Craig I. Peter

POLLINATION SUCCESS IN A DECEPTIVE ORCHID IS ENHANCED BY CO-OCCURRING REWARDING MAGNET PLANTS

It has been debated whether pollination success in nonrewarding plants that flower in association with nectar-producing plants will be diminished by competition for pollinator visits or, alternatively, enhanced through increased local abundance of pollinators (the magnet species effect). We experimentally evaluated these effects using the nonrewarding bumblebee-pollinated orchid Anacamptis morio and associated nectar-producing plants at a site in Sweden. Pollination success (estimated as pollen receipt and pollen removal) in A. morio was significantly greater for individuals translocated to patches of nectar-producing plants (Geum rivale and Allium schoenoprasum) than for individuals placed outside (∼20 m away) such patches. These results provide support for the existence of a facilitative magnet species effect in the interaction between certain nectar plants and A. morio. To determine the spatial scale of these interactions, we correlated the visitation rate to flowers of A. morio with the density of sym...

Comparing models for predicting species' potential distributions: a case study using correlative and mechanistic predictive modelling techniques

Abstract Models used to predict species’ potential distributions have been described as either correlative or mechanistic. We attempted to determine whether correlative models could perform as well as mechanistic models for predicting species potential distributions, using a case study. We compared potential distribution predictions made for a coastal dune plant ( Scaevola plumieri ) along the coast of South Africa, using a mechanistic model based on summer water balance (SWB), and two correlative models (a profile and a group discrimination technique). The profile technique was based on principal components analysis (PCA) and the group-discrimination technique was based on multiple logistic regression (LR). Kappa ( κ ) statistics were used to objectively assess model performance and model agreement. Model performance was calculated by measuring the levels of agreement (using κ ) between a set of testing localities (distribution records not used for model building) and each of the model predictions. Using published interpretive guidelines for the kappa statistic, model performance was “excellent” for the SWB model ( κ =0.852), perfect for the LR model ( κ =1.000), and “very good” for the PCA model ( κ =0.721). Model agreement was calculated by measuring the level of agreement between the mechanistic model and the two correlative models. There was “good” model agreement between the SWB and PCA models ( κ =0.679) and “very good” agreement between the SWB and LR models ( κ =0.786). The results suggest that correlative models can perform as well as or better than simple mechanistic models. The predictions generated from these three modelling designs are likely to generate different insights into the potential distribution and biology of the target organism and may be appropriate in different situations. The choice of model is likely to be influenced by the aims of the study, the biology of the target organism, the level of knowledge the target organism’s biology, and data quality.

MIMICS AND MAGNETS: THE IMPORTANCE OF COLOR AND ECOLOGICAL FACILITATION IN FLORAL DECEPTION

Plants that lack floral rewards can attract pollinators if they share attractive floral signals with rewarding plants. These deceptive plants should benefit from flowering in close proximity to such rewarding plants, because pollinators are locally conditioned on floral signals of the rewarding plants (mimic effect) and because pollinators are more abundant close to rewarding plants (magnet effect). We tested these ideas using the non-rewarding South African plant Eulophia zeyheriana (Orchidaceae) as a study system. Field observations revealed that E. zeyheriana is pollinated solely by solitary bees belonging to a single species of Lipotriches (Halictidae) that appears to be closely associated with the flowers of Wahlenbergia cuspidata (Campanulaceae), a rewarding plant with which the orchid is often sympatric. The pale blue color of the flowers of E. zeyheriana differs strongly from flowers of its congeners, but is very similar to that of flowers of W. cuspidata. Analysis of spectral reflectance patterns using a bee vision model showed that bees are unlikely to be able to distinguish the two species in terms of flower color. A UV-absorbing sunscreen was applied to the flowers of the orchid in order to alter their color, and this resulted in a significant decline in pollinator visits, thus indicating the importance of flower color for attraction of Lipotriches bees. Pollination success in the orchid was strongly affected by proximity to patches of W. cuspidata. This was evident from one of two surveys of natural populations of the orchid, as well as experiments in which we translocated inflorescences of the orchid either into patches of W. cuspidata or 40 m outside such patches. Flower color and location of E. zeyheriana plants relative to rewarding magnet patches are therefore key components of the exploitation by this orchid of the relationship between W. cuspidata and Lipotriches bee pollinators.

Doing the twist: A test of Darwin's cross-pollination hypothesis for pollinarium reconfiguration.

Mating success in plants depends largely on the efficiency of pollen dispersal. For hermaphrodite plants, self-pollination, either within or among flowers, can reduce mating opportunities because of pollen and ovule discounting and inbreeding depression. Self-pollination may be particularly detrimental in plants such as orchids and asclepiads that package each flowers pollen into one or more pollinia which, together with accessory structures, comprise a pollinarium. Darwin proposed that physical reconfiguration of pollinaria serves as a mechanism for reducing the likelihood of self-pollination. To be effective, the time taken for pollinarium reconfiguration would need to exceed that spent by a pollinator on a plant. We investigated pollinarium reconfiguration (including pollinarium bending, pollinium shrinking and anther cap retention) in 19 species and found a strong positive relationship between reconfiguration time and the duration of pollinator visits. Reconfiguration times were also consistently longer than pollinator visit times. These results provide strong support for Darwins idea that this mechanism promotes cross-pollination.

