Personality variation is eroded by simple social behaviours in collective foragers.

The movement of groups can be heavily influenced by 'leader' individuals who differ from the others in some way. A major source of differences between individuals is the repeatability and consistency of their behaviour, commonly considered as their 'personality', which can influence both position within a group as well as the tendency to lead. However, links between personality and behaviour may also depend upon the immediate social environment of the individual; individuals who behave consistently in one way when alone may not express the same behaviour socially, when they may be conforming with the behaviour of others. Experimental evidence shows that personality differences can be eroded in social situations, but there is currently a lack of theory to identify the conditions where we would expect personality to be suppressed. Here, we develop a simple individual-based framework considering a small group of individuals with differing tendencies to perform risky behaviours when travelling away from a safe home site towards a foraging site, and compare the group behaviours when the individuals follow differing rules for aggregation behaviour determining how much attention they pay to the actions of their fellow group-members. We find that if individuals pay attention to the other members of the group, the group will tend to remain at the safe site for longer, but then travel faster towards the foraging site. This demonstrates that simple social behaviours can result in the repression of consistent inter-individual differences in behaviour, giving the first theoretical consideration of the social mechanisms behind personality suppression.

The role of petal transpiration in floral humidity generation.

MAIN CONCLUSION: Using petrolatum gel as an antitranspirant on the flowers of California poppy and giant bindweed, we show that transpiration provides a large contribution to floral humidity generation. Floral humidity, an area of elevated humidity in the headspace of flowers, is believed to be produced predominantly through a combination of evaporation of liquid nectar and transpirational water loss from the flower. However, the role of transpiration in floral humidity generation has not been directly tested and is largely inferred by continued humidity production when nectar is removed from flowers. We test whether transpiration contributes to the floral humidity generation of two species previously identified to produce elevated floral humidity, Calystegia silvatica and Eschscholzia californica. Floral humidity production of flowers that underwent an antitranspirant treatment, petrolatum gel which blocks transpiration from treated tissues, is compared to flowers that did not receive such treatments. Gel treatments reduced floral humidity production to approximately a third of that produced by untreated flowers in C. silvatica, and half of that in E. californica. This confirms the previously untested inferences that transpiration has a large contribution to floral humidity generation and that this contribution may vary between species.

Bumblebees can detect floral humidity.

Floral humidity, a region of elevated humidity in the headspace of the flower, occurs in many plant species and may add to their multimodal floral displays. So far, the ability to detect and respond to floral humidity cues has been only established for hawkmoths when they locate and extract nectar while hovering in front of some moth-pollinated flowers. To test whether floral humidity can be used by other more widespread generalist pollinators, we designed artificial flowers that presented biologically relevant levels of humidity similar to those shown by flowering plants. Bumblebees showed a spontaneous preference for flowers that produced higher floral humidity. Furthermore, learning experiments showed that bumblebees are able to use differences in floral humidity to distinguish between rewarding and non-rewarding flowers. Our results indicate that bumblebees are sensitive to different levels of floral humidity. In this way floral humidity can add to the information provided by flowers and could impact pollinator behaviour more significantly than previously thought.

Cross-modal transfer in visual and nonvisual cues in bumblebees.

Bumblebees Bombus terrestris are good at learning to distinguish between patterned flowers. They can differentiate between flowers that differ only in their patterning of scent, surface texture, temperature, or electrostatic charge, in addition to visual patterns. As recently shown, bumblebees trained to discriminate between nonvisual scent patterns can transfer this learning to visually patterned flowers that show similar spatial patterning to the learnt scent patterns. Bumblebees can, therefore, transfer learnt patterns between different sensory modalities, without needing to relearn them. We used differential conditioning techniques to explore whether cross-modal transfer of learnt patterns also occurred between visual and temperature patterns. Bumblebees that successfully learnt to distinguish rewarding and unrewarding temperature patterns did not show any preferences for the corresponding unlearnt visual pattern. Similarly, bumblebees that learnt visual patterns did not transfer these to temperature patterns, suggesting that they are unable to transfer learning of temperature and visual patterns. We discuss how cross-modality pattern learning may be limited to modalities that have potentially strong neurological links, such as the previously demonstrated transfer between scent and visual patterns.

Bumblebees distinguish floral scent patterns, and can transfer these to corresponding visual patterns.

