The Effect of Digestive Capacity on the Intake Rate of Toxic and Non-Toxic Prey in an Ecological Context

Digestive capacity often limits food intake rate in animals. Many species can flexibly adjust digestive organ mass, enabling them to increase intake rate in times of increased energy requirement and/or scarcity of high-quality prey. However, some prey species are defended by secondary compounds, thereby forcing a toxin limitation on the forager’s intake rate, a constraint that potentially cannot be alleviated by enlarging digestive capacity. Hence, physiological flexibility may have a differential effect on intake of different prey types, and consequently on dietary preferences. We tested this effect in red knots (Calidris canutus canutus), medium-sized migratory shorebirds that feed on hard-shelled, usually mollusc, prey. Because they ingest their prey whole and crush the shell in their gizzard, the intake rate of red knots is generally constrained by digestive capacity. However, one of their main prey, the bivalve Loripes lucinalis, imposes a toxin constraint due to its symbiosis with sulphide-oxidizing bacteria. We manipulated gizzard sizes of red knots through prolonged exposure to hard-shelled or soft foods. We then measured maximum intake rates of toxic Loripes versus a non-toxic bivalve, Dosinia isocardia. We found that intake of Dosinia exponentially increased with gizzard mass, confirming earlier results with non-toxic prey, whereas intake of Loripes was independent of gizzard mass. Using linear programming, we show that this leads to markedly different expected diet preferences in red knots that try to maximize energy intake rate with a small versus a large gizzard. Intra- and inter-individual variation in digestive capacity is found in many animal species. Hence, the here proposed functional link with individual differences in foraging decisions may be general. We emphasize the potential relevance of individual variation in physiology when studying trophic interactions.     

Reinterpretation of gizzard sizes of red knots world-wide emphasises overriding importance of prey quality at migratory stopover sites

The size of digestive organs can be rapidly and reversibly adjusted to ecological circumstances, but such phenotypic flexibility comes at a cost. Here, we test how the gizzard mass of a long-distance migrant, the red knot (Calidris canutus), is adjusted to (i) local climate, (ii) prey quality and (iii) migratory fuelling demands. For eight sites around the world (both wintering and stopover sites), we assembled data on gizzard masses of free-living red knots, the quality of their prey and the local climate. Using an energetic cost-benefit approach, we predicted the gizzard size required for fastest fuelling (net rate-maximization, i.e. expected during migration) and the gizzard size required to balance daily energy budgets (satisficing, expected in wintering birds) at each site. The measured gizzards matched the net rate-maximizing predictions at stopover sites and the satisficing predictions at wintering sites. To our surprise, owing to the fact that red knots selected stopover sites with prey of particularly high quality, gizzard sizes at stopovers and at wintering sites were nevertheless similar. To quantify the benefit of minimizing size changes in the gizzard, we constructed a model incorporating the size-dependent energy costs of maintaining and carrying a gizzard. The model showed that by selecting stopovers containing high-quality prey, metabolic rates are kept at a minimum, potentially reducing the spring migratory period by a full week. By inference, red knots appear to time their stopovers so that they hit local peaks in prey quality, which occur during the reproductive seasons of the intertidal benthic invertebrates.     

Phenotypic flexibility of body composition associated with seasonal acclimatization in passerine birds

