Body temperature and territory selection by males of the speckled wood butterfly (Pararge aegeria): what makes a forest sunlit patch a rendezvous site?

1. Insects locate mobile resources like prey items or mates using either sit‐and‐wait (‘perching’) or active (‘patrolling’) searching strategies. The sit‐and‐wait strategy can be accompanied by defending and monopolising a site through territorial behaviour. 2. The present study focuses on the territorial perching behaviour in males of the speckled wood butterfly (Pararge aegeria L.). Recent studies suggested that the selection of territories (i.e. sunlit patches on the forest floor) is driven by structural characteristics of the site that affect male visual detection. However, given that adult butterflies are heliothermic organisms and that forests provide a diverse array of light environments, it seems likely that thermal aspects may also be used for territory selection. 3. We tested whether used and unused sunlit patches differed in thermal profile under field conditions in a Belgian woodland. We also used dummy butterflies to quantify variation in operative thoracic temperature and to calculate heating rates within (i.e. different vegetation structures) and between patches. 4. Sunlit patches occupied by a territorial male were larger, and were more frequently characterised by low vegetation structures compared with empty sunlit patches. It took longer to reach optimal thorax temperature (starting from a fixed suboptimal body temperature) in small patches compared with large patches. 5. We suggest that aspects of visual detection need to be combined with thermal aspects to fully understand territory selection in the speckled wood butterfly, as synergetic and/or trade‐off effects of ambient temperature, solar radiation, and canopy/vegetation structure may be involved.     

Effect of larval food stress on male adult behaviour, morphology and reproductive investment in the butterfly Pararge aegeria

Males of several animals increase their reproductive success by territorial behaviour. In butterflies, males may defend a territory (i.e., territorial perching tactic), but this is assumed to be an energetically costly way to locate mates. Limitations of the energy budget may affect fight performance, and may, consequently, force males to adopt an alternative non-territorial searching behaviour (i.e., patrolling tactic) to maximize reproductive success. In this study, we tested to what extent behavioural tactics adopted by adult males of the butterfly Pararge aegeria (L.) were affected by the nutritional conditions during the larval stage. We compared the occurrence of territorial versus patrolling behaviour, lipid mass, flight muscle ratio, metabolic rate and spermatophore production of low quality males that were reared as a larva on drought-stressed host plants and control, high quality males. Low quality males were less likely to adopt the territorial perching tactic and emerged as adults with lower lipid mass than high quality males, but they were able to restore their lipid mass through adult feeding (and perhaps the breakdown of flight muscles). Host plant quality also affected spermatophore size. Independent of the larval food treatment, territorial perching males metabolised more lipids than non-territorial males, produced larger spermatophores and copulated for longer than males adopting non-territorial behaviour. We discuss the results relative to the co-existence of the behavioural tactics (perching and patrolling).     

Hostplant-habitat structure and the evolution of butterfly mate-locating behaviour

A hostplant-habitat model is described which explains fundamental differences in mate-locating behaviour between butterfly species, based on a sequence of spatially distinctive activities (habitual patrolling, localized perching, territoriality, lek assembly). It gives equal weight to phylogeny and ecology and is consistent with models for the evolution of territoriality and population structure. Perching repeatedly at characteristic sites, territoriality and lek assembly evolve when resources become associated with distinctive hostplant-habitat structures in which topographic features provide predictable vantage points for acquiring mates. Habitual patrolling is an ancestral behaviour which is suited to widespread and unpredictable resource distributions and to species whose individual apparency detracts from the use of topographic vantage points. Hostplants (emergence sites), nectar and thermoregulation sites typically provide resources and act as precursors to localized perching, territoriality and lek assemblages. But these behaviours fail to evolve in species whose habitats do not include obvious topographic landmarks which provide clear vantage points for males awaiting females and focal points for females searching for males. Where these landmarks occur they result in an increase in receptive female density at those locations compared to the remainder of the habitat.     

Evidence for hilltopping in bumblebees?

