Author Correction: Pollinator importance networks illustrate the crucial value of bees in a highly speciose plant community.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Pollinator importance networks illustrate the crucial value of bees in a highly speciose plant community.

Accurate predictions of pollination service delivery require a comprehensive understanding of the interactions between plants and flower visitors. To improve measurements of pollinator performance underlying such predictions, we surveyed visitation frequency, pollinator effectiveness (pollen deposition ability) and pollinator importance (the product of visitation frequency and effectiveness) of flower visitors in a diverse Mediterranean flower meadow. With these data we constructed the largest pollinator importance network to date and compared it with the corresponding visitation network to estimate the specialisation of the community with greater precision. Visitation frequencies at the community level were positively correlated with the amount of pollen deposited during individual visits, though rarely correlated at lower taxonomic resolution. Bees had the highest levels of pollinator effectiveness, with Apis, Andrena, Lasioglossum and Osmiini bees being the most effective visitors to a number of plant species. Bomblyiid flies were the most effective non-bee flower visitors. Predictions of community specialisation (H2') were higher in the pollinator importance network than the visitation network, mirroring previous studies. Our results increase confidence in existing measures of pollinator redundancy at the community level using visitation data, while also providing detailed information on interaction quality at the plant species level.

Constructing more informative plant-pollinator networks: visitation and pollen deposition networks in a heathland plant community.

Interaction networks are widely used as tools to understand plant-pollinator communities, and to examine potential threats to plant diversity and food security if the ecosystem service provided by pollinating animals declines. However, most networks to date are based on recording visits to flowers, rather than recording clearly defined effective pollination events. Here we provide the first networks that explicitly incorporate measures of pollinator effectiveness (PE) from pollen deposition on stigmas per visit, and pollinator importance (PI) as the product of PE and visit frequency. These more informative networks, here produced for a low diversity heathland habitat, reveal that plant-pollinator interactions are more specialized than shown in most previous studies. At the studied site, the specialization index [Formula: see text] was lower for the visitation network than the PE network, which was in turn lower than [Formula: see text] for the PI network. Our study shows that collecting PE data is feasible for community-level studies in low diversity communities and that including information about PE can change the structure of interaction networks. This could have important consequences for our understanding of threats to pollination systems.

Estimation of thermal constants: the importance of using equilibrium temperature rather than ambient temperature demonstrated with hoverflies (Diptera, Syrphidae, genus Eristalis).

Water evaporation has a marked effect on the passive rates of body temperature change of eristaline hoverflies. It results in the equilibrium temperature of these flies being significantly lower than ambient temperature. Different values for the cooling and warming constants are therefore obtained depending on whether equilibrium or ambient temperature is used as the baseline. Hence, care must be taken when estimating these constants with all animals, especially those of moderate to high permeability. It is recommended that equilibrium temperature be used in such situations. Evaporative cooling is probably also responsible for cooling constants being higher than warming constants in this and other studies.

The roles of colour change and behaviour in the hygrothermal balance of a littoral isopod, Ligia oceanica.

Ligia oceanica can change its colour using melanophores, the animal's reflectance varying between about 2 and 10%. Darker individuals heat up more quickly, and to higher body temperatures, than do pale ones. Colour change shows an underlying circadian rhythm, though the pattern of this rhythm varies with temperature, humidity, background and time of year. In general the rhythm is such as to ensure maximum camouflage at the critical dusk period, but in some conditions hygrothermal needs are overriding and the animals are paler (to stay cool) or darker (to warm up). In addition, animals show short term colour modification; when transferred to differing backgrounds and temperatures their colours initially reflect background matching, but after 30-45 min are modified into thermally appropriate shades, dark at 5° C and pale at 20° C. Field-caught specimens showed body temperatures that varied with colouration, and modification of colour in relation to thermal needs, particularly by being paler than expected when forced into the open by daytime high tides, and darker than expected when active prior to dusk. Animals invariably selected dark backgrounds in choice chambers. However, choice of humidity depended on previous experience; saturated air was normally preferred, but warm animals chose drier air (to allow evaporative cooling) unless also water-stressed. They also tended to disperse to facilitate cooling, whereas aggregation increased with increasing RH and with decreasing temperature. The interactions of colour changes, behavioural choices, and activity patterns in controlling the hygrothermal belance of Ligia in the intertidal environment are discussed in the light of these results.

The role of insect water balance in pollination ecology: Xylocopa and Calotropis.

Two carpenter bees (Xylocopa spp.) in southern Israel both use the asclepiad Calotropis procera as a primary nectar source. This plant genus is coevolved with carpenter bees, and aspects of the insect-flower interaction in Israel suggest that the smaller bee, X. sulcatipes, is the natural co-adapted pollinator, a view borne out by the geographical distributions of the species concerned. There are significant mismatches between the plant and the larger X. pubescens, involving physical fit and behaviour. These mismatches are particularly evident when the physiologies of the bees and the plant are considered. The different sizes and colours of the two bees lead to different daily activity patterns, only X. sulcatipes being thermally suited to, and thus abundant at, times of maximum nectar production by Calotropis. Similarly the water requirements of X. sulcatipes are finely balanced with the water production in the floral nectar; this bee gains just enough water when foraging to restore its blood concentration and production in the floral nectar; for deposition in the nest. X. pubescens does not incur net water loss in flight and gains too much water from Calotropis flowers, necessitating copious urination and 'tonguelashing'. Hence physiological information can be of use in deciphering insect-plant coevolutionary patterns, and the water component of nectar is confirmed as a potentially major determinant of foraging activities. The circumstances where this will be particularly true, and cases where it may not apply, are discussed.

Cellulose: a biopolymeric sorbent for heavy-metal traces in waters.