Improved winter cold tolerance is widespread among small birds overwintering in cold climates and is associated with improved shivering endurance and elevated summit metabolic rate (M_sum). Phenotypic flexibility resulting in elevated M_sum could result from either increased skeletal muscle mass (perhaps with support from similar adjustments in “nutritional organs”) and/or cellular metabolic intensity. We investigated seasonal changes in body composition of three species of passerine birds resident in cold winter climates, all of which show large seasonal variations in M_sum (>25%); white-breasted nuthatch (Sitta carolinensis), black-capped chickadee (Poecile atricapillus), and house sparrow (Passer domesticus). All three species displayed significant winter increases in pectoralis and heart masses, and supracoracoideus mass also increased in winter chickadees. Gizzard mass increased in winter for all three species, but masses of other nutritional organs did not vary consistently with season. These data suggest that winter increases in pectoralis and heart masses are important contributors to elevated thermogenic capacity and cold tolerance, but seasonal variation in nutritional organ masses, other than gizzard, which is likely associated with dietary changes, are not universally associated with seasonal phenotypes. The winter increases in pectoralis and heart masses are consistent with data from other small passerines showing marked seasonal changes in cold tolerance and support the Variable Maximum Model of seasonal phenotypic flexibility, where physiological adjustments that promote improved cold tolerance, also result in elevated M_sum.     

Realized heritability and repeatability of risk-taking behaviour in relation to avian personalities

Personalities are general properties of humans and other animals. Different personality traits are phenotypically correlated, and heritabilities of personality traits have been reported in humans and various animals. In great tits, consistent heritable differences have been found in relation to exploration, which is correlated with various other personality traits. In this paper, we investigate whether or not risk-taking behaviour is part of these avian personalities. We found that (i) risk-taking behaviour is repeatable and correlated with exploratory behaviour in wild-caught hand-reared birds, (ii) in a bi-directional selection experiment on 'fast' and 'slow' early exploratory behaviour, bird lines tend to differ in risk-taking behaviour, and (iii) within-nest variation of risk-taking behaviour is smaller than between-nest variation. To show that risk-taking behaviour has a genetic component in a natural bird population, we bred great tits in the laboratory and artificially selected 'high' and 'low' risk-taking behaviour for two generations. Here, we report a realized heritability of 19.3 +/- 3.3% (s.e.m.) for risk-taking behaviour. With these results we show in several ways that risk-taking behaviour is linked to exploratory behaviour, and we therefore have evidence for the existence of avian personalities. Moreover, we prove that there is heritable variation in more than one correlated personality trait in a natural population, which demonstrates the potential for correlated evolution.     

How do red knots Calidris canutus leave Northwest Australia in May and reach the breeding grounds in June? Predictions of stopover times,fuelling rates and prey quality in the Yellow Sea

In general, Arctic-breeding waders leave non-breeding grounds in Australasia from March (New Zealand) to mid-April (Northwest Australia). Here we provide evidence from radio-tracking and visual observations that many red knots Calidris canutus do not leave Roebuck Bay, Northwest Australia, until early or mid-May. Late-departing red knots probably belong to the subspecies piersmai , which breeds on the New Siberian Islands, 10,400 km from Northwest Australia. Based on comparisons of temperatures on the breeding grounds of different knot subspecies, we predict that piersmai knots would not arrive on the breeding grounds until early June, leaving at most 3–4 weeks refuelling in Asia. Using a model of fuelling capacity in relation to prey quality and gizzard mass, we show that these knots must fuel very differently in Australia and Asia. In Australia, knots have seemingly suboptimal gizzard sizes and deposit fuel slowly. In the Yellow Sea, birds could only fuel up within the available time if they either enlarged their gizzards substantially or encountered prey qualities much higher than in Australia, for which we provide quantitative predictions.     

Gizzard and other lean mass components increase, yet Basal Metabolic Rates decrease, when red knots Calidris canutus are shifted from soft to hard-shelled food

We measured basal metabolic rate (BMR), body mass, lean mass, and gizzard mass of captive red knots Calidris canutus islandica maintained on a trout chow diet (soft-texture, low ash and water content) for several years and then shifted to small mussels Mytilus edulis (hard-texture, high ash and water content). During a 3-week period of feeding on mussels, body mass, lean mass, and gizzard mass increased 7.3 g (+7%), 10.5 g (+12%), and 4.9 g (+213%), respectively, yet BMR decreased from 0.96 to 0.89 W (−8%). Under the new mussel regime, red knots must have reduced the metabolic intensity of some of the tissues. This suggests that the experimental red knots experienced the transition to a mussel diet as stressful and energy limiting, resulting in an energy-saving strategy by reducing BMR in spite of hypertrophy of the gizzard and other organs.     

