorescent nanoparticles (quantum dots). This enabled us to map pollen placement by long- and short-tubed flowers on the only floral visitor, a long-proboscid fly. Furthermore, it allowed us to quantify pollen movement within and betweenshort- and long-tubed flowers. Short- and long-tubed flowers placed pollen on different parts of the pollinator, and long-tubed flowers placed more pollen per visit than short-tubed flowers. This resulted in assortative pollen receipt (most pollen received comes from the same phenotype) and strong but asymmetric reproductive isolation, where short-tubed plants are more reproductively isolated than long-tubed plants. These results suggest that floral-tube length divergence can promote mechanical isolation in plants through divergence in pollen placement sites on pollinators. Consequently, in concert with other reproductive isolation mechanisms, selection for differences in floral-tube length can play an important role in ecological speciation of plants.

Importance of birds versus insects as pollinators of the African shrub Syncolostemon densiflorus (Lamiaceae)

Flowers visited by a broad assemblage of animals may nevertheless be considered to have a specialized pollination system if pollen deposition is effected mainly by a subset of these visitors. The importance of a particular animal group for pollination derives mainly from the relative rate of floral visitation and per-visit effectiveness for pollen deposition. We assessed the relative importance of birds and insects for pollination of the African shrub Syncolostemon densiflorus (Lamiaceae). Flower visitors were observed to evaluate their visitation rates. Virgin flowers were exposed to single visits by nine animal visitor groups to evaluate their effectiveness at depositing pollen on stigmas. We selectively excluded birds and larger insects from flowers to assess the contribution of smaller visitors to pollen deposition and seed set. Sunbirds and insects visited flowers at a similar frequency, but on a per-visit basis, sunbirds deposited more pollen on stigmas than did any of the insect groups, including honeybees, long-proboscid flies, butterflies and day-flying hawkmoths. Selective exclusion of birds and larger insects resulted in a large decline in pollen deposition on stigmas and seed production. These results show that S. densiflorus is pollinated mainly by sunbirds, despite its attraction of a broad assemblage of insect flower visitors. These findings highlight the importance of per-visit pollination effectiveness and selective exclusion for understanding the ecological importance of different flower visitors.

Pollination, mating and reproductive fitness in a plant population with bimodal floral-tube length.

Mating patterns and natural selection play important roles in determining whether genetic polymorphisms are maintained or lost. Here, we document an atypical population of Lapeirousia anceps (Iridaceae) with a bimodal distribution of floral-tube length and investigate the reproductive mechanisms associated with this pattern of variation. Flowers were visited exclusively by the long-proboscid fly Moegistorhynchus longirostris (Nemestrinidae), which exhibited a unimodal distribution of proboscis length and displayed a preference for long-tubed phenotypes. Despite being visited by a single pollinator species, allozyme markers revealed significant genetic differentiation between open-pollinated progeny of long- and short-tubed phenotypes suggesting mating barriers between them. We obtained direct evidence for mating barriers between the floral-tube phenotypes through observations of pollinator foraging, controlled hand pollinations and measurements of pollen competition and seed set. Intermediate tube-length phenotypes produced fewer seeds in the field than either long- or short-tubed phenotypes. Although floral-tube length bimodality may not be a stable state over long timescales, reproductive barriers to mating and low 'hybrid' fitness have the potential to contribute to the maintenance of this state in the short term.