1. Male bumblebees are known to exhibit a range of mate‐location behaviours, including perching on prominent objects and darting at passing queens, patrolling of scent‐marked flight routes, and waiting outside nest entrances for virgin queens to emerge. Here we provide evidence for a fourth strategy, known as hilltopping. This behaviour is widely known from a range of invertebrates, but has not previously been described in bumblebees. 2. We studied the distribution of bumblebees along transects ascending four hills in Scotland and demonstrate that, relative to workers, males of four bumblebee species or species groups (Bombus lapidarius, B. monticola, B. pascuorum, and B. lucorum/magnus/cryptarum/terrestris) tend to congregate at or near the tops of hills. This is, to our knowledge, the first evidence for hilltopping in bumblebees and the first record of any putative mate‐locating behaviour for male B. pascuorum, a very common species in Europe. 3. We note that, in common with most previous studies of mate‐locating behaviour in bumblebees, attraction of virgin queens and mating were not observed.     

Diurnal and seasonal changes in the mate-locating behavior of the satyrine butterfly <Emphasis Type="Italic">Lethe diana</Emphasis>

The time schedule of mate-locating behavior in the male satyrine butterfly Lethe diana was studied. Males displayed territorial behavior only in the afternoon, but it was not known whether they attempted to locate mates in the morning. In order for males of most butterfly species to mate with a virgin female, it would be most effective to initiate mate-locating behavior in the morning, when synchronous eclosions occur. Males of L.diana flew patrols in the morning and displayed territorial behavior in the afternoon. The cost of territorial behavior is probably less than that of patrolling, as territorial flights are shorter than those when patrolling. The temporal switching of mate-locating behavior is theoretically favored, since the reproductive value of females decreases later in the day, when males perform the less costly mate-locating behavior. Thus, the diurnal switching of male mate-locating behavior may be an adaptive strategy in response to the rapid decline in the reproductive value of females over the course of a day.     

Investigating the Consistency of Mate-Locating Behavior in the Territorial Butterfly Hypolimnas bolina (Lepidoptera: Nymphalidae)

The study of butterfly behavior has afforded valuable insights into the evolution of alternative mating tactics. Two hypotheses derived from this area of research contend that (1) territoriality is only viable under low to moderate conspecific densities (due to the costs of site defence) and (2) perching may be employed only when thermal conditions constrain flight activity. These hypotheses were evaluated by investigating mate locating behavior in Hypolimnas bolina, a territorial species that is naturally subject to variation in population density and weather conditions. Male behavior was charted throughout the day during a period of high population density at an encounter site in tropical Australia. Perching was the primary tactic, although a small proportion of individuals patrolled nonaggressively in the afternoon. Population-level male behavior failed to support predictions drawn from either the territory economics or thermal constraint hypotheses. First, the proportion of perching males and the number of aggressive conspecific interactions (per male) increased with increasing male density at the site. Second, few males patrolled at the hottest, brightest time of day (approximately midday), and the diel distribution of perchers did not emulate the U-shaped distribution shown by the occurrence of dorsal basking behavior. These results show that perching in this species is not a suboptimal tactic employed when temperatures constrain flight activity but may represent the best method of locating receptive females. At this stage the reproductive significance of the observed patrolling behavior remains obscure.     

Ecological and phylogenetic perspectives on wing stiffness in nine Theclini species (Lepidoptera: Lycaenidae)

We studied the relationship between wing stiffness and butterfly ecology and phylogeny. Nine species belonging to the tribe Theclini of the family Lycaenidae were selected and examined for the wing stiffness of dried specimens by a three‐point bending test. It was found in Japonica lutea that the wing stiffness was not affected by the humidity to which it had been exposed, but was strongly affected by wing size and sex. Comparisons of sexual differences in four species indicated that females of patrolling species had stiffer wings than conspecific males, but that males of territorial species had stiffer wings than conspecific females. Finally, the wing stiffness was compared among males of nine species that use different mate‐locating tactics, and the results revealed a tendency that males of territorial species have stiffer wings than males of patrolling species. These results, though including a few exceptional cases, are discussed from the perspective of ecological requirements and phylogenetic constraints on the species.     