Flowers act as multisensory billboards to pollinators by using a range of sensory modalities such as visual patterns and scents. Different floral organs release differing compositions and quantities of the volatiles contributing to floral scent, suggesting that scent may be patterned within flowers. Early experiments suggested that pollinators can distinguish between the scents of differing floral regions, but little is known about how these potential scent patterns might influence pollinators. We show that bumblebees can learn different spatial patterns of the same scent, and that they are better at learning to distinguish between flowers when the scent pattern corresponds to a matching visual pattern. Surprisingly, once bees have learnt the spatial arrangement of a scent pattern, they subsequently prefer to visit novel unscented flowers that have an identical arrangement of visual marks, suggesting that multimodal floral signals may exploit the mechanisms by which learnt information is stored by the bee.

Assessing the seasonal prevalence and risk factors for nuchal crest adiposity in domestic horses and ponies using the Cresty Neck Score.

BACKGROUND: Nuchal crest adiposity in horses and ponies has been associated with an enhanced risk of metabolic health problems. However, there is no current information on the prevalence of, and risk factors specific to, nuchal crest adiposity in horses and ponies. In addition, the cresty neck score has not previously been utilised across different seasons within a UK leisure population, it is not know whether nuchal crest adiposity shows the same seasonal trends as general obesity. RESULTS: A Cresty Neck Score (CNS, 0-5) was given to 96 horses with access to pasture (>6 h per day) at the end of winter and at the end of summer in order to obtain two prevalence estimates. Risk factors were assessed using the single outcome cresty neck/no cresty neck in either season (binary), from owner questionnaires and analysed using a mixed effects logistic regression model (outcome variable CNS <3 or CNS ≥3/5). Agreement between winter and summer scores was assessed using weighted Kappa methods. Winter CNS values were significantly higher than summer CNS values (p = 0.002) indicating a systematic bias. The prevalence of a CNS ≥ 3/5 was 45.83% at the end of winter, falling to 33.33% at the end of summer and was higher in ponies (<14.2 hh) than horses (≥14.2 hh) in both seasons. This may reflect a real winter increase in regional fat deposition, or an increased difficulty in obtaining an accurate estimate of regional adiposity in winter months. Breed was the strongest risk factor for CNS ≥3/5 in both seasons, with native UK breeds appearing to be most at risk (p < 0.001). In a separate, small validation study, the CNS showed good inter-observer reliability. CONCLUSIONS: The prevalence of a CNS ≥3/5 was higher at the end of winter than at the end of summer, which was the opposite pattern seasonal variation to that observed for general obesity. Further studies are required to investigate the potential influence of time of year upon CNS interpretation and studies utilising the CNS should consider potential seasonal variability in nuchal crest adiposity.

Dominance rank is associated with body condition in outdoor-living domestic horses (Equus caballus).

The aim of our study was to explore the association between dominance rank and body condition in outdoor group-living domestic horses, Equus caballus. Social interactions were recorded using a video camera during a feeding test, applied to 203 horses in 42 herds. Dominance rank was assigned to 194 individuals. The outcome variable body condition score (BCS) was recorded using a 9-point scale. The variables age and height were recorded and considered as potential confounders or effect modifiers. Results were analysed using multivariable linear and logistic regression techniques, controlling for herd group as a random effect. More dominant (p = 0.001) individuals generally had a higher body condition score (p = 0.001) and this association was entirely independent of age and height. In addition, a greater proportion of dominant individuals fell into the obese category (BCS ≥ 7/9, p = 0.005). There were more displacement encounters and a greater level of interactivity in herds that had less variation in age and height, lending strength to the hypothesis that phenotypic variation may aid cohesion in group-living species. In addition there was a strong quadratic relationship between age and dominance rank (p < 0.001), where middle-aged individuals were most likely to be dominant. These results are the first to link behavioural predictors to body condition and obesity status in horses and should prompt the future consideration of behavioural and social factors when evaluating clinical disease risk in group-living animals.

The behavioural responses of bumblebees Bombus terrestris to simulated rain.

Bumblebee activity typically decreases during rainfall, putting them under the threat of the increased frequency of precipitation due to climate change. A novel rain machine was used within a flight arena to observe the behavioural responses of bumblebees (Bombus terrestris) to simulated rain at both a colony and individual level. During rainfall, a greater proportion of workers left the arena than entered, the opposite of which was seen during dry periods, implying that they compensate for their lack of activity when conditions improve. The proportion of workers flying and foraging decreased while resting increased in rain. This pattern reversed during dry periods, providing further evidence for compensatory activity. The increase in resting behaviour during rain is thought to evade the high energetic costs of flying while wet without unnecessarily returning to the nest. This effect was not repeated in individual time budgets, measured with lone workers, suggesting that the presence of conspecifics accelerates the decision of their behavioural response, perhaps via local enhancement. Bumblebees probably use social cues to strategize their energetic expenditure during precipitation, allowing them to compensate for the reduced foraging activity during rainfall when conditions improve.