Handling Stress as a Measurement of Personality in Great Tit Nestlings (Parus major)

Interest in personality is growing in a wide range of disciplines, but only in a few systems it is possible to assess the survival value of personality. Field studies looking at the relationship between personality and survival value early in life are greatly hampered by the fact that personality can at present only be assessed after individuals become independent from their parents. In passerines, for example, this is often after a period of intensive selection for the survival on fledglings. The main aim of this study is therefore to develop a method to measure personality before this period of selection. For this purpose, we developed the handling stress (HS) test. We measured HS in 14-d-old great tit nestlings by counting the number of breast movements (breath rate) in four subsequent 15-s bouts for 1 min; before and after they were socially isolated from their siblings for 15 min. To calculate the repeatability of HS, we repeated the test 6 mo later. To assess the relationship between HS and exploratory behaviour, we correlated the outcome of both tests. We ran tests both on birds of lines selected for extreme personality and on wild birds from a natural population. We found that birds selected for fast exploration reacted more to HS compared with birds selected for slow exploration and that HS was repeatable in different life phases. We confirmed this by finding an increase in the HS with increasing exploratory scores in wild birds. These results show that we can use the HS test as a measurement of personality, making it a potential tool for studying the relationship between personality and survival value early in life.     

Effects of personality on territory defence in communication networks: a playback experiment with radio-tagged great tits

Individuals often differ consistently in behaviour across time and contexts, and such consistent behavioural differences are commonly described as personality. Personality can play a central role in social behaviour both in dyadic interactions and in social networks. We investigated whether explorative behaviour, as proxy of personality of territorial male great tits (Parus major), predicts their own and their neighbours'' territorial responses towards simulated intruders. Several weeks prior to playback, subjects were taken from the wild to test their exploratory behaviour in a standard context in the laboratory. Exploratory behaviour provides a proxy of personality along a slow–fast explorer continuum. Upon release, males were radio-tracked and subsequently exposed to interactive playback simulating a more or a less aggressive territorial intruder (by either overlapping or alternating broadcast songs with the subjects'' songs). At the same time, we radio-tracked a neighbour of the playback subject. Male vocal responses during playback and spatial movements after playback varied according to male explorative behaviour and playback treatment. Males with lower exploration scores approached the loudspeaker less, and sang more songs, shorter songs and songs with slower element rates than did males with higher exploration scores. Moreover, neighbour responses were related to the explorative behaviour of the subject receiving the playback but not to their own explorative behaviour. Our overall findings reveal for the first time how personality traits affect resource defence within a communication network providing new insights on the cause of variation in resource defence behaviour.     

EVOLUTION OF BASAL METABOLIC RATE AND ORGAN MASSES IN LABORATORY MICE

Animal species of similar body mass vary widely in basal metabolic rate (BMR). A central problem of evolutionary physiology concerns the anatomical/physiological origin and functional significance of that variation. It has been hypothesized that such interspecific differences in wild animals evolved adaptively from differences in relative sizes of metabolically active organs. In order to minimize confounding phenotypic effects and maximize relevant genetic variation, we tested for intraspecific correlations between body-mass-corrected BMR and masses of four organs (heart, kidney, liver, and small intestine) among six inbred strains of mice. We found significant differences between strains in BMR and in masses of all four organs. Strains with exceptionally high (or low) BMR tended to have disproportionately large (or small) organs. The mass of each organ was correlated with the masses of each of the other three organs. Variation in organ masses accounted for 52% of the observed variation in BMR, of which 42% represented between-strain variation, and 10% represented within-strain variation. This conclusion is supported by published measurements of metabolic rates of tissue slices from the four organs. The correlation between BMR and intestine or heart mass arose exclusively from differences between strains, while the correlation between BMR and liver or kidney mass also appeared in comparing individual mice within the same strain. Thus, even though the masses of the four examined organs account for no more than 17% of total body mass, their high metabolic activities or correlated factors account for much of the variation in BMR among mice. We suggest that large masses of metabolically active organs are subject to natural selection through evolutionary trade-offs. On the one hand, they make possible high-energy budgets (advantageous under some conditions), but on the other hand they are energetically expensive to maintain.     