Cross-fostering Does Not Influence the Mate Preferences and Territorial Behaviour of Males in Lake Victoria Cichlids

The ability to recognise conspecifics in contexts of mate choice and territorial defence may have large effects on an individual's fitness. Understanding the development of assortative behaviour may shed light on how species assortative behaviour evolves and how it may influence reproductive isolation. This is the case not only for female mate preferences, but also for male mate preferences and male territorial behaviour. Here we test with a cross-fostering experiment whether early learning influences male mate preferences and male–male aggression biases in two closely related, sympatrically occurring cichlid species Pundamilia pundamilia and Pundamilia nyererei from Lake Victoria. Males that had been fostered, either by a conspecific female or by a heterospecific female, were tested for their aggression bias, as well as for their mate preferences, in two-way choice tests. Males cross-fostered with conspecific and heterospecific foster mothers selectively directed their aggression towards conspecific intruders. The cross-fostering treatment also did not affect male mate preferences. These results are in striking contrast with the finding that females of these species show a sexual preference for males of the foster species.     

Factors influencing mate guarding and territory defence in the stitchbird (hihi)

Socially monogamous male birds are predicted to maximise their reproductive success by pursuing extra-pair copulations (EPCs) while engaging in anti-cuckoldry behaviour such as mate guarding. In the stitchbird, Notiomystis cincta, high levels of forced EPCs and a high proportion of nestlings resulting from extra- pair fertilisations lead to the prediction that males of this species should exhibit intense paternity guarding behaviours. While studying an isolated stitchbird population on Tiritiri Matangi Island New Zealand (3636'S, 17453'E), I collected daily behavioural data throughout the breeding season from 15 males in 2000/01 and 27 males in 2001/02. In this study, male stitchbirds demonstrated clear paternity guarding by exhibiting: (1) an increased likelihood of being close to their mate during her fertile period, (2) an increased initiation of mate contact during her fertile period, (3) switching from site-specific territorial defence during the pre-fertile period to defending an area centring on the their female partners location during her fertile period, and (4) an increased following of the female to communal feeding sites outside the territory during her fertile period. For polygynous males, mate guarding and territorial defence were conditional on which of their females was fertile. Additional evidence supporting the hypothesis that mate guarding in this species is a form of paternity assurance, rather than protection from harassment, is that males protected their partner from harassment by other stitchbird males but did not intervene when females were harassed by male bellbirds, Anthornis melanura. While mate-guarding intensity in many species is conditional on the stage of female fertility, male stitchbirds also modified their behaviour depending on the location of the female and the rate of intrusions by extra-pair males. Resident males adopted a best-of-a-bad-job tactic when they were unable to locate their female by defending an area around her last known location. Furthermore, when the rate of intrusions by extra-pair males increased they traded-off the area they could defend within their territory against their ability to guard the female. Territory takeovers were uncommon, but when they did occur older males displaced younger males and healthy birds displaced sick ones. Contrary to the prevailing view that mate guarding is a male response to female infidelity, male stitchbirds appear to use mate guarding primarily to prevent paternity losses from forced EPCs. Future assessments of mate guarding function should consider the possibility that mate guarding involves a combination of conflict and co-operation between the sexes.     

SEXUAL SELECTION AND BUTTERFLY DESIGN—A COMPARATIVE STUDY

Temperate butterflies of 44 species were examined to determine if their mating system (perching and patrolling) affected flight design. To control for spurious effects due to ancestry, 25 of these species were assigned to eight contrasts within which a change in mating system had occurred. In perching species sexual selection was predicted to favor traits associated with high acceleration ability and speed, while in patrolling species traits associated with flight endurance were predicted. In conformance with these expectations males of perching species had larger thorax/body mass ratios, higher wing loadings, and higher aspect ratios than patrolling species. The male mating system affected females in the same direction in the same variables as males. This could be explained by a genetic correlation with males. When removing the covariance between the sexes, only male design was explained by the mating system. The mating system was also associated with different degrees of sexual dimorphism in wing size. This supported the hypothesis that male design was affected by the mating system.     