Reproductive biology of Acrolophia cochlearis (Orchidaceae): estimating rates of cross-pollination in epidendroid orchids

BACKGROUND AND AIMS Pollen fates strongly influence mating success in plants but are difficult to quantify. By promoting foraging constancy in pollinators, floral rewards such as nectar may enhance the overall efficiency of pollen transfer. However, this can also lead to high levels of geitonogamy. Pollen fates were studied in Acrolophia cochlearis, a member of a terrestrial epidendroid orchid genus that includes both rewarding and deceptive species. METHODS Pollinator observations were conducted. Pollen transfer efficiency (PTE), the proportion of removed pollinia deposited on stigmas, was measured in a large population at regular intervals throughout the 5-month flowering season. The level of cross-pollination in two populations was estimated from the percentage of seeds with embryos in naturally pollinated fruits. KEY RESULTS Acrolophia cochlearis (and a congener A. micrantha) produce minute but concentrated nectar rewards. Observations showed that A. cochlearis is pollinated exclusively by a solitary bee species, Colletes claripes. Although both sexes visited flowers, only males carried pollinaria. Overall levels of pollination and PTE of the rewarding A. cochlearis were much higher than in a deceptive congener, A. capensis. Seeds resulting from self-fertilization had a significantly lower probability of containing viable embryos than did those from cross-fertilization. This dichotomy in fruit quality was used to estimate that cross-pollination occurred in approx. 66 % of A. cochlearis flowers in a large dense population and approx. 10 % in a small sparse population. Traits of A. cochlearis that limit geitonogamy include pollinarium reconfiguration that exceeds the visit time of pollinators and rapid flower senescence following visitation. CONCLUSIONS Presence of a nectar reward in Acrolophia cochlearis results in high levels of PTE. It is estimated that approx. 33-90 % of fruits in natural populations arise from self-pollination in this species.

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.

The invasive 'mothcatcher' (Araujia sericifera Brot.; Asclepiadoideae) co-opts native honeybees as its primary pollinator in South Africa.

Araujia sericifera is a South American milkweed which is an invasive/exotic species in South Africa. This species relies on pollinators for sexual reproduction and we show it has co-opted native South African honeybees as primary pollinators in its adopted country. Moths also visit the flowers of this species, but contribute little to its relatively high pollination success.

Large pollen loads of a South African asclepiad do not interfere with the foraging behaviour or efficiency of pollinating honey bees

The pollen of asclepiads (Asclepiadoideae, Apocynaceae) and most orchids (Orchidaceae) are packaged as large aggregations known as pollinaria that are removed as entire units by pollinators. In some instances, individual pollinators may accumulate large loads of these pollinaria. We found that the primary pollinator of Cynanchum ellipticum (Apocynaceae—Asclepiadoideae), the honey bee Apis mellifera, accumulate very large agglomerations of pollinaria on their mouthparts when foraging on this species. We tested whether large pollinarium loads negatively affected the foraging behaviour and foraging efficiency of honey bees by slowing foraging speeds or causing honey bees to visit fewer flowers, and found no evidence to suggest that large pollinarium loads altered foraging behaviour. C. ellipticum displayed consistently high levels of pollination success and pollen transfer efficiency (PTE). This may be a consequence of efficiently loading large numbers of pollinaria onto pollinators even when primary points of attachment on pollinators are already occupied and doing so in a manner that does not impact the foraging behaviour of pollinating insects.

Environmental limits to the distribution of Scaevola plumieri along the South African coast

Scaevola plumieri is an important pioneer on many tropical and subtropical sand dunes, forming a large perennial subterranean plant with only the tips of the branches emerging above accreting sand. In South Africa it is the dominant pioneer on sandy beaches along the east coast, less abundant on the south coast and absent from the southwest and west coasts. Transpiration rates (E) of S. plumieri are predictably related to atmospheric vapour pressure deficit under a wide range of conditions and can therefore be predicted from measurement of ambient temperature and relative humidity. Scaling measurements of E at the leaf level to the canopy level has been demonstrated previously. Using a geographic information system, digital maps of regional climatic variables were used to calculate digital maps of potential transpiration from mean monthly temperature and relative humidity values, effectively scaling canopy level transpiration rates to a regional level. Monthly potential transpiration was subtracted from the monthly median rainfall to produce a map of mean monthly water balance. Seasonal growth was correlated with seasonal water balance. Localities along the coast with water deficits in summer corresponded with the recorded absence of S. plumieri, which grows and reproduces most actively in the summer months. This suggests that reduced water availability during the summer growth period limits the distribution of S. plumieri along the southwest coast, where water deficits develop in summer. Temperature is also important in limiting the distribution of S. plumieri on the southwest coast of South Africa through its effects on the growth and phenology of the plant.

A Temporal Dimension to the Influence of Pollen Rewards on Bee Behaviour and Fecundity in Aloe tenuior

The net effect of pollen production on fecundity in plants can range from negative – when self-pollen interferes with fecundity due to incompatibility mechanisms, to positive – when pollen availability is associated with increased pollinator visitation and fecundity due to its utilization as a reward. We investigated the responses of bees to pollen and nectar rewards, and the effects of these rewards on pollen deposition and fecundity in the hermaphroditic succulent shrub Aloe tenuior. Self-pollinated plants failed to set fruit, but their ovules were regularly penetrated by self-pollen tubes, which uniformly failed to develop into seeds as expected from ovarian self-incompatibility (or strong early inbreeding depression). Bees consistently foraged for pollen during the morning and early afternoon, but switched to nectar in the late afternoon. As a consequence of this differential foraging, we were able to test the relative contribution to fecundity of pollen- versus nectar-collecting flower visitors. We exposed emasculated and intact flowers in either the morning or late afternoon to foraging bees and showed that emasculation reduced pollen deposition by insects in the morning, but had little effect in the afternoon. Despite the potential for self-pollination to result in ovule discounting due to late-acting self-sterility, fecundity was severely reduced in artificially emasculated plants. Although there were temporal fluctuations in reward preference, most bee visits were for pollen rewards. Therefore the benefit of providing pollen that is accessible to bee foragers outweighs any potential costs to fitness in terms of gender interference in this species.