The cuticular wax composition and crystal coverage of leaves and petals differ in a consistent manner between plant species.

Both leaves and petals are covered in a cuticle, which itself contains and is covered by cuticular waxes. The waxes perform various roles in plants' lives, and the cuticular composition of leaves has received much attention. To date, the cuticular composition of petals has been largely ignored. Being the outermost boundary between the plant and the environment, the cuticle is the first point of contact between a flower and a pollinator, yet we know little about how plant-pollinator interactions shape its chemical composition. Here, we investigate the general structure and composition of floral cuticular waxes by analysing the cuticular composition of leaves and petals of 49 plant species, representing 19 orders and 27 families. We show that the flowers of plants from across the phylogenetic range are nearly devoid of wax crystals and that the total wax load of leaves in 90% of the species is higher than that of petals. The proportion of alkanes is higher, and the chain lengths of the aliphatic compounds are shorter in petals than in leaves. We argue these differences are a result of adaptation to the different roles leaves and petals play in plant biology.

Variations of floral temperature in changing weather conditions.

Floral temperature is a flower characteristic that has the potential to impact the fitness of flowering plants and their pollinators. Likewise, the presence of floral temperature patterns, areas of contrasting temperature across the flower, can have similar impacts on the fitness of both mutualists. It is currently poorly understood how floral temperature changes under the influence of different weather conditions, and how floral traits may moderate these changes. The way that floral temperature changes with weather conditions will impact how stable floral temperatures are over time and their utility to plants and pollinators. The stability of floral temperature cues is likely to facilitate effective plant-pollinator interactions and play a role in the plant's reproductive success. We use thermal imaging to monitor how floral temperatures and temperature patterns of four plant species (Cistus 'snow fire' and 'snow white', Coreopsis verticillata and Geranium psilostemon) change with several weather variables (illumination, temperature; windspeed; cloud cover; humidity and pressure) during times that pollinators are active. All weather variables influenced floral temperature in one or more species. The directionality of these relationships was similar across species. In all species, light conditions (illumination) had the greatest influence on floral temperatures overall. Floral temperature and the extent to which flowers showed contrasting temperature patterns were influenced predominantly by light conditions. However, several weather variables had additional, lesser, influences. Furthermore, differences in floral traits, pigmentation and structure, likely resulted in differences in temperature responses to given conditions between species and different parts of the same flower. However, floral temperatures and contrasting temperature patterns that are sufficiently elevated for detection by pollinators were maintained across most conditions if flowers received moderate illumination. This suggests the presence of elevated floral temperature and contrasting temperature patterns are fairly constant and may have potential to influence plant-pollinator interactions across weather conditions.

The influence of pigmentation patterning on bumblebee foraging from flowers of Antirrhinum majus.

Patterns of pigmentation overlying the petal vasculature are common in flowering plants and have been postulated to play a role in pollinator attraction. Previous studies report that such venation patterning is significantly more attractive to bee foragers in the field than ivory or white flowers without veins. To dissect the ways in which venation patterning of pigment can influence bumblebee behaviour, we investigated the response of flower-naïve individuals of Bombus terrestris to veined, ivory and red near-isogenic lines of Antirrhinum majus. We find that red venation shifts flower colour slightly, although the ivory background is the dominant colour. Bees were readily able to discriminate between ivory and veined flowers under differential conditioning but showed no innate preference when presented with a free choice of rewarding ivory and veined flowers. In contrast, both ivory and veined flowers were selected significantly more often than were red flowers. We conclude that advantages conferred by venation patterning might stem from bees learning of their use as nectar guides, rather than from any innate preference for striped flowers.

Red deer Cervus elaphus blink more in larger groups.

Most animals need to spend time being vigilant for predators, at the expense of other activities such as foraging. Group-living animals can benefit from the shared vigilance effort of other group members, with individuals reducing personal vigilance effort as group size increases. Behaviors like active scanning or head lifting are usually used to quantify vigilance but may not be accurate measures of this. We suggest that measuring an animal's blinking rate gives a meaningful measure of vigilance: increased blinking implies reduced vigilance, as the animal cannot detect predators when its eyes are closed. We describe an observational study of a captive population of red deer, where we measured the blinking rates of individual deer from groups of differing sizes (where mean group size ranged between 1 and 42.7 individuals). We demonstrate that as group size increases in red deer, individuals increase their blink rate, confirming the prediction that vigilance should decrease. Blinking is a simple non-invasive measure and offers a useful metric for assessing the welfare of animals experiencing an increase in perceived predation risk or other stressors.