Long-Proboscid Flies as Pollinators of Cretaceous Gymnosperms

The great evolutionary success of angiosperms has traditionally been explained, in part, by the partnership of these plants with insect pollinators. The main approach to understanding the origins of this pervasive relationship has been study of the pollinators of living cycads, gnetaleans, and basal angiosperms. Among the most morphologically specialized living pollinators are diverse, long-proboscid flies. Early such flies include the brachyceran family Zhangsolvidae, previously known only as compression fossils from the Early Cretaceous of China and Brazil. It belongs to the infraorder Stratiomyomorpha, a group that includes the flower-visiting families Xylomyidae and Stratiomyidae. New zhangsolvid specimens in amber from Spain (ca. 105 mega-annum [Ma]) and Myanmar (100 Ma) reveal a detailed proboscis structure adapted to nectivory. Pollen clumped on a specimen from Spain is Exesipollenites, attributed to a Mesozoic gymnosperm, most likely the Bennettitales. Late Mesozoic scorpionflies with a long proboscis have been proposed as specialized pollinators of various extinct gymnosperms, but pollen has never been observed on or in their bodies. The new discovery is a very rare co-occurrence of pollen with its insect vector and provides substantiating evidence that other long-proboscid Mesozoic insects were gymnosperm pollinators. Evidence is thus now gathering that visitors and probable pollinators of early anthophytes, or seed plants, involved some insects with highly specialized morphological adaptations, which has consequences for interpreting the reproductive modes of Mesozoic gymnosperms and the significance of insect pollination in angiosperm success.

A new long-tubed subspecies of <i>Pelargonium dipetalum</i> (section <i>Hoarea</i>) (Geraniaceae) from the Albertinia-Swellendam area in Western Cape Province, South Africa

Background: Field studies confirmed that unusually long-tubed populations of Pelargonium dipetalum from between Swellendam and Albertinia, Western Cape Province, South Africa, are a distinct ecotype adapted to pollination by the long-proboscid fly, Prosoeca longipennis. The geographical and morphological isolation of these populations suggests that they are reproductively isolated from short-tubed populations, which are pollinated by bees.Objectives: To determine and describe the floral variation in P. dipetalum, with a view to recognising the long-tubed populations at some taxonomic level.Method: All available collections were measured and compared.Results: Populations of P. dipetalum were segregated into a short-tubed form with hypanthium 3 mm – 24 mm long and mostly pink petals that occurs from Betty’s Bay to Knysna, and a long-tubed form with the hypanthium 34 mm – 54 mm long and consistently white petals that is restricted to a small area east of Swellendam between Suurbraak and Albertinia. We described the long-tubed form as the new subspecies P. dipetalum subsp. stenosiphon.Conclusion: The new subspecies increases our understanding of the diversity in P. dipetalum and represents a new taxon of conservation concern.

Floral divergence in closely related Leucospermum tottum (Proteaceae) varieties pollinated by birds and long-proboscid flies

The Proteaceae are renowned for their floral diversity but surprisingly the role of pollinators in driving evolutionary divergence in this family has been underexplored. Here we focus on recently diverged taxa to gain insight into the processes that generate diversity by testing whether two varieties of Leucospermum tottum might have originated by pollinator mediated adaptive divergence. L. tottum var. tottum has pale salmon-coloured horizontally-oriented flowers, long nectar tubes, and small volumes of concentrated nectar. L. tottum var. glabrum has red and yellow vertically oriented flowers, short nectar tubes, and large volumes of dilute nectar. Despite the morphological divergence, the varieties are indistinguishable using eight molecular markers, indicating a very early stage of differentiation. Consistent with their morphologies, L. tottum var. tottum is pollinated by long-proboscid flies (Philoliche rostrata and Philoliche gulosa), Cape sugarbirds (Promerops cafer), and, to a lesser extent, by Orange-breasted sunbirds (Anthobaphes violacea), whereas, L. tottum var. glabrum is pollinated only by Orange-breasted sunbirds. A. violacea visits both varieties, but makes more frequent contact with pollen presenters when foraging on L. tottum var. glabrum. The exclusion of birds caused a steeper reduction in seed production in L. tottum var. glabrum than in L. tottum var. tottum, consistent with specialization for bird-pollination in this variety. Additionally, L. tottum var. glabrum exhibits autogamy, whereas L. tottum var. tottum does not. Floral divergence between the two L. tottum varieties corresponds with divergence in pollinator use.