Do Feeding Resources Induce the Adoption of Resource Defence Polygyny in a Lekking Butterfly?

Among polygynous species, males often compete for the possession of mating sites to increase their reproductive success. Weaker individuals frequently adopt alternative non‐territorial mate‐locating tactics, but the adoption of alternative territorial tactics may also occur. Although alternative tactics with territory defence are less common in arthropods, factors that drive its adoption may provide information to understand the organization of different territorial mating systems in the group. Here we investigate the adoption of resource‐based territoriality as an alternative to a non‐resource‐based one by males of the butterfly Paryphthimoides phronius. Male P. phronius commonly defend sunny clearings lacking feeding resources in the forest edge (non‐resource‐based territoriality). However, after experimentally offering fermenting fruit in previously undefended sites, we showed that males also adopt a resource defence tactic. Males in territories with fermenting fruits apparently feed on this resource when they are not defending the territory. However, males in sites without resources did not migrate to territories with resources when given the opportunity. To our knowledge, this is the first experimental investigation to show a non‐resource‐ and an alternative resource‐based mate‐locating tactic in a butterfly. We suggest that this behavioural flexibility may represent an important step to understand the ecological factors responsible for the organization and evolution of different territorial mating systems in insects.     

Importance of historical factors on species richness and composition of butterfly assemblages (Lepidoptera: Rhopalocera) in a northern Iberian mountain range

ABSTRACT. The butterfly assemblages of three vegetation types (pasture, heathland and rocky outcrop with Genista spp.) and two geographical orientations (a northern slope and a sheltered gorge) were studied along an altitudinal gradient in the Picos de Europa in northern Spain. In order to study the effects or historical climatic changes on current butterfly assemblages, it was examined if changes in richness and faunal composition of assemblages were joined with changes in their biogeographical composition. The three vegetation types showed characteristic butterfly assemblages, but no difference in species richness and biogeographical composition was found. Species richness in the sheltered gorge was higher than in the northern slope. Faunal composition also varied between orientations and there were more widespread and Mediterranean species in the sheltered gorge than in the northern slope. Species richness declined with increasing altitude, though a midaltitudinal peak was observed. There was a faunal segregation between high and low localities. Species richness of widespread and Mediterranean butterflies decreased as altitude increased, whereas the reverse was true for montane species. Moreover, biogeographical elements differed in their climatic tolerances measured as altitudinal ranges. Therefore, changes in richness and composition of butterfly assemblages between both slopes and along the altitudinal gradient were joined in part with variation in their biogeographical composition. These results suggest that current species richness and composition of butterfly assemblages in the Picos de Europa might be the consequence of differential colonization of refuges during the past climatic changes.     

Why Do Males of Parnassius imperator Fight for Bare Rocks but Not the Nectar Flower During Mate Selection?

Territoriality is an important male behavioral trait used in mate selection of butterflies, but the factors determining the outcome of territorial contests remain controversial. We studied the foraging and territorial behaviors of males in the Parnassius imperator butterfly. Adult males were selective about nectar sources, but they did not defend nectar flowers. By contrast, they fought over areas of bare rocks because bare rocks were used as lek locations to attract and court receptive females. The ownership of a lek, rather than sex pheromones or wing pigments, played the key role in attracting females. In an owner replacement experiment, the owner of a territory always defeated intruders, indicating that perching order, not relative body size, determined the outcome of territorial contests. Territories are small areas of bare rocks that lack food resources, so intruders do not challenge the ownership of a resident in a location where they can easily find vacant territories. Males that first perch in a territory have an advantage compared to intruders in that they can forage on flowers in the vicinity of the territory. This may help explain why satiated owners always defeated fatigued intruders. Our findings suggest that early perching on bare rock allows male butterflies of P. imperator to occupy a territory, attract receptive females and exclude later‐arriving competitors.     