Personality and urbanization: behavioural traits and <Emphasis Type="Italic">DRD4</Emphasis> SNP830 polymorphisms in great tits in Barcelona city

Most examples of adaptation to the urban environment relate to plasticity processes rather than to natural selection. Personality, however, defined as consistent individual differences in behaviour related to exploration, caution, and neophobia, is a good behavioural candidate character to study natural selection in relation to the urban habitat due to its heritable variation. The aim of this paper was to analyse variation in personality by comparing urban and forest great tits Parus major using standard tests of exploratory behaviour and boldness. We studied personality in 130 wild great tits captured in Barcelona city and nearby forests and found that urban birds were more explorative and bolder towards a novel object than forest birds. Genotype frequencies of the DRD4 SNP830 polymorphism, a gene region often associated with personality variation, varied significantly between forest and urban birds. Behavioural scores, however, were not correlated with this polymorphism in our population. Exploration scores correlated to boldness for forest birds but not for urban birds. Our findings suggest that the novel selection pressures of the urban environment favour the decoupling of behavioural traits that commonly form behavioural syndromes in the wild.     

Shorebirds' seasonal adjustments in thermogenic capacity are reflected by changes in body mass: how preprogrammed and instantaneous acclimation work together

Phenotypic flexibility in shorebirds has been studied mainly in the context of adjustments to migration and to quality of food; little is known on how birds adjust their phenotype to harsh winter conditions. We showed earlier that red knot (Calidris canutus islandica) can acclimate to cold by elevating body mass. This goes together with larger pectoral muscles, i.e., greater shivering machinery, and thus, better thermogenic capacity. Here, we present results of a yearlong experiment with indoor captive knots to determine whether this strategy is part of their natural seasonal phenotypic cycle. We maintained birds under three thermal regimes: constant cold (5 °C), constant thermoneutrality (25 °C) and natural seasonal variation between these extremes (9-22 °C). Each month we measured variables related to the birds' endurance to cold and physiological maintenance [body mass, thickness of pectoral muscles, summit metabolic rate (M(sum)), food intake, gizzard size, basal metabolic rate (BMR)]. Birds from all treatments expressed synchronized and comparable variation in body mass in spite of thermal treatments, with a 17-18% increase between the warmest and coldest months of the year; which appeared regulated by an endogenous driver. In addition, birds living in the cold exhibited a 10% higher average body mass than did those maintained at thermoneutrality. Thickness of the pectoral muscle tracked changes in body mass in all treatments and likely contributed to greater capacity for shivering in heavier birds. Consequently, M(sum) was 13% higher in cold-acclimated birds compared to those experiencing no thermoregulation costs. However, our data also suggest that part of maximal heat production comes from nonshivering processes. Birds facing cold conditions ate up to 25% more food than did birds under thermoneutral conditions, yet did not develop larger gizzards. Seasonal variation in BMR followed changes in body mass, probably reflecting changes in mass of metabolically active tissues. Just as cold-exposed birds, red knots in the variable treatment increased body mass in winter, thereby improving cold endurance. During summer, however, they maintained a lower body mass and thermogenic capacity compared to cold-exposed birds, similar to individuals kept at thermoneutrality. We conclude that red knots acclimate to seasonal variations in ambient temperature by modulating body mass, combining a preprogrammed increase in mass during winter with a capacity for fine-tuning body mass and thermogenic capacity to temperature variations.     