Time management and nectar flow: flower handling and suction feeding in long-proboscid flies (Nemestrinidae: Prosoeca)

A well-developed suction pump in the head represents an important adaptation for nectar-feeding insects, such as Hymenoptera, Lepidoptera and Diptera. This pumping organ creates a pressure gradient along the proboscis, which is responsible for nectar uptake. The extremely elongated proboscis of the genus Prosoeca (Nemestrinidae) evolved as an adaptation to feeding from long, tubular flowers. According to the functional constraint hypothesis, nectar uptake through a disproportionately elongated, straw-like proboscis increases flower handling time and consequently lowers the energy intake rate. Due to the conspicuous length variation of the proboscis of Prosoeca, individuals with longer proboscides are hypothesised to have longer handling times. To test this hypothesis, we used field video analyses of flower-visiting behaviour, detailed examinations of the suction pump morphology and correlations of proboscis length with body length and suction pump dimensions. Using a biomechanical framework described for nectar-feeding Lepidoptera in relation to proboscis length and suction pump musculature, we describe and contrast the system in long-proboscid flies. Flies with longer proboscides spent significantly more time drinking from flowers. In addition, proboscis length and body length showed a positive allometric relationship. Furthermore, adaptations of the suction pump included an allometric relationship between proboscis length and suction pump muscle volume and a combination of two pumping organs. Overall, the study gives detailed insight into the adaptations required for long-proboscid nectar feeding, and comparisons with other nectar-sucking insects allow further considerations of the evolution of the suction pump in insects with sucking mouthparts.Electronic supplementary materialThe online version of this article (doi:10.1007/s00114-013-1114-6) contains supplementary material, which is available to authorized users.

Matching floral and pollinator traits through guild convergence and pollinator ecotype formation.

Pollinator landscapes, as determined by pollinator morphology/behaviour, can vary inter- or intraspecifically, imposing divergent selective pressures and leading to geographically divergent floral ecotypes. Assemblages of plants pollinated by the same pollinator (pollinator guilds) should exhibit convergence of floral traits because they are exposed to similar selective pressures. Both convergence and the formation of pollination ecotypes should lead to matching of traits among plants and their pollinators. We examined 17 floral guild members pollinated in all or part of their range by Prosoeca longipennis, a long-proboscid fly with geographic variation in tongue length. Attractive floral traits such as colour, and nectar properties were recorded in populations across the range of each species. The length of floral reproductive parts, a mechanical fit trait, was recorded in each population to assess possible correlation with the mouthparts of the local pollinator. A multiple regression analysis was used to determine whether pollinators or abiotic factors provided the best explanation for variation in floral traits, and pollinator shifts were recorded in extralimital guild member populations. Nine of the 17 species were visited by alternative pollinator species in other parts of their ranges, and these displayed differences in mechanical fit and attractive traits, suggesting putative pollination ecotypes. Plants pollinated by P. longipennis were similar in colour throughout the pollinator range. Tube length of floral guild members co-varied with the proboscis length of P. longipennis. Pollinator shifts have resulted in geographically divergent pollinator ecotypes across the ranges of several guild members. However, within sites, unrelated plants pollinated by P. longipennis are similar in the length of their floral parts, most probably as a result of convergent evolution in response to pollinator morphology. Both of these lines of evidence suggest that pollinators play an important role in selecting for certain floral traits.