Female choice of sites versus mates in a coral reef fish,Thalassoma bifasciatum

To understand the dynamics of sexual selection, one needs to identify the basis of female choice in mating. Daily censuses showed that most spawns of individual female bluehead wrasses, Thalassoma bifasciatum, occurred at one of several available mating sites, either in a group-spawning aggregation of small males or with a large territorial male. Most females were not located near their principal mating site during the non-spawning period of the day. Females were thus exposed to other males and sites and should have had the opportunity for choice. When a resident territorial male disappeared or was removed experimentally from a mating site, females did not alter their fidelity to that site. Furthermore, manipulations showed that when a territorial male changed the location of his mating activity to an adjacent site, females did not follow him but continued to mate at their customary site. Thus mating sites, rather than the males occupying them, appeared to be the objects of female choice. The bright coloration and courtship behaviour of larger males may demonstrate the safety of a particular site rather than indicating some aspect of male quality.     

Landmark Territoriality in the Neotropical Paper Wasps Polistes canadensis (L.) and P. carnifex (F.) (Hymenoptera: Vespidae)

Male Polistes canadensis and P. carnifex aggregate along crests of prominent ridges in Santa Rosa National Park, Costa Rica. At these sites males of both species defend territories (trees and shrubs) by chasing conspecific rivals. Territories do not contain nests or resources that are collected by females. Chasing by territorial males reduces the amount of time spent by intruders in a territory. I describe and contrast male territorial behavior of both species. Some male P. canadensis are territorial while others in the same area exhibit patrolling behavior, flying from one occupied territory to another. Males of P. carnifex exhibit territoriality only. Patrolling in P. canadensis is an outcome of relatively high male density along the ridge, rendering territories in short supply, as shown by the observation that experimentally vacated territories are seized rapidly by formerly patrolling males. Due to a high intraspecific intrusion rate, territorial male P. canadensis spend less time perching and more time flying and chasing intruders from their territories than do male P. carnifex. Males of these two species also differ in the placement of their territories along the ridgeline; P. canadensis occupy territories in saddles while P. carnifex occupy those at peaktops. I show that this divergent spatial pattern is not maintained by competitive exclusion of either species by the other, and I discuss alternative explanations for their separate spatial distributions. Comparative data suggest that males are territorial because females restrict matings to within territories, and I discuss alternative hypotheses to explain this bias in female behavior.     

Thermoregulatory differences between phenotypes in the speckled wood butterfly: hot perchers and cold patrollers?

Males of the speckled wood butterfly Pararge aegeria L. (Satyrinae), actively search for females (“patrolling”) or wait for them at particular places (“perching”). Darker males are more likely to patrol than pale ones, which are mainly territorial perchers. We studied whether this morphological variation relates to thermoregulatory differences. The relationship between thoracic temperature and ambient temperature differed between the colour types under natural conditions: darker males had on average lower body temperatures than paler males. Different activities (e.g. resting, flying) and behavioural strategies (perching or patrolling) were associated with differences in thoracic temperature: patrolling males which mainly engaged in long flights and periods of basking afterwards, had lower thoracic temperatures than perching males which engaged in very short flights, fights and basking. When resting for a while thoracic temperatures did not differ between males practising different strategies. Under laboratory conditions, darker males heated up faster than pale males but there was no difference in the thoracic temperature at which they started to fly. These results indicate that thermal requirements (or general conditions) differ between the behavioural strategies, and that behavioural differences between phenotypes (colour types) relate to differences in thermal ecology. This supports the idea that darker males are better adapted to patrolling. There is no evidence that one mate-locating strategy is always superior to the other, which coincides with the observation that both strategies co-exist. More generally, this study shows that relatively small differences in colour can have a considerable effect on thermoregulation and hence on the behavioural strategies a heliothermic insect will adopt. Received: 15 August 1997 / Accepted: 15 December 1997     

Evidence for Mate Guarding Behavior in the Taylor’s Checkerspot Butterfly

Discerning the intricacies of mating systems in butterflies can be difficult, particularly when multiple mating strategies are employed and are cryptic and not exclusive. We observed the behavior and habitat use of 113 male Taylor’s checkerspot butterflies (Euphydryas editha taylori). We confirmed that two distinct mating strategies were exhibited; patrolling and perching. These strategies varied temporally in relation to the protandrous mating system employed. Among perching males, we recorded high site fidelity and aggressive defense of small (<5 m²) territories. This territoriality was not clearly a function of classic or non-classic resource defense (i.e., host plants or landscape), but rather appeared to constitute guarding of female pupae (virgin females). This discrete behavior is previously undocumented for this species and has rarely been observed in butterflies.     