Seasonal variation in body composition in an Afrotropical passerine bird: increases in pectoral muscle mass are,unexpectedly, associated with lower thermogenic capacity

Phenotypic flexibility in avian metabolic rates and body composition have been well-studied in high-latitude species, which typically increase basal metabolic rate (BMR) and summit metabolism (M_sum) when acclimatized to winter conditions. Patterns of seasonal metabolic acclimatization are more variable in lower-latitude birds that experience milder winters, with fewer studies investigating adjustments in avian organ and muscle masses in the context of metabolic flexibility in these regions. We quantified seasonal variation (summer vs winter) in the masses of organs and muscles frequently associated with changes in BMR (gizzard, intestines and liver) and M_sum (heart and pectoral muscles), in white-browed sparrow-weavers (Plocepasser mahali). We also measured pectoral muscle thickness using a portable ultrasound system to determine whether we could non-lethally estimate muscle size. A concurrent study measured seasonal changes in BMR and M_sum in the same population of sparrow-weavers, but different individuals. There was no seasonal variation in the dry masses of the gizzard, intestines or liver of sparrow-weavers, and during the same period, BMR did not vary seasonally. We found significantly higher heart (~ 18% higher) and pectoral muscle (~ 9% higher) dry mass during winter, although ultrasound measurements did not detect seasonal changes in pectoral muscle size. Despite winter increases in pectoral muscle mass, M_sum was ~ 26% lower in winter compared to summer. To the best of our knowledge, this is the first study to report an increase in avian pectoral muscle mass but a concomitant decrease in thermogenic capacity.

     

Ambient temperature does not affect fuelling rate in absence of digestive constraints in long-distance migrant shorebird fuelling up in captivity

Pre-flight fuelling rates in free-living red knots Calidris canutus, a specialized long-distance migrating shorebird species, are positively correlated with latitude and negatively with temperature. The single published hypothesis to explain these relationships is the heat load hypothesis that states that in warm climates red knots may overheat during fuelling. To limit endogenous heat production (measurable as basal metabolic rate BMR), birds would minimize the growth of digestive organs at a time they need. This hypothesis makes the implicit assumption that BMR is mainly driven by digestive organ size variation during pre-flight fuelling. To test the validity of this assumption, we fed captive knots with trout pellet food, a diet previously shown to quickly lead to atrophied digestive organs, during a fuelling episode. Birds were exposed to two thermal treatments (6 and 24°C) previously shown to generate different fuelling rates in knots. We made two predictions. First, easily digested trout pellet food rather than hard-shelled prey removes the heat contribution of the gut and would therefore eliminate an ambient temperature effect on fuelling rate. Second, if digestive organs were the main contributors to variations in BMR but did not change in size during fuelling, we would expect no or little change in BMR in birds fed ad libitum with trout pellets. We show that cold-acclimated birds maintained higher body mass and food intake (8 and 51%) than warm-acclimated birds. Air temperature had no effect on fuelling rate, timing of fuelling, timing of peak body mass or BMR. During fuelling, average body mass increased by 32% while average BMR increased by 15% at peak of mass and 26% by the end of the experiment. Our results show that the small digestive organs characteristic of a trout pellet diet did not prevent BMR from increasing during premigratory fuelling. Our results are not consistent with the heat load hypothesis as currently formulated.     

Seasonal acclimatization of metabolism in Eurasian tree sparrows (Passer montanus)