The sorption of dissolved heavy-metal traces on natural cellulose is characterized for the lower mug l . range. It proceeds relatively rapidly, with half-times of about 1.5 min. At neutral pH-values the distribution coefficients (K(d)) of many cations, of various types, e.g., Al(3+), Be(2+), Cd(2+), Cr(3+), Fe(3+), Pb(3+), Zn(2+), between cellulose and electrolyte solutions (e.g., Nad) are of the order 10(1)-10(4). In alkaline salt solutions K(d) as high as 10(5) can be attained. The role of cellulose as a sorbent for metal traces in natural waters is discussed.

Thermoregulatory mechanisms in Sarcophaga.

The flesh fly, Sarcophaga, is frequently seen feeding on flowers during periods of high radiation when other flies of comparable size avoid exposure because of the dangers of overheating. Sarcophaga is able to maintain its intermittent flower visits due to a cuticle of high thermal reflectance, giving low intrinsic heating rates, and to an ability to shunt blood between thorax and abdomen according to its needs. The fly thus achieves partial thermoregulation and can keep its body temperature within the preferred range for longer periods than its potential entomophilous competitors.

The nectar of Justicia and Columnea: Composition and concentration in a humid tropical climate.

Studies in Costa Rica on two ornithophilous flowers, Justicia aurea (Acanthaceae) and Columnea glabra (Gesneriaceae) showed a constancy of nectar solute concentrations that was attributed to microclimatic protection by the tubular corolla and to copious nectar secretion, helped by waterproofing by a lipid film on the nectar surface in Justicia and by preferential compass orientation of the flowers of Columnea.Most of the corollas in the patch of Justicia had been pierced by nectar-robbers. A consequence of this damage, together with local microclimate effects, was flower-to-flower variation in the amount and accessibility of nectar and in the nature and concentration of its minor components, notably amino acids.McDade and Kinsman's (1980) finding that nectar secretion could be suppressed by repeated sampling or by nectar-robbing was confirmed.

Temporal and microclimatic partitioning of the floral resources of Justicia aurea amongst a concourse of pollen vectors and nectar robbers.

The flowers of Justicia aurea, morphologically characteristic of ornithophily, attracted a diverse array of foragers where they occurred as a dense stand in the tropical forests at La Selva, Costa Rica, and so provided an arena for this study of competition and coexistence. Two hummingbird species (Phaethornis superciliosus and Campylopterus hemileucurus) visited the flowers legally early in the morning, and defended the nectar resource; a third smaller bird (P. longuemareus) foraged for nectar throughout the day but collected it illegally by piercing the corollas. In addition, nectar was harvested illegally by four species of stingless bee (Trigona) and by ants, creating a further drain on the limited floral resources.Consideration of the diurnal patterns of foraging activities in combination with a spatial axis (defined here in terms of microclimate and insolation) nevertheless showed a good separation of flower visits for the different nectarivores. Hummingbirds visited flowers in zones where the reward was highest, while insects foraged to minimise their energetic costs; each of these factors could in turn be related to microclimatic considerations. A comprehensive scheme of resource utilisation could therefore be extracted from the field observations and interpreted in these terms. The limited area of niche overlap thus revealed corresponded closely with the situations where direct interference competition was observed, between hummingbird species or between bees and aggressive ants.

Field analyses of insect heat budgets: Reflectance, size and heating rates.

This paper outlines simple techniques for determining rates of heat gain and loss in relation to the weight and reflectance of insects caught in their natural habitats. In particular the construction of a new 'reflectometer' is described. The results thus obtained permit estimates of the relative importance of size and colour in determining rates of heat exchange and temperature excesses, so allowing better predictions of heat budgets for a given species.

The effects of insect visitors on nectar constituents in temperate plants.

The floral nectar of angiosperms is primarily a solution of simple sugars, but contains detectable amounts of other solutes, particularly amino acids. These have been regarded as diagnostic, for phylogenetic and taxonomic purposes, and their mean concentrations may be significant in relation to the pollination syndrome of a plant species. However, in several temperate flowers the amino acid concentration varied by an order of magnitude when measured at intervals through a single day. In open, cup-shaped flowers, this was partly due to post-secretory equilibration with the varying humidity of the air. But the effects of floral visitation by insects on nectar amino acids were also important, both in open flowers and in those with long corollas and more enclosed nectar. Visitors could add amino acids to nectar by direct contact, by salivation, by damaging the neighbouring tissues causing cell leakage, and by dislodging pollen into the nectar; the importance of each of these effects varied according to corolla morphology and the spectrum of visitors.

Physiologically induced changes in intramembranous particle frequency in the axons of an osmoconforming bivalve.

Freeze-fractured axonal membrane surfaces from the connectives of Mytilus edulis show an increment in particle frequency of 52% (fixed tissues) or 68% (unfixed tissues) after long-term adaptation to low salinity. Particle size distribution was unaffected by osmotic adaptation, but was significantly different in fixed and unfixed material. The possibility that these structural changes reflect the known increase in sodium pump frequency in this osmoconforming tissue is considered.

Hormonal control of integumentary water-loss: evidence for a novel neuroendocrine system in an insect (Periplaneta americana).

An accelerated water-loss was observed in decapitated individuals, no equivalent increase being obtained following severance of the nervous connectives in the neck. Injection of brain and, to a lesser extent, corpus cardiacum extract resulted in a significant reduction in the rate of loss of water from decapitated individuals. The accelerated water-loss observed following decapitation appeared not to result from significant increase in excretory output or loss of water through the spiracles. It is suggested that integumentary transpiration may be affected by a blood-borne factor, or factors, which originate in the brain and corpus cardiacum.