Multimodal floral recognition by bumblebees.

Flowers present information to their insect visitors in multiple simultaneous sensory modalities. Research has commonly focussed on information presented in visual and olfactory modalities. Recently, focus has shifted towards additional 'invisible' information, and whether information presented in multiple modalities enhances the interaction between flowers and their visitors. In this review, we highlight work that addresses how multimodality influences behaviour, focussing on work conducted on bumblebees (Bombus spp.), which are often used due to both their learning abilities and their ability to use multiple sensory modes to identify and differentiate between flowers. We review the evidence for bumblebees being able to use humidity, electrical potential, surface texture and temperature as additional modalities, and consider how multimodality enhances their performance. We consider mechanisms, including the cross-modal transfer of learning that occurs when bees are able to transfer patterns learnt in one modality to an additional modality without additional learning.

The effects of dominance on leadership and energetic gain: a dynamic game between pairs of social foragers.

Although social behaviour can bring many benefits to an individual, there are also costs that may be incurred whenever the members of a social group interact. The formation of dominance hierarchies could offer a means of reducing some of the costs of social interaction, but individuals within the hierarchy may end up paying differing costs dependent upon their position within the hierarchy. These differing interaction costs may therefore influence the behaviour of the group, as subordinate individuals may experience very different benefits and costs to dominants when the group is conducting a given behaviour. Here, a state-dependent dynamic game is described which considers a pair of social foragers where there is a set dominance relationship within the pair. The model considers the case where the subordinate member of the pair pays an interference cost when it and the dominant individual conduct specific pairs of behaviours together. The model demonstrates that if the subordinate individual pays these energetic costs when it interacts with the dominant individual, this has effects upon the behaviour of both subordinate and the dominant individuals. Including interaction costs increases the amount of foraging behaviour both individuals conduct, with the behaviour of the pair being driven by the subordinate individual. The subordinate will tend to be the lighter individual for longer periods of time when interaction costs are imposed. This supports earlier suggestions that lighter individuals should act as the decision-maker within the pair, giving leadership-like behaviours that are based upon energetic state. Pre-existing properties of individuals such as their dominance will be less important for determining which individual makes the decisions for the pair. This suggests that, even with strict behavioural hierarchies, identifying which individual is the dominant one is not sufficient for identifying which one is the leader.

The Ability of Bumblebees Bombus terrestris (Hymenoptera: Apidae) to Detect Floral Humidity is Dependent Upon Environmental Humidity.

Flowers produce local humidity that is often greater than that of the surrounding environment, and studies have shown that insect pollinators may be able to use this humidity difference to locate and identify suitable flowers. However, environmental humidity is highly heterogeneous, and is likely to affect the detectability of floral humidity, potentially constraining the contexts in which it can be used as a salient communication pathway between plants and their pollinators. In this study, we use differential conditioning techniques on bumblebees Bombus terrestris audax (Harris) to explore the detectability of an elevated floral humidity signal when presented against different levels of environmental noise. Artificial flowers were constructed that could be either dry or humid, and individual bumblebees were presented with consistent rewards in either the humid or dry flowers presented in an environment with four levels of constant humidity, ranging from low (~20% RH) to highly saturated (~95% RH). Ability to learn was dependent upon both the rewarding flower type and the environment: the bumblebees were able to learn rewarding dry flowers in all environments, but their ability to learn humid rewarding flowers was dependent on the environmental humidity, and they were unable to learn humid rewarding flowers when the environment was highly saturated. This suggests that floral humidity might be masked from bumblebees in humid environments, suggesting that it may be a more useful signal to insect pollinators in arid environments.

Phylogenetically-controlled correlates of primate blinking behaviour.