Acclimatization to winter conditions is an essential prerequisite for survival of small passerines of the northern temperate zone. Changes in photoperiod, ambient temperature and food availability trigger seasonal acclimatization in physiology and behavior of many birds. In the present study, seasonal adjustments in several physiological, hormonal, and biochemical markers were examined in wild-captured Eurasian tree sparrows (Passer montanus) from the Heilongjiang Province in China. In winter sparrows had higher body mass and basal metabolic rate (BMR). Consistently, the dry mass of liver, heart, gizzard, small intestine, large intestine and total digestive tract were higher in winter than in that in summer. The contents of mitochondrial protein in liver, and state-4 respiration and cytochrome c oxidase (COX) activity in liver and muscle increased significantly in winter. Circulating level of serum triiodothyronine (T₃) was significantly higher in winter than in summer. Together, these data suggest that tree sparrows mainly coped with cold by enhancing thermogenic capacities through increased organ masses and heightened activity of respiratory enzymes activities. The results support the view that prominent winter increases in BMR are manifestations of winter acclimatization in tree sparrows and that seasonal variation in metabolism in sparrows is similar to that in other small temperate-wintering birds.     

Additive and nonadditive genetic variation in avian personality traits

Individuals of all vertebrate species differ consistently in their reactions to mildly stressful challenges. These typical reactions, described as personalities or coping strategies, have a clear genetic basis, but the structure of their inheritance in natural populations is almost unknown. We carried out a quantitative genetic analysis of two personality traits (exploration and boldness) and the combination of these two traits (early exploratory behaviour). This study was carried out on the lines resulting from a two-directional artificial selection experiment on early exploratory behaviour (EEB) of great tits (Parus major) originating from a wild population. In analyses using the original lines, reciprocal F(1) and reciprocal first backcross generations, additive, dominance, maternal effects ands sex-dependent expression of exploration, boldness and EEB were estimated. Both additive and dominant genetic effects were important determinants of phenotypic variation in exploratory behaviour and boldness. However, no sex-dependent expression was observed in either of these personality traits. These results are discussed with respect to the maintenance of genetic variation in personality traits, and the expected genetic structure of other behavioural and life history traits in general.     

Environmental transmission of a personality trait: foster parent exploration behaviour predicts offspring exploration behaviour in zebra finches

Consistent behavioural differences among individuals are common in many species and can have important effects on offspring fitness. To understand such ‘personality’ variation, it is important to determine the mode of inheritance, but this has been quantified for only a few species. Here, we report results from a breeding experiment in captive zebra finches, Taeniopygia guttata, in which we cross-fostered offspring to disentangle the importance of genetic and non-genetic transmission of behaviour. Genetic and foster-parents’ exploratory type was measured in a novel environment pre-breeding and offspring exploratory type was assessed at adulthood. Offspring exploratory type was predicted by the exploratory behaviour of the foster but not the genetic parents, whereas offspring size was predicted by genetic but not foster-parents’ size. Other aspects of the social environment, such as rearing regime (uni- versus biparental), hatching position, brood size or an individual''s sex did not influence offspring exploration. Our results therefore indicate that non-genetic transmission of behaviour can play an important role in shaping animal personality variation.     

Intraspecific variation in reproductive physiology and egg quality in the European Starling Sturnus vulgaris

Egg mass shows large intraspecific variation in birds and is repeatable within individuals. The mechanisms underlying this variation are unknown. We hypothesized that measures of egg quality (the mass of yolk protein, yolk lipid, and albumen protein) would be positively correlated with the plasma pools of the yolk precursor vitellogenin, and the masses of the oviduct, metabolic machinery (liver, heart, lungs, kidneys, gizzard, small intestine and pancreas), and endogenous stores of protein and lipid. We tested these predictions in European Starlings Sturnus vulgaris collected at the peak of egg production effort. In contrast to our predictions, both yolk protein and yolk lipid were negatively correlated with plasma vitellogenin levels. Albumen protein was positively related to oviduct mass, but other aspects of body composition failed to explain variation in egg quality. Hence, while we observed correlations between egg composition and peripheral systems (circulating precursor pools and the oviduct), we found no evidence that egg quality is determined by more general processes, i.e., the supply and processing of nutrients.     