ADAPTIVE DIVERGENCE IN DARWIN'S RACE: HOW COEVOLUTION CAN GENERATE TRAIT DIVERSITY IN A POLLINATION SYSTEM

Understanding how reciprocal selection shapes interacting species in Darwin's coevolutionary race is a captivating pursuit in evolutionary ecology. Coevolving traits can potentially display following three patterns: (1) geographical variation in matched traits, (2) bias in trait matching, and (3) bimodal distribution of a trait in certain populations. Based on the framework of adaptive dynamics, we present an evolutionary model for a coevolving pollination system involving the long-proboscid fly (Moegistorhynchus longirostris) and the long-tubed iris (Lapeirousia anceps). The model successfully demonstrates that Darwin's hypothesis can lead to all three patterns if costs are involved. Geographical variation in matched traits could be driven by geographical variation in environmental factors that affect the cost rate of trait escalation. Unequal benefits derived from the interaction by the fly and the flower could potentially cause the bias in trait matching of the system. Different cost rates to trait elongation incurred by the two species and weak assortative interactions in the coevolutionary race can drive divergent selection (i.e., an evolutionary branching) that leads to the bimodal distribution of traits. Overall, the model highlights the importance of assortative interactions and the balance of costs incurred by coevolving species as factors determining the eventual phenotypic outcome of coevolutionary interactions.

Preliminary observations of insect pollination in Protea punctata (Proteaceae)

Available information on the phylogeny and pollination systems in Protea suggests that bird-pollination is ancestral and occurs in the majority of species, and that there have been several shifts to rodent pollination and a single shift to pollination by cetoniine beetles in this large African genus. Here we report that Protea punctata plants in a population in the Swartberg mountains are pollinated primarily by long-proboscid flies and butterflies. The threadlike pollen presenter on an unusually flexible style facilitates insect pollination in this species. The length of the style matches that of the proboscides of its two most common visitors, the nemestrinid fly Prosoeca longipennis and the nymphalid butterfly Aeropetes tulbaghia. Spectral reflectance of the involucral bracts is similar to that of flowers of other plants visited by long-proboscid flies. P. punctata occupies a recently diverged position in a clade (the “white proteas”) in which all the other species appear to be bird-pollinated, and may represent a shift to insect pollination in the genus.

Competition versus facilitation: conspecific effects on pollinator visitation and seed set in the iris Lapeirousia oreogena

Ecological interactions between conspecific plants can range from facilitative to competitive depending on the spacing and abundance of individuals. Competitive interactions are expected when plants flower en‐masse and availability of pollinators is limited. We tested this prediction using Lapeirousia oreogena, a mass‐flowering South African iris that is pollinated by a single species of long‐proboscid fly. Controlled hand‐pollination experiments showed that L. oreogena is self‐compatible, but reliant on pollinator visits for seed set. Seed production per flower showed a significant negative relationship with patch size (and the correlated measure of number of individuals per patch), but was not affected by flower density or distance to neighbouring patches. There was a tendency for fly abundance to increase with patch size, but the rate of visits to individual flowers by flies was not affected by patch size. Seed set of hand‐pollinated flowers did not differ for plants in and out of dense patches, indicating that the large differences in seed set among patches were likely to reflect pollinator visits, rather than the genetic or physiological capacity of plants to produce seeds. The reduced fecundity of L. oreogena in large patches with a greater numbers of flowers is consistent with the idea that plants with highly specialized pollination systems can experience intra‐specific competition for pollination.

Diverse pollination systems of the twin-spurred orchid genus Satyrium in African grasslands

The large terrestrial orchid genus Satyrium underwent evolutionary radiations in the Cape floral region and the grasslands of southern and eastern Africa. These radiations were accompanied by tremendous diversification of the unusual twin-spurred flowers that characterize the genus, but pollination data required to interpret these patterns of floral evolution have been lacking for grassland species in the genus. Here we document pollinators, nectar properties, and levels of pollination success for 11 grassland Satyrium species in southern and south-central Africa. Pollinators of these species include bees, beetles, butterflies, hawkmoths, noctuid moths, long-proboscid flies, and sunbirds. Most species appear to be specialized for pollination by one functional pollinator group. Long-proboscid fly pollination systems are reported for the first time in Satyrium (in S. macrophyllum and a high-altitude form of S. neglectum). Floral morphology, especially spur length and rostellum structure, differs markedly among plants with different pollinators, while nectar volume, concentration, and sugar composition are fairly uniform across species. Most taxa exhibited high levels of pollination success (>50% of flowers pollinated), a trend that can be attributed to the presence of nectar in the twin spurs.