Eye blinking is an essential maintenance behaviour for many terrestrial animals, but is also a risky behaviour as the animal is unable to scan the environment and detect hazards while its eyes are temporarily closed. It is therefore likely that the length of time that the eyes are closed and the length of the gap between blinks for a species may reflect aspects of the ecology of that species, such as its social or physical environment. An earlier published study conducted a comparative study linking blinking behaviour and ecology, and detailed a dataset describing the blinking behaviour of a large number of primate species that was collected from captive animals, but the analysis presented did not control for the nonindependence of the data due to common evolutionary history. In the present study, the dataset is reanalysed using phylogenetic comparative methods, after reconsideration of the parameters describing the physical and social environments of the species. I find that blink rate is best described by the locomotion mode of a species, where species moving through arboreal environments blink least, ground-living species blink most, and species that use both environments show intermediate rates. The duration of a blink was also related to locomotion mode, and positively correlated with both mean species group size and mean species body mass, although the increase in relation to group size is small. How a species moves through the environment therefore appears to be important for determining blinking behaviour, and suggests that complex arboreal environments may require less interruption to visual attention. Given that the data were collected with captive individuals, caution is recommended for interpreting the correlations found.

Floral infrared emissivity estimates using simple tools.

BACKGROUND: Floral temperature has important consequences for plant biology, and accurate temperature measurements are therefore important to plant research. Thermography, also referred to as thermal imaging, is beginning to be used more frequently to measure and visualize floral temperature. Accurate thermographic measurements require information about the object's emissivity (its capacity to emit thermal radiation with temperature), to obtain accurate temperature readings. However, there are currently no published estimates of floral emissivity available. This is most likely to be due to flowers being unsuitable for the most common protocols for emissivity estimation. Instead, researchers have used emissivity estimates collected on vegetative plant tissue when conducting floral thermography, assuming these tissues to have the same emissivity. As floral tissue differs from vegetative tissue, it is unclear how appropriate and accurate these vegetative tissue emissivity estimates are when they are applied to floral tissue. RESULTS: We collect floral emissivity estimates using two protocols, using a thermocouple and a water bath, providing a guide for making estimates of floral emissivity that can be carried out without needing specialist equipment (apart from the thermal camera). Both protocols involve measuring the thermal infrared radiation from flowers of a known temperature, providing the required information for emissivity estimation. Floral temperature is known within these protocols using either a thermocouple, or by heating the flowers within a water bath. Emissivity estimates indicate floral emissivity is high, near 1, at least across petals. While the two protocols generally indicated the same trends, the water bath protocol gave more realistic and less variable estimates. While some variation with flower species and location on the flower is observed in emissivity estimates, these are generally small or can be explained as resulting from artefacts of these protocols, relating to thermocouple or water surface contact quality. CONCLUSIONS: Floral emissivity appears to be high, and seems quite consistent across most flowers and between species, at least across petals. A value near 1, for example 0.98, is recommended for accurate thermographic measurements of floral temperature. This suggests that the similarly high values based on vegetation emissivity estimates used by previous researchers were appropriate.

Phylogenetic signal in floral temperature patterns.

OBJECTIVES: Floral structures may be warmer than their environment, and can show thermal patterning, where individual floral structures show different temperatures across their surface. Pollinators can differentiate between artificial flowers that mimic both naturally warmed and thermally patterned ones, but it has yet to be demonstrated that these patterns are biologically meaningful. To explore the relationship between pollinators and temperature patterning, we need to know whether there is diversity in patterning, and that these patterns are not simply a by-product of floral architecture constrained by ancestry. We analysed a dataset of 97 species to explore whether intrafloral temperature differences were correlated within clades (phylogenetic signal), or whether the variation seen was diverse enough to suggest that floral temperature patterns are influenced by the abiotic or pollinator-related niches to which plant species are adapted. RESULTS: Some phylogenetic signal was observed, with both the Asteraceae and species of Pelargonium being more similar than expected by chance, but with other species surveyed not showing signal. The Asteraceae tend to have large temperature differences across the floral surface, which may be due to floral architecture constraints within the family. Other families show no correlation, suggesting that patterning is influenced by pollinators and the environment.

Using radio frequency identification and locomotor activity monitoring to assess sleep, locomotor, and foraging rhythmicity in bumblebees.

Bumblebees are a key pollinator. Understanding the factors that influence the timing of sleep and foraging trips is important for efficient foraging and pollination. Here, we illustrate how individual locomotor activity monitoring and colony-wide radio frequency identification tracking can be combined to analyze the effects of agrochemicals like neonicotinoids on locomotor and foraging rhythmicity and sleep quantity/quality in bumblebees. We also highlight aspects of the design that can be adapted for other invertebrates or agrochemicals, allowing broader application of these techniques. For complete details on the use and execution of this protocol, please refer to Tasman et al. (2020).