Fibre-induced feed sorting in King Quail (Coturnix chinensis): behavioural plasticity elicited by a physiological challenge

We examined the effect of an abrupt change in diet fibre content on the feed intake, gastrointestinal morphology and utilisation of gastroliths by a small (ca. 40 g body mass) herbivorous bird, the King Quail (Coturnix chinensis). King Quail were acclimated for 14 days on a low-fibre (LF) pullet starter diet. Following acclimation, half the quail population was immediately switched to a 23 % wood-shaving diluted high-fibre (HF) diet for a further 14 days. Contrary to expectations, we found no differences in feed intake, gut morphology or gastrolith mass between the LF- and HF-fed quail. However, when switched from the LF to HF diet, the quail commenced feed-sorting behaviours that permitted HF-fed animals to maintain body condition (mass, abdominal fat mass) without adjustments to intestinal organ sizes or gastrolith mass. Feed sorting was initiated only after exposure to the HF diet, which corresponded with an immediate reduction in food intake, suggesting that the sorting behaviour was cued by a physiological challenge associated with the HF diet. This challenge apparently induced preferential sorting behaviour and was possibly due to abrupt changes in the rate of food passage, impacting satiation or other internal cues.     

Rapid changes in the size of different functional organ and muscle groups during refueling in a long-distance migrating shorebird.

The adaptive value of size changes in different organ and muscle groups was studied in red knots (Calidris canutus islandica) in relation to their migration. Birds were sampled on five occasions: at arrival in Iceland in May 1994, two times during subsequent refueling, at departure toward, and on return from, the high arctic breeding grounds. During their 24-d stopover in May, body mass increased from 144.3 to 214.5 g. Mass gains were lowest over the first week (0.85 g/d, only fat-free tissue deposited). Over the subsequent 10 d, average mass increased by 5.0 g/d (fat contributing 78%), and over the last week before takeoff, it increased by 2.0 g/d (fat contributing over 100% because of loss of lean components). There were no sex differences in body and fat mass gains. Over the first interval, lean masses of heart, stomach, and liver increased. During the middle 10 d, sizes of leg muscle, intestine, liver, and kidneys increased. Stomach mass decreased over the same interval. In the last interval before takeoff, the stomach atrophied further and the intestine, leg muscles, and liver became smaller too, but pectoral muscles and heart increased in size. Sizes of "exercise organs" such as pectoral muscle and heart were best correlated with body mass, whereas sizes of organs used during foraging (leg muscles) and nutrient extraction (intestine, liver) were best correlated with rate of mass gain. Kidneys changed little before takeoff, which suggests that they are needed as much during flight as during refueling.     

Individual‐ and population‐level personalities in a floriphilic katydid

Behaviour, including personality, informs us about the response of animals towards their changing environment. Despite the widespread occurrence of florivorous insects and the important but often underrated ecological roles that they play, the study of florivore behaviour is neglected relative to that of pollinators and other herbivores. Specifically, we do not know how different personality types can develop among florivores and enable them to persist in habitats with an ephemeral and dynamic availability of food resources. To address this knowledge gap, we investigated the following questions: whether the (a) inter‐individual differences of exploration and boldness are consistent; (b) inter‐population differences of exploration and boldness are consistent; (c) exploration and boldness are correlated. We collected individuals of the polyphagous floriphilic katydid, Phaneroptera brevis from four populations from wasteland sites in Singapore and performed a personality assay conducted in an insectary to investigate the exploratory and boldness levels of the individuals and populations. The major novel finding was that the floriphilic P. brevis katydids exhibit population‐level personality types for boldness, but not for exploration. Some katydid individuals were consistently more exploratory and bolder than other individuals. However, contrary to our predictions, we did not find any evidence of behavioural syndromes in the katydid individuals, as the boldness level for individuals was not significantly correlated with exploration for individuals. This suggests that an individual which is more exploratory may not be equally keen to take risks and consume novel food that it encounters. Our findings also suggest that boldness and exploration are linked to ecologically important behaviours, but more studies are needed to better understand population‐level personality and how and why natural selection may favour the evolution of personality in certain populations.