Reproductive success in male sunbleak, a recent invasive fish species in the U.K.

The spawning behaviour of male nest guarding sunbleak Leucaspius delineatus , a recent invasive species in southern England, was studied and quantified for the first time. In the absence of physical differences between territorial and non‐territorial males ( i.e . colour, size, etc .), the reproductive behaviour of territorial males was analysed and related to reproductive success. The results showed that females preferred high‐courting and highly aggressive males. The initial cue in female mate choice, however, was based on courtship, while aggression was the decisive behavioural trait in influencing mate choice, providing a direct signal of physical condition and 'paternal competence'. Some males picked nest sites which were subsequently preferred by other males taking over the nest of a previous male ('communal nest'), with the new territorial male adopting the eggs already present at the nest. It appears that either due to female preference for nests already containing eggs or lower rates of sired egg predation by dilution among unrelated eggs, sunbleak males have adopted the mating strategy of allopaternal care.     

Mate-locating behaviour, habitat-use, and flight morphology relative to rainforest disturbance in an Afrotropical butterfly

To cope with environmental constraints, organisms can show variation in phenotype, either by genetic adaptation or phenotypic plasticity. These patterns are especially pronounced in ecosystems that are under anthropogenic influence. Due to human-induced disturbances such as logging and deforestation, tropical forests comprise such a system. To date, most studies have dealt with ecological responses at the community level relative to forest disturbance or degradation. However, the evolutionary consequences of tropical forest deterioration on behaviour and functional morphology have received far less attention compared to temporal regions. From a resource-point of view, light conditions are essential for heliotherms such as butterflies. Because degradation of tropical cloud forests in the Taita Hills (Kenya) is very pronounced, the present study tested whether this induced changes in mate-location strategies, habitat-use, and functional flight morphology in a forest butterfly, Salamis parhassus. According to predictions from temperate regions, it was hypothesized that the species would change its mate location strategy from perching to patrolling in more disturbed forests, that this higher mobility results in a faster occupancy of light gaps, and that it accords with a higher wing loading within populations from undisturbed forests. These hypotheses were confirmed by field surveys and experiments. The present study demonstrates that degradation of tropical forests does not only affect communities (e.g. species richness), but also the behaviour and functional morphology of individual species.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 830–839.     

Functions of duetting in the tropical boubou, Laniarius aethiopicus: territorial defence and mutual mate guarding

Duetting, found in many animal taxa, is still a poorly understood form of signalling behaviour despite numerous hypotheses to explain its function. One contentious issue is whether duetting is a cooperative endeavour or signals conflict between the sexes. We studied the function of duetting in the tropical boubou, using interactive playback experiments. We staged encounters by presenting paired birds with four variants of each of four duet types (neighbour and stranger solos and neighbour and stranger duets). Multiple lines of evidence suggest that both joint territorial defence and mutual mate guarding are important functions of duetting, and that both cooperation and conflict between males and females have shaped duetting behaviour. Of a repertoire of 12 duet types, eight were used for joint territorial defence. Two of these eight duet types were probably also used for mutual mate guarding, suggesting that individual duets can have multiple functions. We found an unusual response for duetting birds in that females often synchronized notes with male solo playback to form precise duets. In turn, males attempted to jam the song of rivals when females joined the song of male solo playback, providing further evidence for the importance of acoustic mate guarding. Song jamming by males has not previously been described for duetting species. Finally, we suggest that the repertoire of duet types is used to fine-tune territorial encounters in a network environment of multiple interactants.